Probability Of Breeding Research Articles

Survival and Breeding Transitions for a Reintroduced Bison Population: a Multistate Approach

The iconic plains bison (Bison bison) have been reintroduced to many places in their former range, but there are few scientific data evaluating the success of these reintroductions or guiding the continued management of these populations. Relying on mark–recapture data, we used a multistate model to estimate bison survival and breeding transition probabilities while controlling for the recapture process. We tested hypotheses in these demographic parameters associated with age, sex, reproductive state, and environmental variables. We also estimated biological process variation in survival and breeding transition probabilities by factoring out sampling variation. The recapture rate of females and calves was high (0.78 ± 0.15 [SE]) and much lower for males (0.41 ± 0.23), especially older males (0.17 ± 0.15). We found that overall bison survival was high (>0.8) and that males (0.80 ± 0.13) survived at lower rates than females (0.94 ± 0.04), but as females aged survival declined (0.89 ± 0.05 for F ≥15 yr old). Lactating and non-lactating females survived at similar rates. We found that females can conceive early (approx. 1.5 yr of age) and had a high probability (approx. 0.8) of breeding in consecutive years, until age 13.5 years, when females that were non-lactating tended to stay in that state. Our results suggest senescence in reproduction and survival for females. We found little support for the effect of climatic covariates on demographic rates, perhaps because the park's current population management goals were predicated from drought-year conditions. This reintroduction has been successful, but continued culling actions will need to be employed and an adaptive management approach is warranted. Our demographic approach can be applied to other heavily managed large-ungulate systems with few or no natural predators.

Alloparental behaviour and long-term costs of mothers tolerating other members of the group in a plurally breeding mammal

Cooperative-breeding studies tend to focus on a few alloparental behaviours in highly cooperative species exhibiting high reproductive skew and the associated short-term, but less frequently long-term, fitness costs. We analysed a suite of alloparental behaviours (assessed via filming) in a kin-structured, high-density population of plurally breeding European badgers, Meles meles, which are not highly cooperative. Group members, other than mothers, performed alloparental behaviour; however, this was not correlated with their relatedness to within-group young. Furthermore, mothers babysat, allogroomed cubs without reciprocation, and allomarked cubs more than other members of the group (controlling for observation time). For welfare reasons, we could not individually mark cubs; however, the number observed pre-independence never exceeded that trapped. All 24 trapped cubs, in three filmed groups, were assigned both parents using 22 microsatellites. Mothers may breed cooperatively, as the time they babysat their assigned, or a larger, litter size did not differ. Furthermore, two mothers probably allonursed, as they suckled more cubs than their assigned litter size. An 18-year genetic pedigree, however, detected no short-term (litter size; maternal survival to the following year) or long-term (offspring breeding probability; offspring lifetime breeding success) fitness benefits with more within-group mothers or other members of the group. Rather, the number of other members of the group (excluding mothers) correlated negatively with long-term fitness. Mothers may tolerate other members of the group, as nonbreeders undertook more digging. Our study highlights that alloparental care varies on a continuum from that seen in this high-density badger population, where alloparenting behaviour is minimal, through to species where alloparental care is common and provides fitness benefits.

Discrete two-sex models of population dynamics: On modelling the mating function

Although sexual reproduction has long been a central subject of theoretical ecology, until recently its consequences for population dynamics were largely overlooked. This is now changing, and many studies have addressed this issue, showing that when the mating system is taken into account, the population dynamics depends on the relative abundance of males and females, and is non-linear. Moreover, sexual reproduction increases the extinction risk, namely due to the Allee effect. Nevertheless, different studies have identified diverse potential consequences, depending on the choice of mating function. In this study, we investigate the consequences of three alternative mating functions that are frequently used in discrete population models: the minimum; the harmonic mean; and the modified harmonic mean. We consider their consequences at three levels: on the probability that females will breed; on the presence and intensity of the Allee effect; and on the extinction risk. When we consider the harmonic mean, the number of times the individuals of the least abundant sex mate exceeds their mating potential, which implies that with variable sex-ratios the potential reproductive rate is no longer under the modeller's control. Consequently, the female breeding probability exceeds 1 whenever the sex-ratio is male-biased, which constitutes an obvious problem. The use of the harmonic mean is thus only justified if we think that this parameter should be re-defined in order to represent the females' breeding rate and the fact that females may reproduce more than once per breeding season. This phenomenon buffers the Allee effect, and reduces the extinction risk. However, when we consider birth-pulse populations, such a phenomenon is implausible because the number of times females can reproduce per birth season is limited. In general, the minimum or modified harmonic mean mating functions seem to be more suitable for assessing the impact of mating systems on population dynamics.

