Bottom‐up processes in a declining yellow‐footed rock‐wallaby (Petrogale xanthopus celeris) population

Populations of large herbivores are generally considered to be food limited, escaping the regulatory effects of predation through their large body size, migratory behaviour and/or the occurrence of alternate prey species. In the Australian arid and semi‐arid zones, the availability of forage biomass is considered to be the primary driver of fluctuations in kangaroo abundance. However, little is known about the population dynamics of the smaller sympatric macropods. We examined the demographic traits of a large colony of yellow‐footed rock‐wallabies (Petrogale xanthopus celeris), following a 2‐year period of above average rainfall. The population was located within a conservation reserve that was subject to a predator control program around its perimeter and on neighbouring properties. The low predator abundance provided an opportunity to gauge the strength of bottom‐up population processes. During the two years of the study, the population declined in size by 53%, resulting from both the virtual absence of juvenile recruitment and the loss of adult wallabies. Although reproductive output was high, low pouch young and juvenile survival rates resulted in few individuals progressing into the adult population. With minimal recruitment, the rate of population decline (r = 0.77) matched the observed adult survival rate (Φ = 0.76). Despite average rainfall conditions during the study, survival rates across all age‐classes were equivalent to those reported for other rock‐wallaby populations during periods of scarcity. The reduced survival rates were attributed to low levels of forage resources, particularly around the wallabies' refuge sites, suggesting the bottom‐up regulation of the colony at high densities. The data suggest that the colony was at temporarily high abundance, following a rainfall driven pulse of recruitment. Conservation management actions for this species should focus on increasing juvenile survival rates within declining populations, through the control of feral goats (Capra hircus), rabbits (Oryctolagus cuniculus) and red foxes (Vulpes vulpes).     

Survival estimates for the Australian sea lion: Negative correlation of sea surface temperature with cohort survival to weaning

The Australian sea lion (Neophoca cinerea) population at Seal Bay Conservation Park, South Australia, is estimated to be declining at a rate of 1.14% per breeding season. To better understand the potential causes of this decline, survival rates were examined to 14 yr of age for eight cohorts marked as pups (aged 0.17 yr) between 1991 and 2002. Apparent yearly survival rates (Φ) varied by cohort for pups from marking to weaning at 1.5 yr (Φ= 0.30–0.67). Postweaning juvenile survival (1.5–3 yr) was 0.89 and survival from 3 to 14 yr was constant (Φ female:male = 0.96:0.89). Φ of pup cohorts was negatively correlated to local sea surface temperature where the sea lions forage (SST) and was especially low for cohort 7 in 2000 (0.30). It is possible that periods of unusually warm oceanographic conditions may be limiting primary production and inhibiting maternal provisioning to pups. Pup survival to weaning is relatively low compared to other otariid species, is likely to limit recruitment, and may be contributing to the decline in pup abundance observed in the colony.     

Slow life history and rapid extreme flood: demographic mechanisms and their consequences on population viability in a threatened amphibian

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The population increase of common guillemots Uria aalge on Skomer Island is explained by intrinsic demographic properties

The population of common guillemots Uria aalge on Skomer Island, Wales has been monitored since 1963, and in the last 30 yr has increased at an almost constant rate of 5% yr^?1. A previous attempt to model the population based on intrinsic demographic parameters estimated over just five years failed to explain the observed population increase, probably because the estimate of juvenile survival was too low. This raised the possibility that immigration fuelled the population increase. Here we use > 30 yr of detailed field observations to re‐estimate key population parameters (productivity, adult survival and juvenile survival) in order to model the population. We show that the observed rate of increase can be explained by these intrinsic parameters, and we therefore conclude that immigration is not necessary to generate the observed population growth.     

Offspring size at weaning affects survival to recruitment and reproductive performance of primiparous gray seals

Offspring size affects survival and subsequent reproduction in many organisms. However, studies of offspring size in large mammals are often limited to effects on juveniles because of the difficulty of following individuals to maturity. We used data from a long‐term study of individually marked gray seals (Halichoerus grypus; Fabricius, 1791) to test the hypothesis that larger offspring have higher survival to recruitment and are larger and more successful primiparous mothers than smaller offspring. Between 1998 and 2002, 1182 newly weaned female pups were branded with unique permanent marks on Sable Island, Canada. Each year through 2012, all branded females returning to the breeding colony were identified in weekly censuses and a subset were captured and measured. Females that survived were significantly longer offspring than those not sighted, indicating size‐selective mortality between weaning and recruitment. The probability of female survival to recruitment varied among cohorts and increased nonlinearly with body mass at weaning. Beyond 51.5 kg (mean population weaning mass) weaning mass did not influence the probability of survival. The probability of female survival to recruitment increased monotonically with body length at weaning. Body length at primiparity was positively related to her body length and mass at weaning. Three‐day postpartum mass (proxy for birth mass) of firstborn pups was also positively related to body length of females when they were weaned. However, females that were longer or heavier when they were weaned did not wean heavier firstborn offspring.     

