High survival in poor years: life history tactics adapted to mast seeding in the edible dormouse

Edible dormice (Glis glis) reproduce in years with beech mast seeding, but entire populations may skip reproduction in years when tree seeds, a major food resource of this small hibernator, are absent. We tested the hypothesis that the year-to-year variability in reproductive effort caused by this breeding strategy should lead to detectable differences in yearly survival rates. Therefore, we analyzed capture-recapture data from animals occupying nest boxes, collected over nine years at two study sites in Germany. Among fully grown adults (aged two years or older), survival probabilities were significantly lower (0.32 +/- 0.04) after reproductive years (n = 5) compared to years (n = 4) with absent or below-average reproduction (0.58 +/- 0.07) on both study sites. This trade-off between reproduction and subsequent survival was observed in both females and males and appears to be a relatively rare case in which costs of reproduction in terms of longevity are detectable at the population level. Effects of reproduction on survival were less pronounced when yearlings (with a generally lower reproductive effort) were included and were more distinct in a suboptimal habitat. Of those females breeding in nest boxes, 96.5% had only one or two litters within the study period. Considering these and previously published results, including a report of extremely high mean longevities (9-12 years) of dormice in a habitat with infrequent mast seeding, we conclude that edible dormice flexibly adjust life history tactics to local mast patterns. Long stretches of mast failures can in fact lead to relative semelparity, i.e., a strategy in which dormice "sit tight" for several years until environmental conditions are favorable for reproduction.     

Catastrophic Reproductive Failure, Terrestrial Survival, and Persistence of the Marbled Salamander

Abstract:  Wide variation in reproductive success is common among amphibians that breed in seasonal ponds, but persistence of adults can buffer against these fluctuations, particularly for long-lived species. We hypothesized that the frequent episodes of catastrophic failure of the marbled salamander ( Ambystoma opacum ) enhance the importance of high terrestrial survival. At Rainbow Bay in South Carolina reproductive success was poor (<1 metamorph/breeding female) in nearly half of the 22 years that the species bred. Complete failure occurred in 6 of 22 years. To study catastrophic failure, we adapted an age-structured, individual-based model with density-dependent growth and survival of larvae. The model was based on extensive data from local field studies and experiments. With consistently good survival in the pond stages, the simulated population required survival probabilities in the upland stages (juveniles and adults) near 0.5/year to persist and near 0.8/year to achieve the increases observed. Catastrophic failure, occurring randomly with probability 0.5/year, created additional fluctuations in the population, raised the thresholds of survival required for persistence, and caused extinction under conditions that were otherwise favorable. The marbled salamander at Rainbow Bay is not at great risk of extinction because of catastrophic failure, but the risk increases dramatically if life span is decreased or frequency of failure is increased. Any reduction in terrestrial survival will have deleterious consequences by reducing the breeding populations at equilibrium, even if it does not jeopardize persistence. Our model provides assessments of risk that can be applied to poorly studied species with similar life histories, such as the endangered flatwoods salamander ( A. cingulatum ).     

Sex differences in survival costs of reproduction in a promiscuous primate

In sexually promiscuous mammals, female reproductive effort is mainly expressed through gestation, lactation, and maternal care, whereas male reproductive effort is mainly manifested as mating effort. In this study, we investigated whether reproduction has significant survival costs for a seasonally breeding, sexually promiscuous species, the rhesus macaque, and whether these costs occur at different times of the year for females and males, namely in the birth and the mating season, respectively. The study was conducted with the rhesus macaque population on Cayo Santiago, Puerto Rico. Data on 7,402 births and 922 deaths over a 45-year period were analyzed. Births were concentrated between November and April, while conceptions occurred between May and October. As predicted, female mortality probability peaked in the birth season whereas male mortality probability peaked in the mating season. Furthermore, as the onset of the birth season gradually shifted over the years in relation to climatic changes, there was a concomitant shift in the seasonal peaks of male and female mortality. Taken together, our findings provide the first evidence of sex differences in the survival costs of reproduction in nonhuman primates and suggest that reproduction has significant fitness costs even in environments with abundant food and absence of predation.     

