Intermittent Breeding Research Articles

The Ne/N ratio in applied conservation.

Recent developments within the IUCN and the Convention on Biological Diversity have affirmed the increasingly key role that effective population size (N e) and the effective size: census size ratio (N e/N) play in applied conservation and management of global biodiversity. This paper reviews and synthesizes information regarding the definition of N e and demographic and genetic methods for estimating effective size, census size, and their ratio. Emphasis is on single-generation estimates of contemporary N e/N, which are the most informative for practical applications. It is crucial to clearly define which individuals are included in the census size (N). Defining N as the number of adults alive at a given time facilitates comparisons across species. For a wide range of applications and experimental designs, inbreeding N e is simpler to calculate and interpret than variance N e. Effects of skewed sex ratio are generally modest, so most reductions to N e/N arise from overdispersed (greater-than-Poisson) variance in offspring number (). Even when fecundity changes with age, overdispersed within-age variance generally contributes most to overall , and both random and deterministic (mediated by selection) factors can be important. Most species are age-structured, so it is important to distinguish between effective size per generation (N e) and the effective number of breeders in one season or year (N b). Both N e and N b are important for applied conservation and management. For iteroparous species, a key metric is variance in lifetime reproductive success (), which can be affected by a variety of additional factors, including variation in longevity, skip or intermittent breeding, and persistent individual differences in reproductive success. Additional factors that can be important for some species are also discussed, including mating systems, population structure, sex reversal, reproductive compensation, captive propagation, and delayed maturity.

Accounting for unobserved population dynamics and aging error in close-kin mark-recapture assessments.

Obtaining robust estimates of population abundance is a central challenge hindering the conservation and management of many threatened and exploited species. Close-kin mark-recapture (CKMR) is a genetics-based approach that has strong potential to improve the monitoring of data-limited species by enabling estimates of abundance, survival, and other parameters for populations that are challenging to assess. However, CKMR models have received limited sensitivity testing under realistic population dynamics and sampling scenarios, impeding the application of the method in population monitoring programs and stock assessments. Here, we use individual-based simulation to examine how unmodeled population dynamics and aging uncertainty affect the accuracy and precision of CKMR parameter estimates under different sampling strategies. We then present adapted models that correct the biases that arise from model misspecification. Our results demonstrate that a simple base-case CKMR model produces robust estimates of population abundance with stable populations that breed annually; however, if a population trend or non-annual breeding dynamics are present, or if year-specific estimates of abundance are desired, a more complex CKMR model must be constructed. In addition, we show that CKMR can generate reliable abundance estimates for adults from a variety of sampling strategies, including juvenile-focused sampling where adults are never directly observed (and aging error is minimal). Finally, we apply a CKMR model that has been adapted for population growth and intermittent breeding to two decades of genetic data from juvenile lemon sharks (Negaprion brevirostris) in Bimini, Bahamas, to demonstrate how application of CKMR to samples drawn solely from juveniles can contribute to monitoring efforts for highly mobile populations. Overall, this study expands our understanding of the biological factors and sampling decisions that cause bias in CKMR models, identifies key areas for future inquiry, and provides recommendations that can aid biologists in planning and implementing an effective CKMR study, particularly for long-lived data-limited species.

Earlier springs increase goose breeding propensity and nesting success at Arctic but not at temperate latitudes.

Intermittent breeding is an important tactic in long-lived species that trade off survival and reproduction to maximize lifetime reproductive success. When breeding conditions are unfavourable, individuals are expected to skip reproduction to ensure their own survival. Breeding propensity (i.e. the probability for a mature female to breed in a given year) is an essential parameter in determining reproductive output and population dynamics, but is not often studied in birds because it is difficult to obtain unbiased estimates. Breeding conditions are especially variable at high latitudes, potentially resulting in a large effect on breeding propensity of Arctic-breeding migratory birds, such as geese. With a novel approach, we used GPS-tracking data to determine nest locations, breeding propensity and nesting success of barnacle geese, and studied how these varied with breeding latitude and timing of arrival on the breeding grounds relative to local onset of spring. Onset of spring at the breeding grounds was a better predictor of breeding propensity and nesting success than relative timing of arrival. At Arctic latitudes (>66° N), breeding propensity decreased from 0.89 (95% CI: 0.65-0.97) in early springs to 0.22 (95% CI: 0.06-0.55) in late springs, while at temperate latitudes, it varied between 0.75 (95% CI: 0.38-0.93) and 0.89 (95% CI: 0.41-0.99) regardless of spring phenology. Nesting success followed a similar pattern and was lower in later springs at Arctic latitudes, but not at temperate latitudes. In early springs, a larger proportion of geese started breeding despite arriving late relative to the onset of spring, possibly because the early spring enabled them to use local resources to fuel egg laying and incubation. While earlier springs due to climate warming are considered to have mostly negative repercussions on reproductive success through phenological mismatches, our results suggest that these effects may partly be offset by higher breeding propensity and nesting success.