Postharvesting population dynamics of the South American sea lion (Otaria byronia) in the southwestern Atlantic

Many pinniped populations precipitously declined during the 19th and 20th centuries due to overharvesting. In Uruguay, the South American sea lion (SASL) was harvested until 1986. Birth rates in two nearby breeding colonies have had opposite trends for at least 20 yr. We assessed different mechanisms that could explain opposite trends in birth rates in the two SASL colonies. We compared feeding habits (δ¹⁵N and δ¹³C) of breeding females, birth mass, individual growth rate and early survival of pups and the social structure between colonies. Breeding females from the two colonies did not differ in their feeding habits. However, male and female pups grew faster but had a lower survival in the second month in the smallest colony. We found differences in the social structures, with a higher proportion of males in the smallest colony. The latter is important because peripheral SASL males may abduct and kill pups, which may explain the lower survival of pups in smaller colonies. We believe that the cumulative effects of population extractions have lowered the local SASL population size and disrupted its social structure to the point where Allee‐like effects could become important and hamper the recovery of the Uruguayan SASL population.     

Long‐term population dynamics reveal that survival and recruitment of tropical boobies improve after a hurricane

Variability in population numbers is a central issue in evolutionary ecology and also in biodiversity conservation. However, for most seabirds this information is lacking and tropical populations are virtually unstudied. Long‐term studies are warranted because world's seabird populations exhibit an overall declining trend since 1950. Using data spanning 23 yr, we investigated how adult survival, local recruitment, and their relative contributions to population growth (λ) vary over time in the blue‐footed booby Sula nebouxii, a long‐lived locally foraging seabird that breeds in tropical waters. In addition, we investigated whether booby demographic rates exhibit the same declining trend observed in other seabirds, whether these rates are impacted by hurricanes, and whether these potential impacts differ between sexes. Our analysis of 4608 capture–recapture histories revealed that survival and recruitment were nearly equal between males and females, exhibited a declining trend over the last 23 yr, and in both sexes, these vital rates improved after a hurricane. The declining trend in recruitment was slightly more attenuated in males. These results add to the current evidence for an overall declining trend in world's seabird populations and extend its confirmation to the warm eastern tropical Pacific. Moreover, they provide the first evidence that hurricanes may favor natural populations. As a result of the declining trend and variation in survival and recruitment, λ exhibited a slight decline and substantial variation over the 23 yr. However, most λ values were equal to or higher than 1, and the long‐term average indicates population increase. The ability of blue‐footed boobies to maintain a positive population balance despite of negative trends in their vital rates might result from canalization of adult survival (the vital rate that contributes most to λ and shows lower variation compared to recruitment) against environmental variability.     

Lifetime survival rates and senescence in northern elephant seals

The aim of this study was to extend 40 yr of prior demographic work on northern elephant seals (Mirounga angustirostris) at Año Nuevo, California, by including the oldest animals. We used a Bayesian mark‐recapture analysis to estimate lifelong survival and lifespan of a cohort of 372 weaned pups branded in 1985–1987 and resighted until 2008. Annual survival probability of females averaged 86.3%/yr at ages 5–16, then declined until age 21, the age of the oldest female. Male survival was lower, averaging 67.7%/yr from age 1 to age 15, the age of the oldest male. Northern elephant seal females in the expanding population at Año Nuevo live longer than southern elephant seal females (M. leonina) at colonies whose populations are declining. This comparison suggests that high survival of females is a key factor in population growth.     

Variations in age‐ and sex‐specific survival rates help explain population trend in a discrete marine mammal population

Understanding the drivers underlying fluctuations in the size of animal populations is central to ecology, conservation biology, and wildlife management. Reliable estimates of survival probabilities are key to population viability assessments, and patterns of variation in survival can help inferring the causal factors behind detected changes in population size. We investigated whether variation in age‐ and sex‐specific survival probabilities could help explain the increasing trend in population size detected in a small, discrete population of bottlenose dolphins Tursiops truncatus off the east coast of Scotland. To estimate annual survival probabilities, we applied capture–recapture models to photoidentification data collected from 1989 to 2015. We used robust design models accounting for temporary emigration to estimate juvenile and adult survival, multistate models to estimate sex‐specific survival, and age models to estimate calf survival. We found strong support for an increase in juvenile/adult annual survival from 93.1% to 96.0% over the study period, most likely caused by a change in juvenile survival. Examination of sex‐specific variation showed weaker support for this trend being a result of increasing female survival, which was overall higher than for males and animals of unknown sex. Calf survival was lower in the first than second year; a bias in estimating third‐year survival will likely exist in similar studies. There was some support first‐born calf survival being lower than for calves born subsequently. Coastal marine mammal populations are subject to the impacts of environmental change, increasing anthropogenic disturbance and the effects of management measures. Survival estimates are essential to improve our understanding of population dynamics and help predict how future pressures may impact populations, but obtaining robust information on the life history of long‐lived species is challenging. Our study illustrates how knowledge of survival can be increased by applying a robust analytical framework to photoidentification data.     