Life history costs and consequences of rapid reproductive maturation in female rhesus macaques

Summary Life history theory suggests that reproduction at one point in time involves costs in terms of energy, reduced survival, or probability of reproduction at a future point in time. ln long-lived iteroparous organisms, initiating reproduction at a relatively young age may exact a cost in terms of reduced survivorship, but an early age of first reproduction could be beneficial if it lengthens the breeding lifespan. Data collected over 30 years from one population of rhesus macaques, Macaca mulatta, were analyzed to determine the fertility and survivorship costs of initiating reproduction at a relatively young age. Low population density and high social status increased the chances of accelerating age at first parturition, but high dominance rank was not associated with greater lifetime reproductive success. Rapid reproductive maturation neither reduced short-term survivorship nor decreased lifespan. Fertility costs arose if young females reared a male, but not female, offspring. The fitness consequences of rapid reproductive maturation depend upon longevity, with age at death having a significantly greater impact on lifetime reproductive success than age at first parturition.Correspondence to: F.B. Bercovitch     

Spatial Patterns of Breeding Success of Grizzly Bears Derived from Hierarchical Multistate Models

Conservation programs often manage populations indirectly through the landscapes in which they live. Empirically, linking reproductive success with landscape structure and anthropogenic change is a first step in understanding and managing the spatial mechanisms that affect reproduction, but this link is not sufficiently informed by data. Hierarchical multistate occupancy models can forge these links by estimating spatial patterns of reproductive success across landscapes. To illustrate, we surveyed the occurrence of grizzly bears (Ursus arctos) in the Canadian Rocky Mountains Alberta, Canada. We deployed camera traps for 6 weeks at 54 surveys sites in different types of land cover. We used hierarchical multistate occupancy models to estimate probability of detection, grizzly bear occupancy, and probability of reproductive success at each site. Grizzly bear occupancy varied among cover types and was greater in herbaceous alpine ecotones than in low‐elevation wetlands or mid‐elevation conifer forests. The conditional probability of reproductive success given grizzly bear occupancy was 30% (SE = 0.14). Grizzly bears with cubs had a higher probability of detection than grizzly bears without cubs, but sites were correctly classified as being occupied by breeding females 49% of the time based on raw data and thus would have been underestimated by half. Repeated surveys and multistate modeling reduced the probability of misclassifying sites occupied by breeders as unoccupied to <2%. The probability of breeding grizzly bear occupancy varied across the landscape. Those patches with highest probabilities of breeding occupancy—herbaceous alpine ecotones—were small and highly dispersed and are projected to shrink as treelines advance due to climate warming. Understanding spatial correlates in breeding distribution is a key requirement for species conservation in the face of climate change and can help identify priorities for landscape management and protection. Patrones Espaciales del Éxito Reproductivo de Osos Pardos, Derivados de Modelos Jerárquicos Multi‐Estado     

The interaction between reproductive cost and individual quality is mediated by oceanic conditions in a long-lived bird

Environmental variability, costs of reproduction, and heterogeneity in individual quality are three important sources of the temporal and interindividual variations in vital rates of wild populations. Based on an 18-year monitoring of an endangered, recently described, long-lived seabird, Monteiro's Storm-Petrel (Oceanodroma monteiroi), we designed multistate survival models to separate the effects of the reproductive cost (breeders vs. nonbreeders) and individual quality (successful vs. unsuccessful breeders) in relation to temporally variable demographic and oceanographic properties. The analysis revealed a gradient of individual quality from nonbreeders, to unsuccessful breeders, to successful breeders. The survival rates of unsuccessful breeders (0.90 +/- 0.023, mean +/- SE) tended to decrease in years of high average breeding success and were more sensitive to oceanographic variation than those of both (high-quality) successful breeders (0.97 +/- 0.015) and (low-quality) nonbreeders (0.83 +/- 0.028). Overall, our results indicate that reproductive costs act on individuals of intermediate quality and are mediated by environmental harshness.     