Conservation implications of a mismatch between data availability and demographic impact.

Cost-effective use of limited conservation resources requires understanding which data most contribute to alleviating biodiversity declines. Interventions might reasonably prioritise life-cycle transitions with the greatest influence on population dynamics, yet some contributing vital rates are particularly challenging to document. This risks managers making decisions without sufficient empirical coverage of the spatiotemporal variation experienced by the species. Here, we aimed to explore whether the number of studies contributing estimates for a given life-stage transition aligns with that transition's demographic impact on population growth rate, λ. We parameterised a matrix population model using meta-analysis of vital rates for the common eider (Somateria mollissima), an increasingly threatened yet comparatively data-rich species of seaduck, for which some life stages are particularly problematic to study. Female common eiders exhibit intermittent breeding, with some established breeders skipping one or more years between breeding attempts. Our meta-analysis yielded a breeding propensity of 0.72, which we incorporated into our model with a discrete and reversible 'nonbreeder' stage (to which surviving adults transition with a probability of 0.28). The transitions between breeding and nonbreeding states had twice the influence on λ than fertility (summed matrix-element elasticities of 24% and 11%, respectively), whereas almost 15 times as many studies document components of fertility than breeding propensity (n = 103 and n = 7, respectively). The implications of such mismatches are complex because the motivations for feasible on-the-ground conservation actions may be different from what is needed to reduce uncertainty in population projections. Our workflow could form an early part of the toolkit informing future investment of finite resources, to avoid repeated disconnects between data needs and availability thwarting evidence-led conservation.

Intermittent breeding is associated with breeding group turnover in a cooperatively breeding bird.

Intermittent breeding, in which an adult skips a breeding opportunity, can represent a non-adaptive constraint or an adaptive response to the tradeoff between current and future reproduction. In group-living animals, the social group may also affect the frequency of reproduction, but this possibility has received little attention. Here we use an 11-year data set to investigate intermittent breeding in the greater ani (Crotophaga major), a tropical bird that nests in stable breeding groups containing several unrelated co-breeding females. Population-wide, an average of 62% of females laid eggs in a given year (range 35-84%), and the average female failed to lay eggs once every 3.2years. We found little support for the hypothesis that intermittent breeding reflects a tradeoff between current and future reproduction: breeding in year t did not affect a female's likelihood of breeding in year t + 1, and clutch size in year t did not affect clutch size in year t + 1. Increases in clutch size were associated with decreases in egg mass for eggs laid at the end of that clutch, but this did not affect subsequent nesting attempts. However, reproductive skipping was associated with changes in group membership. Females whose groups changed in composition after year t were significantly less likely to breed in year t + 1 than females whose groups remained stable. These results indicate that breeding group stability influences the frequency of reproduction, suggesting that transitions between groups may be costly to females and their mates.

Reproductive skipping as an optimal life history strategy in the southern elephant seal, Mirounga leonina.

Intermittent breeding by which organisms skip some current reproductive opportunities in order to enhance future reproductive success is a common life history trade‐off among long‐lived, iteroparous species. The southern elephant seal Mirounga leonina engages in intermediate breeding when body condition is low. While it is anticipated that this strategy may increase the lifetime reproductive output of this species, the conditions under which reproductive skipping are predicted to occur are not clear. Here I develop a dynamic state variable model based on published data that examines when southern elephant seals are predicted to optimally skip reproduction in order to maximize lifetime reproductive output as a function of current body mass, maternal age, and survivorship. I demonstrate that the optimal reproductive strategy for this species can include reproductive skipping, and that the conditions where this is optimal depend on patterns of mass‐dependent adult female survival. I further show that intermittent breeding can increase lifetime reproductive output, and that the magnitude of this benefit increases with the ability of individual animals to replenish depleted body mass through foraging. Finally, I show that when the environment is variable and foraging is reduced in bad years, the benefit of adopting an optimal strategy that includes reproductive skipping increases asymptotically with the frequency of bad years. These results highlight the importance of characterizing the pattern of adult survival in this species, as well as the need to identify other factors that may influence the prevalence and benefits of reproductive skipping.