Decreased winter severity increases viability of a montane frog population

Many proximate causes of global amphibian declines have been well documented, but the role that climate change has played and will play in this crisis remains ambiguous for many species. Breeding phenology and disease outbreaks have been associated with warming temperatures, but, to date, few studies have evaluated effects of climate change on individual vital rates and subsequent population dynamics of amphibians. We evaluated relationships among local climate variables, annual survival and fecundity, and population growth rates from a 9-year demographic study of Columbia spotted frogs (Rana luteiventris) in the Bitterroot Mountains of Montana. We documented an increase in survival and breeding probability as severity of winter decreased. Therefore, a warming climate with less severe winters is likely to promote population viability in this montane frog population. More generally, amphibians and other ectotherms inhabiting alpine or boreal habitats at or near their thermal ecological limits may benefit from the milder winters provided by a warming climate as long as suitable habitats remain intact. A more thorough understanding of how climate change is expected to benefit or harm amphibian populations at different latitudes and elevations is essential for determining the best strategies to conserve viable populations and allow for gene flow and shifts in geographic range.

Effects of treatment of anestrous dairy cows with gonadotropin-releasing hormone, prostaglandin, and progesterone

Cows anestrous at the start of a seasonal breeding period have lesser probability of breeding, lesser conception rates, and a longer interval to conception than cycling herdmates. Historically, treatment included estradiol benzoate, which is no longer available. Consequently, alternative programs are required. Hence, a study was undertaken to assess new treatment regimens for these cows. The presence or absence of a corpus luteum was determined using ultrasonography in cows (n=2,222 from 12 herds) that were not detected in estrus by 9 d before the start of breeding. Cows were then randomly assigned to one of 4 treatments within each herd. Treatments were (1) 100 μg of gonadorelin, followed 7 d later by 500 μg of sodium cloprostenol, followed 54 to 56h later by 100 μg of gonadorelin, followed by fixed-time artificial insemination at 13 to 18h after the final GnRH injection (Ovsynch); (2) as for (1) but with placement of an intravaginal progesterone (P4)-releasing insert between the initial GnRH and PGF2α (Ovsynch-56+P4); (3) as for (2) but with the final GnRH treatment delayed until 71h after PGF2α and P4 insert removal with fixed-time artificial insemination 0 to 5h after GnRH treatment and with insemination of those cows detected in estrus before the second GnRH injection (Cosynch-72+P4); and (4) untreated controls (control). Day 0 was defined as the day of the second GnRH injection. Milk samples were collected from 154 and 152 cows from the Ovsynch and Ovsynch-56+P4 treatments, respectively, at d 0, 7, and 14 for P4 concentration determination. This was to test the hypothesis that inclusion of P4 would result in a greater proportion of cows having normal luteal function after treatment in these 2 groups that differed only in the inclusion of P4 in the Ovsynch-56+P4 treatment. All treatments resulted in shorter intervals from first day of breeding to conception compared with the controls. The Ovsynch-56+P4 treatment resulted in start of breeding to conception intervals 3, 6, and 16 d shorter than those of Cosynch-72+P4, Ovsynch, or controls, respectively, and the positive effect of the Ovsynch-56+P4 treatment occurred both in corpus-luteum-positive and in corpus-luteum-negative cows. The Ovsynch-56+P4 treatment resulted in fewer short interestrus intervals than did Ovsynch (i.e., <18 d; 16 vs. 31%) and more cows with elevated (>1 ng/mL) milk P4 concentrations at d 7 (88 vs. 74%) and d 14 (80 vs. 60%). It was concluded that treatment of anestrous cows before the start of breeding resulted in earlier conception than no treatment but had no effect on the final pregnancy rate. The addition of P4 to the Ovsynch program resulted in earlier conception and in more cows with normal subsequent luteal-phase lengths.