The demographic consequences of the cost of reproduction in ungulates

The cost of reproduction can generate covariation between demographic rates that can potentially influence demography and population dynamics in long-lived iteroparous species. However, there has been relatively little work linking the survival cost of reproduction and population dynamics. The apparent scarcity of information on this important link is potentially due to covariation between vital rates, which can substantially influence fluctuations in population size. In this paper we examine the opportunity for survival costs of reproduction to leave a dynamic signature using a simulation model based broadly on an ungulate life history. We find that an increase in the cost delays the onset of reproduction and reduces reproductive rates of young, but not of prime-age, females. Accordingly, the number of offspring produced declines and the interval between reproductive events increases among young females experiencing high cost. These effects are translated to an age structure skewed toward young ages and reduced population density. These results suggest that, by delaying reproduction when conditions deteriorate, females protect their survival during the critical first three years of life, after which the negative effect of reproduction on survival declines. Unless conditions for reproduction are severe, it is not profitable to delay reproduction beyond age 3 years due to the high risk of death before having a chance to reproduce. We also demonstrate that lack of adjustment of reproductive strategies to elevated levels of the cost of reproduction, for example due to rapid changes in environmental conditions, results in lower average density and longevity compared to females that have sufficient time to adjust to changes in the cost. This suggests that even moderate costs of reproduction may have a major negative effect on population dynamics of ungulates.     

Age,size, dominance and reproduction among male kudus: mating enhancement by attrition of rivals

Summary Size dimorphism with males larger than females has been related to the benefits for males of enhanced dominance and hence reproductive success. However, mating gains must outweigh the fitness costs of deferred reproduction and the mortality associated with further growth. The relationships between male age, size and reproduction were assessed for greater kudus (Tragelaphus strepsiceros) in the Kruger National Park in South Africa. Individually identifiable animals were monitored over 10 years, with detailed observations made during six breeding seasons. In the non-breeding season males formed loose all-male groups. Horn grappling and low intensity agonistic interactions fostered dominance rankings. Dominance was age-graded, until males reached full weight at 6 years of age. Males aged 6 and 7 years monopolized courtship and mating, but 5-year-old males secured about 10% of mating opportunities. Few males survived beyond 7 years. Male mortality rate rose steeply with age, so that the functional sex ratio of fertile females per mature male was about 14:1. During the breeding season many female groups remained unattended by a mature male. Reproductive sorting among males occurred largely through variation in survival to full size and maturity. Increased size enhances fighting success and hence dominance. Further growth ceases when the functional sex ratio exceeds the number of mating opportunities that males can effectively achieve during a breeding season. Predation amplifies the mortality cost of continued growth. In the absence of large predators, male-male interactions may be atypically exaggerated.     

Effects of Multiple Levels of Social Organization on Survival and Abundance

Abstract: Identifying how social organization shapes individual behavior, survival, and fecundity of animals that live in groups can inform conservation efforts and improve forecasts of population abundance, even when the mechanism responsible for group‐level differences is unknown. We constructed a hierarchical Bayesian model to quantify the relative variability in survival rates among different levels of social organization (matrilines and pods) of an endangered population of killer whales (Orcinus orca). Individual killer whales often participate in group activities such as prey sharing and cooperative hunting. The estimated age‐specific survival probabilities and survivorship curves differed considerably among pods and to a lesser extent among matrilines (within pods). Across all pods, males had lower life expectancy than females. Differences in survival between pods may be caused by a combination of factors that vary across the population's range, including reduced prey availability, contaminants in prey, and human activity. Our modeling approach could be applied to demographic rates for other species and for parameters other than survival, including reproduction, prey selection, movement, and detection probabilities.     