To breed or not to breed: drivers of intermittent breeding in a seabird under increasing predation risk and male bias.

Intermittent breeding may be adaptive for long-lived species subjected to large accessory reproductive costs, but it may also reflect reduced adaptation to the environment, reducing population growth. Nevertheless, environmental influences on breeding propensity, particularly that of predation risk, remain poorly understood and difficult to study, because non-breeders are typically not identified. Female eiders Somateria mollissima from the Baltic Sea provide an excellent testbed, because nesting females have been exposed to intensifying predation and growing male bias that may increase female harassment. We based our study on long-term data (14years) on females captured and marked at the nest, and females individually identified at sea irrespective of capture status. We hypothesized that breeding propensity decreases with increasing predation risk and male bias, and increases with breeder age. Consistent with our hypotheses, females nesting on islands with higher nest predation risk were more likely to skip breeding, and breeding probability increased with age. In contrast, the steep temporal decline in breeding propensity could not be reliably attributed to annual adult sex ratio or to the abundance of white-tailed sea eagles (Haliaeetus albicilla), the main predator on females, at the nearby Hanko Bird Observatory. Breeding probability showed significant consistent individual variation. Our results demonstrate that spatiotemporal variation in predation risk affects the decision to breed and high incidence of non-breeding was associated with low fledging success. The increased frequency of intermittent breeding in this declining population should be explicitly considered in demographic models, and emphasis placed on understanding the preconditions for successful reproduction.

Optimizing lifetime reproductive output: Intermittent breeding as a tactic for females in a long-lived, multiparous mammal.

In iteroparous species, intermittent breeding is an important life-history tactic that can greatly affect animal population growth and viability. Despite its importance, few studies have quantified the consequences of breeding pauses on lifetime reproductive output, principally because calculating lifetime reproductive output requires knowledge of each individual's entire reproductive history. This information is extremely difficult to obtain in wild populations. We applied novel statistical approaches that account for uncertainty in state assessment and individual heterogeneity to an 18-year capture-recapture dataset of 6,631 female southern elephant seals from Macquarie Island. We estimated survival and breeding probabilities, and investigated the consequences of intermittent breeding on lifetime reproductive output. We found consistent differences in females' demographic performance between two heterogeneity classes. In particular, breeding imbued a high cost on survival in the females from the heterogeneity class 2, assumed to be females of lower quality. Individual quality also appeared to play a major role in a female's decision to skip reproduction with females of poorer quality more likely to skip breeding events than females of higher quality. Skipping some breeding events allowed females from both heterogeneity classes to increase lifetime reproductive output over females that bred annually. However, females of lower quality produced less offspring over their lifetime. Intermittent breeding seems to be used by female southern elephant seals as a tactic to offset reproductive costs on survival and enhance lifetime reproductive output but remains unavoidable and driven by individual-specific constraints in some other females.

Extreme climate events and individual heterogeneity shape life‐history traits and population dynamics

Extreme climatic conditions and their ecological impacts are currently emerging as critical features of climate change. We studied extreme sea ice condition (ESIC) and found it impacts both life‐history traits and population dynamics of an Antarctic seabird well beyond ordinary variability.The Southern Fulmar (Fulmarus glacialoides) is an ice‐dependent seabird, and individuals forage near the ice edge. During an extreme unfavorable year (when sea ice area is reduced and distance between ice edge and colony is high), observed foraging trips were greater in distance and duration. As a result, adults brought less food to their chicks, which fledged in the poorest body condition. During such unfavorable years, breeding success was extremely low and population growth rate (λ) was greatly reduced. The opposite pattern occurred during extreme favorable years.Previous breeding status had a strong influence on life‐history traits and population dynamics, and their responses to extreme conditions. Successful breeders had a higher chance of breeding and raising their chick successfully during the following breeding season as compared to other breeding stages, regardless of environmental conditions. Consequently, they coped better with unfavorable ESIC. The effect of change in successful breeder vital rates on λ was greater than for other stages' vital rates, except for pre‐breeder recruitment probabilities, which most affected λ.For environments characterized by ordinary sea ice conditions, interindividual differences were more likely to persist over the life of individuals and randomness in individual pathways was low, suggesting individual heterogeneity in vital rates arising from innate or acquired phenotypic traits. Additionally, unfavorable ESIC tended to exacerbate individual differences in intrinsic quality, expressed through differences in reproductive status.We discuss the strong effects of ESIC on Southern Fulmar life‐history traits in an evolutionary context. ESICs strongly affect fitness components and act as potentially important agents of natural selection of life histories related to intrinsic quality and intermittent breeding. In addition, recruitment is a highly plastic trait that, if heritable, could have a critical role in evolution of life histories. Finally, we find that changes in the frequency of extreme events may strongly impact persistence of Southern Fulmar populations.