Recruitment in a Colorado population of big brown bats: breeding probabilities, litter size, and first-year survival

We used mark–recapture estimation techniques and radiography to test hypotheses about 3 important aspects of recruitment in big brown bats (Eptesicus fuscus) in Fort Collins, Colorado: adult breeding probabilities, litter size, and 1st-year survival of young. We marked 2,968 females with passive integrated transponder (PIT) tags at multiple sites during 2001–2005 and based our assessments on direct recaptures (breeding probabilities) and passive detection with automated PIT tag readers (1st-year survival). We interpreted our data in relation to hypotheses regarding demographic influences of bat age, roost, and effects of years with unusual environmental conditions: extreme drought (2002) and arrival of a West Nile virus epizootic (2003). Conditional breeding probabilities at 6 roosts sampled in 2002–2005 were estimated as 0.64 (95% confidence interval [95% CI] 5 0.53–0.73) in 1-year-old females, but were consistently high (95% CI 5 0.94–0.96) and did not vary by roost, year, or prior year breeding status in older adults. Mean litter size was 1.11 (95% CI 5 1.05–1.17), based on examination of 112 pregnant females by radiography. Litter size was not higher in older or larger females and was similar to results of other studies in western North America despite wide variation in latitude. First-year survival was estimated as 0.67 (95% CI 5 0.61–0.73) for weaned females at 5 maternity roosts over 5 consecutive years, was lower than adult survival (0.79; 95% CI 5 0.77–0.81), and varied by roost. Based on model selection criteria, strong evidence exists for complex roost and year effects on 1st-year survival. Firstyear survival was lowest in bats born during the drought year. Juvenile females that did not return to roosts as 1year-olds had lower body condition indices in late summer of their natal year than those known to survive. DOI: 10.1644/08-MAMM-A-295.1.

Survival and breeding of polar bears in the southern Beaufort Sea in relation to sea ice

1. Observed and predicted declines in Arctic sea ice have raised concerns about marine mammals. In May 2008, the US Fish and Wildlife Service listed polar bears (Ursus maritimus) - one of the most ice-dependent marine mammals - as threatened under the US Endangered Species Act. 2. We evaluated the effects of sea ice conditions on vital rates (survival and breeding probabilities) for polar bears in the southern Beaufort Sea. Although sea ice declines in this and other regions of the polar basin have been among the greatest in the Arctic, to date population-level effects of sea ice loss on polar bears have only been identified in western Hudson Bay, near the southern limit of the species' range. 3. We estimated vital rates using multistate capture-recapture models that classified individuals by sex, age and reproductive category. We used multimodel inference to evaluate a range of statistical models, all of which were structurally based on the polar bear life cycle. We estimated parameters by model averaging, and developed a parametric bootstrap procedure to quantify parameter uncertainty. 4. In the most supported models, polar bear survival declined with an increasing number of days per year that waters over the continental shelf were ice free. In 2001-2003, the ice-free period was relatively short (mean 101 days) and adult female survival was high (0.96-0.99, depending on reproductive state). In 2004 and 2005, the ice-free period was longer (mean 135 days) and adult female survival was low (0.73-0.79, depending on reproductive state). Breeding rates and cub litter survival also declined with increasing duration of the ice-free period. Confidence intervals on vital rate estimates were wide. 5. The effects of sea ice loss on polar bears in the southern Beaufort Sea may apply to polar bear populations in other portions of the polar basin that have similar sea ice dynamics and have experienced similar, or more severe, sea ice declines. Our findings therefore are relevant to the extinction risk facing approximately one-third of the world's polar bears.