Brood size manipulation affects frequency of second clutches in the blue tit

The reproductive trade-off hypothesis predicts that the investment made in current reproduction determines the breeders’ future fitness as a consequence of intra-or inter-generational reproductive costs. Long-lived species are expected to favour their own reproductive value at the expense of their offspring, hence incurring in inter-generational costs, whereas short-lived species are expected to invest in the current breeding attempt even at the expense of their own survival, thus incurring in intra-generational costs. We tested whether intensity of current reproductive effort has intra-or inter-generational costs in a short-lived bird, the blue tit Parus caeruleus, with a brood size manipulation experiment. We expected more intra-generational (parental reproduction and/or survival) than inter-generational (offspring quality and survival) reproductive costs. We found that parental effort, measured as the hourly rate of parental visits to nests, increased gradually with experimental manipulation. Brood size manipulation resulted in a gradual increase in the number of fledglings per nest from reduced to increased treatments. We found an effect of the manipulation on the probability of making a second clutch, with adults rearing enlarged broods being less likely to undertake such a second reproduction during the season compared to those rearing control or decreased broods. We found no evidence of other reproductive costs; neither as adult weight after manipulation, apparent parental local survival, apparent offspring local survival or local recruitment. Although the results seem to support the a priori expectations, alternative explanations are discussed.Communicated by M. Soler     

Forest Roads as Partial Barriers to Terrestrial Salamander Movement

Abstract:  Roads can fragment animal populations by disrupting movement among formerly continuous habitats. Although models have demonstrated that disrupted movement can contribute to long-term extinction, there are few empirical data on the effects of roads on animal movement. We used displacement and homing experiments to determine whether forest roads are barriers to the movement of terrestrial salamanders. We displaced 1471 red-backed salamanders ( Plethodon cinereus ) across five forest roads and compared return rates to those of salamanders displaced equal distances toward the forest interior. Roads significantly reduced the return rate of salamanders, with a mean reduction of 51%. Steep roadside verges further reduced return rates, particularly for salamanders moving downhill across verges. The permeability of roads to salamander movement did not appear to be related to road surface type. Gravel roads had both the highest and lowest observed permeability with the two paved roads intermediate between these. We conclude that narrow forest roads are partial barriers to salamander movement and that steep roadside verges may exacerbate these effects.     

Male mating effort in a polygynous ungulate

Survival and mating success are key fitness components of mammalian males. Because energy is limited, life history theory predicts trade-offs between current and future reproduction. While many studies have examined fitness trade-off in females, we know little about trade-offs faced by males. In polygynous species, male mating success is largely dependent upon intrasexual competition. Consequently, males have greater uncertainty over the benefits of a given allocation than over its costs, and the correlation between mating effort and success is likely much weaker in males than in females. We analyzed 14 years of data on the mating effort and survival of marked bighorn rams to investigate fitness trade-offs. Dominant rams defended single estrous ewes (“tending”) while subordinates attempted to copulate after separating the tending pair (“coursing”). We estimated the participation in tending and coursing for each ram and the effort in searching for breeding opportunities by each ram each year. We compared these three behavioral indices of male mating effort to demographic parameters, individual characteristics, and both yearly and long-term survival. Mating effort during the rut was unrelated to ram overwinter survival, but longevity was positively correlated with mating effort between 2 and 5 years of age. Persistent variation among rams is likely to explain this pattern, suggesting that in natural populations a few high quality males enjoy both high mating success and high survival.An erratum to this article can be found at     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Patterns in reproductive dynamics of burrowing ghost shrimp Trypaea australiensis from small to intermediate scales