The influence of environmental conditions on the age pattern in breeding performance in a transequatorial migratory seabird

Several studies of marine top predators, above all of seabirds, have analysed the effects of either individual age or environmental fluctuations on reproduction; nevertheless, little is known about the age patterns in breeding performance in a variable environment. To investigate the simultaneous influence of age and environmental conditions on laying dates and egg volumes, we tested different climate and food availability indices in a transequatorial migratory seabird using female data from a 23-year study. Our results show an initial improvement in breeding parameters with age (i.e. earlier laying dates and greater egg volumes) but no pattern of senescence in older age groups. The best models showed an interaction of time and age in breeding performance, i.e. the age pattern of breeding performance changed each year likely as a result of environmental variability. Nevertheless, climatic indexes used here explained part of that annual variability: NAO and SOI index accounted for 24% and 20% of deviances in laying dates and egg volume respectively. Part of that unexplained variability might be related to other processes such as intermittent breeding and the individual quality of breeders, which were not assessed in our study.

INTERMITTENT BREEDING AND CONSTRAINTS ON LITTER SIZE: CONSEQUENCES FOR EFFECTIVE POPULATION SIZE PER GENERATION (Ne) AND PER REPRODUCTIVE CYCLE (Nb)

In iteroparous species, it is easier to estimate Nb (effective number of breeders in one reproductive cycle) than Ne (effective population size per generation). Nb can be used as a proxy for Ne and also can provide crucial insights into eco-evolutionary processes that occur during reproduction. We used analytical and numerical methods to evaluate effects of intermittent breeding and litter/clutch size on inbreeding Nb and Ne . Fixed or random litter sizes ≥ 3 have little effect on either effective-size parameter; however, in species (e.g., many large mammals) in which females can produce only one offspring per cycle, female Nb = ∞ and overall Nb = 4Nb (male) . Intermittent breeding reduces the pool of female breeders, which reduces both female and overall Nb ; reductions are larger in high-fecundity species with high juvenile mortality and increase when multiple reproductive cycles are skipped. Simulated data for six model species showed that both intermittent breeding and litter-size constraints increase Ne , but only slightly. We show how to quantitatively account for these effects, which are important to consider when (1) using Nb to estimate Ne , or (2) drawing inferences about male reproductive success based on estimates of female Nb .

Genetic variation and selection within glandless cotton germplasm

There is a renewed interest in research and use of glandless (free of gossypol) Upland cotton (Gossypium hirsutum L.) that can produce edible seeds for human food and animal feed. However, there was a lack of information on yield potential of existing glandless germplasm since intermittent breeding activities for glandless cotton were stopped in the U.S. before 2000. The objectives of the present study were to test obsolete and exotic glandless germplasm for possible production and evaluate progress in direct pedigree selection within the existing glandless germplasm in eight field tests. In multiple tests, the glandless cotton Acala GLS yielded only 65–80 % lint of the glanded control Acala 1517-08 and 46–75 % lint of commercial transgenic cultivars. Most of obsolete glandless germplasm and their selections yielded <70 % of Acala 1517-08 and only three selections yielded 82–89 % of the control. Genetic variation in yield and fiber quality traits was seen from significant differences between selections within the same glandless germplasm, indicating the existence of residual genetic variation. Most importantly, three selections from exotic glandless germplasm out-yielded Acala 1517-08 by 4–9 %, one of which had desirable fiber quality traits. These promising glandless lines with comparable yields to commercial cotton make it possible to revitalize the glandless cotton industry. More field tests will be needed to evaluate their yield potential and adaptability in the U.S. Cotton Belt.