Influence of maternal characteristics and oceanographic conditions on survival and recruitment probabilities of Weddell seals

For long‐lived animals, maternal age and breeding experience can vary widely and affect offspring survival and recruitment probabilities. In addition, these vital rates may be influenced by annual variation in environmental conditions. We evaluated various hypotheses regarding how offspring survival and recruitment probabilities vary as functions of maternal characteristics and oceanographic conditions, using 25 years of data from a study of individually‐marked Weddell seals in Erebus Bay, Antarctica. We predicted that survival and recruitment would be positively related to maternal age and experience up to some threshold value and considered three hypothesized shapes for the relationship beyond the threshold age (steadily increasing, pseudo‐threshold, or decreasing). We predicted an inverse relationship between maternal age at first reproduction and offspring survival and recruitment probabilities. We predicted that sea‐ice extent, which positively influences primary productivity, would be positively related to annual recruitment probabilities. Results revealed contrasting influences of maternal age on probabilities of survival and recruitment of young. Survival rate was best modeled by a pseudo‐threshold relationship with maternal age, e.g. in 1999, survival rate was estimated as 0.61, 0.69 and 0.72, respectively, for seals born to 6‐, 14‐ and 22‐yr‐old mothers. In contrast, estimated recruitment probability was highest for seals born to young mothers, e.g. recruitment probability for a 7‐yr‐old who had not yet had a pup was estimated as 0.51 vs 0.30, respectively, if she was born to a 6‐ versus a 14‐yr‐old mother. The combined results for offspring survival and recruitment suggest countervailing selection where genotypes favored for reproductive success are generally selected against as juveniles, resulting in high recruitment probabilities for individuals that had low juvenile survival rates. Finally, we found support for our prediction that oceanographic conditions affected annual recruitment rates, but not survival rates. Specifically, annual recruitment probability was positively related to the sea‐ice extent in September of the previous year.     

Inferring causal factors for a declining population of bottlenose dolphins via temporal symmetry capture–recapture modeling

We applied temporal symmetry capture–recapture (TSCR) models to assess the strength of evidence for factors potentially responsible for population decline in bottlenose dolphins (Tursiops truncatus) in Doubtful Sound, New Zealand from 1995 to 2008. Model selection was conducted to estimate recruitment and population growth rates. There were similar levels of support for three different models, each reflecting distinct trends in recruitment. Modeling yielded low overall estimates of recruitment (0.0249, 95% CI: 0.0174–0.0324) and population growth rate (0.9642, 95% CI: 0.9546–0.9737). The TSCR rate of population decline was consistent with an estimate derived from trends in abundance (lambda = 0.9632, 95% CI: 0.9599–0.9665). The TSCR model selection confirmed the influence of a decline in the survival of calves (<1 yr old) since 2002 for population trends. However, TSCR population growth rates did not exceed 1 in any year between 1995 and 2008, indicating the population was declining prior to 2002. A separate reduction in juvenile survival (1–3 yr old) prior to 2002 was identified as a likely contributing factor in the population decline. Thus, TSCR modeling indicated the potential cause of the population decline in Doubtful Sound: cumulative impacts on individuals <3 yr old resulting in a reduced recruitment.     

Demographic buffering and compensatory recruitment promotes the persistence of disease in a wildlife population

Demographic buffering allows populations to persist by compensating for fluctuations in vital rates, including disease‐induced mortality. Using long‐term data on a badger (Meles meles Linnaeus, 1758) population naturally infected with Mycobacterium bovis, we built an integrated population model to quantify impacts of disease, density and environmental drivers on survival and recruitment. Badgers exhibit a slow life‐history strategy, having high rates of adult survival with low variance, and low but variable rates of recruitment. Recruitment exhibited strong negative density‐dependence, but was not influenced by disease, while adult survival was density independent but declined with increasing prevalence of diseased individuals. Given that reproductive success is not depressed by disease prevalence, density‐dependent recruitment of cubs is likely to compensate for disease‐induced mortality. This combination of slow life history and compensatory recruitment promotes the persistence of a naturally infected badger population and helps to explain the badger's role as a persistent reservoir of M. bovis.     