Many studies have examined latitudinal differences in reproduction of marine invertebrates, but few have measured variation at small to intermediate scales (kilometres to hundreds of kilometres), which may confound comparisons across broader geographic regions. Here, we examined variation in the reproductive biology of a little-studied species of burrowing ghost shrimp (Trypaea australiensis) at spatial scales ranging from km (between sites within estuaries) to 100s of km (among estuaries), over a 2-year period in south-eastern Australia. Sex ratios of populations were consistently biased towards females through time and space. Although reproduction started in summer months across all spatial scales, there was a pattern of earlier spawning from southern to northern estuaries. Integration of results from previous studies of T. australiensis supported a similar pattern of earlier breeding from high to low latitudes. Fecundity of shrimp increased linearly with female size, but the relationship varied inconsistently across the different spatial scales. Similarly, sizes at maturity varied from small to intermediate scales and observed patterns were not consistent with general predictions e.g. shrimp were smaller and ovigerous at smaller sizes at sites in the southern-most estuary, compared to estuaries further north. We found no differences in the sizes of embryos across the different spatial scales, but confirm that T. australiensis employs a strategy of high fecundity and small embryo size compared to other thalassinidean shrimp. Our results suggest that factors at smaller scales (e.g. food availability) may be important in affecting reproductive dynamics of T. australiensis, but further research is needed in testing hypotheses about patterns observed here. A lack of similar studies on other marine organisms remains an impediment to understanding life-history strategies and the sustainable management and conservation of populations.     

Effects of Urbanization on Occupancy of Stream Salamanders

Abstract: Urban development is the most common form of land conversion in the United States. Using a before–after control‐impact study design, we investigated the effects of urbanization on larval and adult stages of southern two‐lined salamanders (Eurycea cirrigera) and northern dusky salamanders (Desmognathus fuscus). Over 5 years, we estimated changes in occupancy and probabilities of colonization and survival in 13 stream catchments after urbanization and in 17 catchments that were not urbanized. We also examined effects of proportion of urbanized area in a catchment and distance of the salamander population to the nearest stream on probabilities of colonization and survival. Before urbanization, adult and larval stages of the two salamander species occupied nearly all surveyed streams, with occupancy estimates ranging from 1.0 to 0.78. Four years after urbanization mean occupancy of larval and adult two‐lined salamanders had decreased from 0.87 and 0.78 to 0.57 and 0.39, respectively. Estimates of mean occupancy of larval northern dusky salamanders decreased from 1.0 to 0.57 in urban streams 4 years after urbanization; however, adult northern dusky salamander occupancy remained close to 1.0 in urban streams over 5 years. Occupancy estimates in control streams were similar for each species and stage over 5 years. Urbanization was associated with decreases in survival probabilities of adult and larval two‐lined salamanders and decreases in colonization probabilities of larval dusky salamanders. Nevertheless, proportion of impervious surface and distance to nearest stream had little effect on probabilities of survival and colonization. Our results imply that in the evaluation of the effects of urbanization on species, such as amphibians, with complex life cycles, consideration of the effects of urbanization on both adult and larval stages is required.     

Floater mortality within settlement areas can explain the Allee effect in breeding populations

The Allee effect (the positive relationship between population growth rate and population size) is a constraint of some animal populations at low numbers, which increases their likelihood of extinction because of a decrease in reproduction and/or survival. We were able to demonstrate that the Allee effect can be the result of a mortality increase affecting floaters (i.e. dispersing individuals able to enter as breeders in the reproductive population when a breeding territory or a potential mate – owner of a suitable breeding territory – becomes available). Previously, potential mechanisms underlying Allee effects were always related to the breeding portion of a population only. In contrast, our understanding of or solutions to population declines due to the Allee effects can reside elsewhere, away from breeding territories.     