Intermittent breeding in the absence of a large cost of reproduction: evidence for a non‐migratory, iteroparous salmonid

In long‐lived organisms, intermittent breeding likely evolves as a resource allocation strategy for coping with environmental uncertainty or individual heterogeneity in condition. In fishes, the phenomenon of intermittent breeding is referred to as skipped spawning, and appears to be more common at high latitudes or in migratory species with high accessory costs of reproduction. We used long‐term monitoring data on lake trout (Salvelinus namaycush) to test whether key predictions about the frequency of skipped spawning hold in a mid‐latitude population of a species lacking any obvious costs of reproduction beyond the production and fertilization of gametes. We first developed a threshold‐based method to classify skipped spawners based on gonad size, fish size, and fish age. Consistent with life history theory, age‐specific frequencies of skipped spawning were higher in females than males. The frequency of skipped spawning varied among years and was higher in 1994–2011 than in 1938–1959, perhaps because of food web changes over the past century. In temperate lakes, food web structure may be sufficiently variable to favor intermittent breeding in long‐lived iteroparous fishes, despite low accessory costs of reproduction.

Intermittent breeding and the dynamics of resource allocation to reproduction, growth and survival

Summary In iteroparous organisms, the timing of reproduction varies greatly and reproduction may be intermittent. Here, we investigate the proximate determinants of intermittent breeding and their evolutionary significance in a viviparous snake, the meadow viper Vipera ursinii ursinii. We used individual life‐history data collected over a 28‐year capture–mark–recapture field study. Among viviparous snakes, the meadow viper's almost exclusively insectivorous diet is remarkable. Breeding females maintain substantial foraging activity during gestation. The life cycle is iteroparous, and female breeding frequency is essentially biennial. Breeding status is strongly associated with a threshold on female body condition prior to vitellogenesis. Non‐breeding females grow and store fat reserves needed for future reproduction. Breeding females convert reserves into eggs, the number of which is determined by maternal size and body condition prior to vitellogenesis. During pregnancy, females stop growing. Females survive breeding and non‐breeding years with equal probabilities. Our results suggest that income resources during pregnancy are used to fully cover the costs of gestation. Integration of our data into a matrix population model allowed us to relate breeding pattern to fitness. According to the model, biennial breeding is strongly favoured by natural selection if consecutive breeding decreases maternal survival and/or fecundity. Growth cessation during breeding years generates weak selection pressures for intermittent breeding.

The evolution of intermittent breeding

A central issue in life history theory is how organisms trade off current and future reproduction. A variety of organisms exhibit intermittent breeding, meaning sexually mature adults will skip breeding opportunities between reproduction attempts. It's thought that intermittent breeding occurs when reproduction incurs an extra cost in terms of survival, energy, or recovery time. We have developed a matrix population model for intermittent breeding, and use adaptive dynamics to determine under what conditions individuals should breed at every opportunity, and under what conditions they should skip some breeding opportunities (and if so, how many). We also examine the effect of environmental stochasticity on breeding behavior. We find that the evolutionarily stable strategy (ESS) for breeding behavior depends on an individual's expected growth and mortality, and that the conditions for skipped breeding depend on the type of reproductive cost incurred (survival, energy, recovery time). In constant environments there is always a pure ESS, however environmental stochasticity and deterministic population fluctuations can both select for a mixed ESS. Finally, we compare our model results to patterns of intermittent breeding in species from a range of taxonomic groups.

Now you see him, now you don't: experience, not age, is related to reproduction in kittiwakes

In long-lived species, individuals can skip reproduction. The proportion of breeders affects population growth rate and viability, there is a need to investigate the factors influencing intermittent breeding. The theory predicts that if lack of experience is an important constraint, breeding probabilities should increase with experience for individuals of the same age, whereas under the so-called restraint hypothesis, breeding probabilities should increase with age regardless of experience. However, because the probability of detecting individuals in the wild is generally less than 1, it is difficult to know exactly the number of previous breeding episodes (breeding experience). To cope with this issue, we developed a hidden process model to incorporate experience as a latent state possibly influencing the probability of breeding. Using a 22-year mark-recapture dataset involving 9970 individuals, we analysed simultaneously experience and age effects on breeding probabilities in the kittiwake (Rissa tridactyla). We did not detect an influence of age on adult breeding probabilities. We found that inexperienced birds breed less frequently than experienced birds. Our approach enables us to highlight the key role of experience on adults breeding probabilities and can be used for a wide range of organisms for which detection is less than 1.

To breed or not to breed: a seabird's response to extreme climatic events

Intermittent breeding is an important life-history strategy that has rarely been quantified in the wild and for which drivers remain unclear. It may be the result of a trade-off between survival and reproduction, with individuals skipping breeding when breeding conditions are below a certain threshold. Heterogeneity in individual quality can also lead to heterogeneity in intermittent breeding. We modelled survival, recruitment and breeding probability of the red-footed booby (Sula sula), using a 19 year mark-recapture dataset involving more than 11,000 birds. We showed that skipping breeding was more likely in El-Niño years, correlated with an increase in the local sea surface temperature, supporting the hypothesis that it may be partly an adaptive strategy of birds to face the trade-off between survival and reproduction owing to environmental constraints. We also showed that the age-specific probability of first breeding attempt was synchronized among different age-classes and higher in El-Niño years. This result suggested that pre-breeders may benefit from lowered competition with experienced breeders in years of high skipping probabilities.