Influence of climate and reproductive timing on demography of little brown myotis Myotis lucifugus

1. Estimating variation in demographic rates, such as survival and fecundity, is important for testing life-history theory and identifying conservation and management goals. 2. We used 16 years (1993-2008) of mark-recapture data to estimate age-specific survival and breeding probabilities of the little brown myotis Myotis lucifugus LeConte in southern New Hampshire, USA. Using Kendall & Nichols' (1995) full-likelihood approach of the robust design to account for temporary emigration, we tested whether survival and breeding propensity is influenced by regional weather patterns and timing of reproduction. 3. Our results demonstrate that adult female survival of M. lucifugus ranged from 0.63 (95% CL = 0.56, 0.68) to 0.90 (95% CL = 0.77, 0.94), and was highest in wet years with high cumulative summer precipitation. First-year survival [range: 0.23 (95% CL = 0.14, 0.35) to 0.46 (95% CL = 0.34, 0.57)] was considerably lower than adult survival and depended on pup date of birth, such that young born earlier in the summer (c. late May) had a significantly higher probability of surviving their first year than young born later in the summer (c. mid-July). Similarly, the probability of young females returning to the maternity colony to breed in the summer following their birth year was higher for individuals born earlier in the summer [range: 0.23 (95% CL = 0.08, 0.50) to 0.53 (95% CL = 0.30, 0.75)]. 4. The positive influence of early parturition on 1st-year survival and breeding propensity demonstrates significant fitness benefits to reproductive timing in this temperate insectivorous bat. 5. Climatic factors can have important consequences for population dynamics of temperate bats, which may be negatively affected by summer drying patterns associated with global climate change. 6. Understanding long-term demographic trends will be important in the face of a novel disease phenomenon (White-Nose Syndrome) that is associated with massive mortalities in hibernating bat species, including M. lucifugus, in the northeastern United States.

Maximizing re-introduction success: Assessing the immediate cost of release in a threatened waterfowl

Translocations have become one of the most commonly used tools for biodiversity restoration worldwide, however one out of three re-introduction plans fails to create a viable population or to successfully reinforce the existing one. We used results from the analysis of individual-based information on the re-introduction of a threatened waterfowl species, the crested coots Fulica cristata, to provide guidelines to maximise re-introduction success. We found that about a third of the post-release mortality took place within the first month after release. This immediate ‘cost of release’ in terms of local survival or ‘release risk factor’ seems to be a common feature of re-introduction projects, and it is likely due to the inexperience of captive-born individuals to face the new environment. This hypothesis was supported by the positive association between survival and time spent in the wild Results suggested that coots released between February and May have a slightly higher survival. A joint measure of survival and breeding probabilities indicated that birds released in late winter (February-March) had a higher chance to survive and reproduce compared to birds released later in the year. From an applied perspective our results can be used within an adaptive management framework to determine the optimum period of release, providing substantial support for future decision-making in the management of waterfowl, and other long-term projects of re-introduction of threatened vertebrate species.

Lesser Scaup Breeding Probability and Female Survival on the Yukon Flats, Alaska

ABSTRACT Information on the ecology of waterfowl breeding in the boreal forest is lacking, despite the boreal region's importance to continental waterfowl populations and to duck species that are currently declining, such as lesser scaup (Aythya affinis). We estimated breeding probability and breeding season survival of female lesser scaup on the Yukon Flats National Wildlife Refuge, Alaska, USA, in 2005 and 2006. We captured and marked 93 lesser scaup with radiotransmitters during prelaying and nesting periods. Although all marked lesser scaup females were paired throughout prelaying and incubation periods, we estimated breeding probability over both years as 0.12 (SE = 0.05, n = 67) using telemetry. Proportion of lesser scaup females undergoing rapid follicle growth at capture in 2006 was 0.46 (SE = 0.11, n = 37), based on concentration of yolk precursors in blood plasma. By combining methods based on telemetry, yolk precursors, and postovulatory follicles, we estimated maximum breeding probability as 0.68 (SE = 0.08, n = 37) in 2006. Notably, breeding probability was positively related to female body mass. Survival of female lesser scaup during the nesting and brood‐rearing periods was 0.92 (SE = 0.05) in 2005 and 0.86 (SE = 0.08) in 2006. Our results suggest that breeding probability is lower than expected for lesser scaup. In addition, the implicit assumption of continental duck‐monitoring programs that all paired females attempt to breed should be reevaluated. Recruitment estimates based on annual breeding‐pair surveys may overestimate productivity of scaup pairs in the boreal forest.