Lethal Effects of Water Quality on Threatened California Salamanders but Not on Co‐Occurring Hybrid Salamanders

Biological invasions and habitat alteration are often detrimental to native species, but their interactions are difficult to predict. Interbreeding between native and introduced species generates novel genotypes and phenotypes, and human land use alters habitat structure and chemistry. Both invasions and habitat alteration create new biological challenges and opportunities. In the intensively farmed Salinas Valley, California (U.S.A.), threatened California tiger salamanders (Ambystoma californiense) have been replaced by hybrids between California tiger salamander and introduced barred tiger salamanders (Ambystoma tigrinum mavortium). We conducted an enclosure experiment to examine the effects habitat modification and relative frequency of hybrid and native California tiger salamanders have on recruitment of salamanders and their prey, Pacific chorus frogs (Pseudacris regilla). We tested whether recruitment differed among genetic classes of tiger salamanders (hybrid or native) and pond hydroperiod (seasonal or perennial). Roughly 6 weeks into the experiment, 70% (of 378 total) of salamander larvae died in 4 out of 6 ponds. Native salamanders survived (n = 12) in these ponds only if they had metamorphosed prior to the die‐offs. During die‐offs, all larvae of native salamanders died, whereas 56% of hybrid larvae died. We necropsied native and hybrid salamanders, tested water quality, and queried the California Department of Pesticide Regulation database to investigate possible causes of the die‐offs. Salamander die‐offs, changes in the abundance of other community members (invertebrates, algae, and cyanobacteria), shifts in salamander sex ratio, and patterns of pesticide application in adjacent fields suggest that pesticide use may have contributed to die‐offs. That all survivors were hybrids suggests that environmental stress may promote rapid displacement of native genotypes. Efectos Letales de la Calidad del Agua sobre Salamandras de California Amenazadas pero no sobre Salamandras Híbridas Concurrentes     

Antipredator defenses along a latitudinal gradient in Rana temporaria

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.     

Range-wide effects of breeding- and nonbreeding-season climate on the abundance of a Neotropical migrant songbird

Geographic variation in the population dynamics of a species can result from regional variability in climate and how it affects reproduction and survival. Identifying such effects for migratory birds requires the integration of population models with knowledge of migratory connectivity between breeding and nonbreeding areas. We used Bayesian hierarchical models with 26 years of Breeding Bird Survey data (1982-2007) to investigate the impacts of breeding- and nonbreeding-season climate on abundance of American Redstarts (Setophaga ruticilla) across the species range. We focused on 15 populations defined by Bird Conservation Regions, and we included variation across routes and observers as well as temporal trends and climate effects. American Redstart populations that breed in eastern North America showed increased abundance following winters with higher plant productivity in the Caribbean where they are expected to overwinter. In contrast, western breeding populations showed little response to conditions in their expected wintering areas in west Mexico, perhaps reflecting lower migratory connectivity or differential effects of winter rainfall on individuals across the species range. Unlike the case with winter climate, we found few effects of temperature prior to arrival in spring (March-April) or during the nesting period (May-June) on abundance the following year. Eight populations showed significant changes in abundance, with the steepest declines in the Atlantic Northern Forest (-3.4%/yr) and the greatest increases in the Prairie Hardwood Transition (4%/yr). This study emphasizes how the effects of climate on populations of migratory birds are context dependent and can vary depending on geographic location and the period of the annual cycle. Such knowledge is essential for predicting regional variation in how populations of a species might vary in their response to climate change.     

Older can be better: physiological costs of paternal investment in the Florida scrub-jay

In species that undergo actuarial senescence, the value of current reproduction is predicted to increase relative to the value of future reproduction with age, as the probability of survival to another reproductive event is reduced. Therefore, life history theory predicts that aging animals should increase their investment in reproduction. However, an increase in reproductive investment may carry significant costs to the breeding individuals. We recorded provisioning rates of Florida scrub-jay male breeders, followed by their immediate capture to assess body condition and collect blood for an in vitro test of immunocompetence and an assay of baseline corticosterone for a measure of stress. Older males provisioned offspring and brooding mates at the highest rates. There was no evidence of any physiological deficits in males with high provisioning rates, independent of age. It appears that birds that survive to old age are high quality birds that maintain good physiological condition, which complements the value of experience and permits maximal investment in offspring.