Intermittent breeding as a cost of site fidelity

Intermittent breeding (skipping a breeding season) can be the result of an adaptive decision by a focal individual, trading current reproductive success in favour of future reproductive success (residual reproductive value hypothesis). In contrast, an individual can also be forced by conspecifics to abandon the familiar breeding site and refrain from breeding due to lack of suitable alternative breeding sites or mates (competition hypothesis). I studied intermittent breeding in the territorial and site-faithful Eurasian oystercatcher Haematopus ostralegus, using a dataset covering 20 years. Intermittent breeding (in total 86 cases) occurred among breeders that formerly bred in high- as well as low-quality territories. The main factor associated with intermittent breeding in high-quality sites was death of a mate, while in low quality sites divorce was the most prominent factor. In 93% of the cases birds were forced to cease breeding due to pressure from conspecifics consistent with the competition hypothesis. There was no association between intermittent breeding and promotion to a territory of better quality. Instead, oystercatchers returned to breeding habitat of similar quality and at a very close distance (median distance 128 m) from the previous breeding location. Breeding absences lasted on average 2.4 years, with a maximum of 9 years, and the quality of the territory obtained after the absence varied with the duration of it. Birds who re-bred in a high-quality territory acquired this on average faster than those that re-bred in a low-quality territory, indicating that birds in high-quality sites are better competitors.

Trade-off between current reproductive effort and delay to next reproduction in the leatherback sea turtle

The trade-off between current and future reproduction plays an important role in demographic analyses. This can be revealed by the relationship between the number of years without reproduction and reproductive investment within a reproductive year. However, estimating both the duration between two successive breeding season and reproductive effort is often limited by variable recapture or resighting effort. Moreover, a supplementary difficulty is raised when nonbreeder individuals are not present sampling breeding grounds, and are therefore unobservable. We used capture-recapture (CR) models to investigate intermittent breeding and reproductive effort to test a putative physiological trade-off in a long-lived species with intermittent breeding, the leatherback sea turtle. We used CR data collected on breeding females on Awa:la-Ya:lima:po beach (French Guiana, South America) from 1995 to 2002. By adding specific constraints in multistate (MS) CR models incorporating several nonobservable states, we modelled the breeding cycle in leatherbacks and then estimated the reproductive effort according to the number of years elapsed since the last nesting season. Using this MS CR framework, the mean survival rate was estimated to 0.91 and the average resighting probability to 0.58 (ranged from 0.30 to 0.99). The breeding cycle was found to be limited to 3 years. These results therefore suggested that animals whose observed breeding intervals are greater than 3 years were most likely animals that escaped detection during their previous nesting season(s). CR data collected in 2001 and 2002 allowed us to compare the individual reproductive effort between females that skipped one breeding season and females that skipped two breeding seasons. These inferences led us to conclude that a trade-off between current and future reproduction exists in leatherbacks nesting in French Guiana, likely linked to the resource provisioning required to invest in reproduction.

Adult survival and temporary emigration in the common toad

In vertebrates exhibiting intermittent breeding, breeding activity is a factor of critical importance in capture–recapture studies using data from individually marked animals. Nonbreeders can be absent from locations used by breeders and can be considered "temporary emigrants". We addressed the influence of sex on survival in common toads, Bufo bufo (L., 1758), using the Joly–Seber model and the existance of temporary emigration in male common toads by assessing trap-dependence and by conducting a robust design analysis. We addressed the hypothesis that the probability of the presence of an individual in the study area depends on the presence of the individual the year before (i.e., that transitions between reproductive states are a first-order Markovian process). Results provided support for the hypotheses of random temporary emigration, of sex-specific differences in survival, and of the presence of "transients" in males. Females had intermediate survival compared with the groups of "transients + residents" and "resident" males. Females had lower recapture probability under the Joly–Seber model, which may be interpreted as evidence of lower breeding probability or lower detectability of breeding females. Behaviour may explain this difference in that females may attend ponds for shorter periods. This may be common in species where females aggregate to seek fertilization and lay eggs in locations attended by males and in species with a "resource-based lek" mating system.