Survival and Breeding Frequency in Marbled Salamanders (Ambystoma opacum): Implications for Spatio-temporal Population Dynamics

Despite known population-level sensitivity to adult vital rates, there is a shortage of robust estimates of adult survival and breeding frequency for pond-breeding amphibians. Evaluating the applicability of metapopulation principles to amphibians has also been constrained by the lack of demographic studies scaled beyond individual breeding populations. We investigate spatial and temporal demographic variability among six proximate breeding populations of Marbled Salamanders (Ambystoma opacum) in western Massachusetts, USA, focusing on the terrestrial adult life stage. Between 1999 and 2004, we captured and photographed approximately 1700 breeding adults, each between one and 12 times, at drift fences encircling breeding sites. After constructing individual capture histories from field data, we applied maximum likelihood approaches to estimate parameters for survival in the pond basins during breeding periods, survival in upland habitat during breeding and non-breeding periods, and both breeding and capture probabilities. Models selected using Akaike's Information Criteria suggested that there was moderate variability in pond survival across years, but that generally, pond and upland survival varied little among populations within years. This suggests that asynchronous variability indicative of metapopulation dynamics is unlikely to be significant in the adult stage, at least among nearby populations in similar upland forests. An integrated measure of annual survival was 0.66 (SE = 0.02) for males and 0.62 (SE = 0.01) for females. Average breeding probabilities were very high (0.96; SE = 0.01) for males and high (0.67; SE = 0.02) for females, resulting in estimates of 2.5 and 2.0 lifetime breeding attempts, respectively. These parameter estimates will be used to develop spatially explicit population models to guide conservation and forest management strategies for A. opacum and other pond-breeding amphibians with similar life histories. Additional empirical work that estimates the effects of alternative management strategies on these vital rates will greatly improve the utility of these models.

Costs of reproduction in a long-lived bird: large clutch size is associated with low survival in the presence of a highly virulent disease

Fitness costs of reproduction are expected to be more pronounced when the environmental conditions deteriorate. We took advantage of a natural experiment to investigate the costs of reproduction among common eiders (Somateria mollissima) nesting at a site in the Arctic, where an avian cholera epizootic appeared at different magnitudes. We tested the predictions that larger reproductive effort (clutch size) is associated with lower survival or breeding probability the following year, and that this relationship was more pronounced under heightened exposure to the disease. Our results indicate that large clutch sizes were associated with lower survival of female eider ducks, but only when there was heightened exposure to avian cholera, as indexed by eider mortality on site. No cost was observed when cholera was absent or when lesser exposure was evident. This supports the hypothesis that fitness costs of high reproductive effort are higher under unfavourable conditions such as a disease epizootic, and further indicates that being a conservative breeder can increase survival probability, given the presence of a highly virulent disease.

Retrospective investigation of captive red wolf reproductive success in relation to age and inbreeding

The critically endangered red wolf (Canis rufus) has been subject to a strictly managed captive breeding program for three decades. A retrospective demographic analysis of the captive population was performed based on data from the red wolf studbook. Data analyses revealed a decrease in the effective population size relative to the total population size, and changes in age structure and inbreeding coefficients over time. To varying degrees, the probability of successful breeding and litter sizes declined in association with increasing dam age and sire inbreeding coefficients. Neonate survival also declined with increasing dam age. Recent changes in strategies regarding breed-pair recommendations have resulted in moderate increases in reproductive success.

Body Mass, Territory Size, and Life-History Tactics in a Socially Monogamous Canid, the Red FoxVulpes vulpes

Male-biased sexual size dimorphism is typical of polygynous mammals, where the degree of dimorphism in body mass is related to male intrasexual competition and the degree of polygyny. However, the importance of body mass in monogamous mammals is largely unknown. We investigated the effect of body mass on life-history parameters and territory size in the red fox (Vulpes vulpes), a socially monogamous canid with slight sexual dimorphism. Increased body size in males appeared to confer an advantage in territory acquisition and defense contests because heavier males held larger territories and exerted a greater boundary pressure on smaller neighbors. Heavier male foxes invested more effort in searching for extrapair matings by moving over a wider area and farther from their territories, leading to greater reproductive success. Males that sired cubs outside their own social group appeared to be heavier than males that only sired cubs within their social group or that were cuckolded, but our results should be treated with caution because sample sizes were small. Territory size, boundary pressure, and paternity success were not related to age of males. In comparison, body mass of females was not related to territory size, probability of breeding, litter size, or cub mass. Only age affected probability of breeding in females: younger females reproduced significantly less than did older females, although we did not measure individual nutritional status. Thus, body mass had a significant effect on life-history traits and territory size in a socially monogamous species comparable to that reported in polygynous males, even in the absence of large size dimorphism.

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine, the tree swallowThachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallowTachycineta bicolorusing 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.

Spatial variation in age‐specific probabilities of first reproduction for Weddell seals

Spatial variation in vital rates can affect the dynamics and persistence of a population. We evaluated the prediction that age‐specific probabilities of survival and first reproduction for Weddell seals would vary as a function of birth location in Erebus Bay, Antarctica. We used multi‐state mark–resight models and 25 years of data to estimate demographic rates for female seals. We predicted that probabilities of survival and first reproduction would be higher for seals born at near‐shore colonies or more southerly‐located colonies with consistent ice conditions. Contrary to predictions, results revealed higher age‐specific probabilities of first reproduction at offshore colonies relative to near‐shore colonies and no spatial variation in survival rates. For 7‐year old females (average age at 1st reproduction=7.6 years old) born at offshore colonies to mothers aged 10.8 years (average maternal age), probability of first reproduction was 0.43 (SE=0.07), whereas probability of first reproduction for females born at near‐shore colonies was 0.30 (SE=0.05) based on estimates from our top‐ranked model. Breeding probabilities following first reproduction were also higher at offshore colonies. Thus, our results (1) provide evidence of spatial variation in breeding probabilities, (2) reveal the importance of birth location on a female's vital rates, and (3) suggest that the effect persisted for many years. Birth‐colony effects may be attributed to spatial variation in prey availability, or to heterogeneity in female quality in this population. If females who are superior competitors consistently chose offshore colonies for pupping, pups born at these locations may have inherited those superior qualities and displayed higher probabilities of first reproduction, relative to seals born at other colonies. Further research into physical or food‐related differences among colonies may offer insight into spatial variation in breeding probabilities documented in this paper.

Fidelity and breeding probability related to population density and individual quality in black brent geese Branta bernicla nigricans

1. Patterns of temporary emigration (associated with non-breeding) are important components of variation in individual quality. Permanent emigration from the natal area has important implications for both individual fitness and local population dynamics. 2. We estimated both permanent and temporary emigration of black brent geese (Branta bernicla nigricans Lawrence) from the Tutakoke River colony, using observations of marked brent geese on breeding and wintering areas, and recoveries of ringed individuals by hunters. We used the likelihood developed by Lindberg, Kendall, Hines & Anderson 2001 (Combining band recovery data and Pollock's robust design to model temporary and permanent emigration. Biometrics, 57, 273-281) to assess hypotheses and estimate parameters. 3. Temporary emigration (the converse of breeding) varied among age classes up to age 5, and differed between individuals that bred in the previous years vs. those that did not. Consistent with the hypothesis of variation in individual quality, individuals with a higher probability of breeding in one year also had a higher probability of breeding the next year. 4. Natal fidelity of females ranged from 0.70 +/- 0.07-0.96 +/- 0.18 and averaged 0.83. In contrast to Lindberg et al. (1998), we did not detect a relationship between fidelity and local population density. Natal fidelity was negatively correlated with first-year survival, suggesting that competition among individuals of the same age for breeding territories influenced dispersal. Once females nested at the Tutakoke River, colony breeding fidelity was 1.0. 5. Our analyses show substantial variation in individual quality associated with fitness, which other analyses suggest is strongly influenced by early environment. Our analyses also suggest substantial interchange among breeding colonies of brent geese, as first shown by Lindberg et al. (1998).

Recruitment processes in long‐lived species with delayed maturity: estimating key demographic parameters

Long‐lived species show delayed maturity and generally skip breeding sites on a given year, causing difficulties in estimating demographic parameters. A novel multi‐state capture – recapture model (model G for general) is proposed to estimate survival and recruitment. Model G considers long and short periods of non‐attendance at the breeding site. Model G is compared against a reduced model (model R) to test if pre‐breeders skip the studied site a given year, potentially prospecting other breeding habitats.This model is applied to the vulnerable Cory's shearwaters seabird species. As for many species, natal philopatry, pre‐breeding attendance and prospecting play an important role for recruitment into the breeding population. Model G shows that all Cory's shearwaters come back to their natal colony, and attend the colony before recruiting into the breeding population. Moreover, model G is preferred against model R, highlighting that prospecting plays a potentially role in the recruitment processes. Return and recruitment probabilities show similar time variation, suggesting an environmental forcing. The maximum probability of the first breeding is reached at the age of seven years. During the first year the annual survival rate is 0.52 (IC95%=[0.40, 0.64]) and pre‐breeders and adult annual survival rates do not differ, being respectively 0.88, IC95%=[0.83, 0.92] and 0.89, IC95%=[0.88, 0.9].Model G provides an improvement to estimate demographic parameters for long‐lived species life cycle and an adequate framework to test the influence of covariates. Model G is structured by age and breeding status categories, allowing easy further population models.

Survival and Breeding Advantages of Larger Black Brant (Branta Bernicla Nigricans) Goslings: Within- and Among-Cohort Variation

AbstractWe examined the relationship between mass late in the first summer and survival and return to the natal breeding colony for 12 cohorts (1986-1997) of female Black Brant (Branta bernicla nigricans). We used Cormack-Jolly-Seber methods and the program MARK to analyze capture-recapture data. Models included two kinds of residuals from regressions of mass on days after peak of hatch when goslings were measured; one based on the entire sample (12 cohorts) and the other based only on individuals in the same cohort. Some models contained date of peak of hatch (a group covariate related to lateness of nesting in that year) and mean cohort residual mass. Finally, models allowed survival to vary among cohorts. The best model of encounter probability included an effect of residual mass on encounter probability and allowed encounter probability to vary among age classes and across years. All competitive models contained an effect of one of the estimates of residual mass; relatively larger goslings survived their first year at higher rates. Goslings in cohorts from later years in the analysis tended to have lower first-year survival, after controlling for residual mass, which reflected the generally smaller mean masses for these cohorts but was potentially also a result of population-density effects additional to those on growth. Variation among cohorts in mean mass accounted for 56% of variation among cohorts in first-year survival. Encounter probabilities, which were correlated with breeding probability, increased with relative mass, which suggests that larger goslings not only survived at higher rates but also bred at higher rates. Although our findings support the well-established linkage between gosling mass and fitness, they suggest that additional environmental factors also influence first-year survival.Supervivencia y Ventajas Reproductivas de los Pichones de Mayor Tamaño de Branta bernicla nigricans: Variación entre y dentro de las Cohortes

THE COST OF REPRODUCTION INDUCED BY BODY SIZE AT BIRTH AND BREEDING DENSITY

Body size at birth has implications for the quality of individuals throughout their life. Although large body size is generally considered an advantage, the relationship between body size at birth and long-term fitness is often complicated. Under spatial or temporal variation in environmental conditions, such as the seasonally changing densities of Fennoscandian vole populations, selection should favor variation in offspring phenotypes, as different qualities may be beneficial in different conditions. We performed an experiment in which a novel hormonal manipulation method was used to increase phenotypic variance in body size at birth in the bank vole (Myodes glareolus). The effects of body size on the future fitness of young males and females were then studied at varying population densities in outdoor enclosures. Our results show that small body size at birth and high breeding density increase the survival costs of reproduction. However, there was no interaction between the effects of body size and density on survival, which suggests that the fitness effects of body size were strong enough to persist under environmental variation. Moreover, litter size and the probability of breeding were more sensitive to variation in breeding density than offspring size. Therefore, it is unlikely that individual fitness could be optimized by adjusting offspring body size to the prevailing population density through adaptive maternal effects. Our results highlight the significance of the costs of reproduction in the evolution of life-history traits, and give strong experimental support for the long-term fitness effects of body size at birth.