Comparing survey and multiple recruitment–mortality models to assess growth rates and population projections

Estimation of population trends and demographic parameters is important to our understanding of fundamental ecology and species management, yet these data are often difficult to obtain without the use of data from population surveys or marking animals. The northeastern Minnesota moose (Alces alces Linnaeus, 1758) population declined 58% during 2006–2017, yet aerial surveys indicated stability during 2012–2017. In response to the decline, the Minnesota Department of Natural Resources (MNDNR) initiated studies of adult and calf survival to better understand cause‐specific mortality, calf recruitment, and factors influencing the population trajectory. We estimated population growth rate (λ) using adult survival and calf recruitment data from demographic studies and the recruitment–mortality (R‐M) Equation and compared these estimates to those calculated using data from aerial surveys. We then projected population dynamics 50 years using each resulting λ and used a stochastic model to project population dynamics 30 years using data from the MNDNR's studies. Calculations of λ derived from 2012 to 2017 survey data, and the R‐M Equation indicated growth (1.02 ± 0.16 [SE] and 1.01 ± 0.04, respectively). However, the stochastic model indicated a decline in the population over 30 years (λ = 0.91 ± 0.004; 2014–2044). The R‐M Equation has utility for estimating λ, and the supporting information from demographic collaring studies also helps to better address management questions. Furthermore, estimates of λ calculated using collaring data were more certain and reflective of current conditions. Long‐term monitoring using collars would better inform population performance predictions and demographic responses to environmental variability.     

Forest Fragmentation Alters the Population Dynamics of a Late‐successional Tropical Tree

Tropical late‐successional tree species are at high risk of local extinction due to habitat loss and fragmentation. Population‐growth rates in fragmented populations are predicted to decline as a result of reduced fecundity, survival and growth. We examined the demographic effects of habitat fragmentation by comparing the population dynamics of the late‐successional tree Poulsenia armata (Moraceae) in southern Mexico between a continuous forest and several forest fragments using integral projection models (IPMs) during 2010–2012. Forest fragmentation did not lead to differences in population density and even resulted in a higher population‐growth rate (λ) in fragments compared to continuous forests. Habitat fragmentation had drastic effects on the dynamics of P. armata, causing the population structure to shift toward smaller sizes. Fragmented populations experienced a significant decrease in juvenile survival and growth compared to unaltered populations. Adult survival and growth made the greatest relative contributions to λ in both habitat types during 2011–2012. However, the relative importance of juvenile survival and growth to λ was highest in the fragmented forest in 2010–2011. Our Life Table Response Experiment analysis revealed that positive contributions of adult fecundity explained most of the variation of λ between both habitats and annual periods. Finally, P. armata has a relatively slow speed of recovery after disturbances, compromising persistence of fragmented populations. Developing a mechanistic understanding of how forest fragmentation affects plant population dynamics, as done here, will prove essential for the preservation of natural areas.     

Trend changes in sympatric Subantarctic and Antarctic fur seal pup populations at Marion Island,Southern Ocean

Recent pup population estimates of sympatric Subantarctic (Arctocephalus tropicalis) and Antarctic fur seals (A. gazella) at Marion Island are presented. Published pup population estimates of A. tropicalis (1995 and 2004) with an unpublished total island count in 2013, and annual counts on subsets of rookeries (2007–2015) were analyzed using a hierarchical Bayesian model. The pup population declined by 46% (95% credible interval CI: 43%–48%) between 2004 (mean = 15,260, CI: 14,447–16,169 pups) and 2013 (mean = 8,312, CI: 7,983–8,697), mirrored by a 58%–60% decline at rookeries counted annually (2007–2015). Population decline was highest at high‐density west and north coast rookeries, despite negligible change in female attendance patterns, pup mortality or median pupping date over the previous 25 yr. A better understanding of foraging behavior and its effects on reproductive success and survival in this A. tropicalis population is needed before we can attribute population decline to any external factors. In contrast, total island counts of A. gazella pups in 2007, 2010, and 2013, suggest that this population is still increasing although the annual intrinsic rate of population growth decreased from 17.0% (1995–2004, 744 pups) to 4.0% (2010–2013, 1,553 pups). The slowed growth of A. gazella is likely the result of saturation at the main rookery.     

Demography of the Pacific walrus (Odobenus rosmarus divergens) in a changing Arctic

The Pacific walrus (Odobenus rosmarus divergens) is a candidate to be listed as an endangered species under United States law, in part, because of climate change‐related concerns. While the population was known to be declining in the 1980s and 1990s, its recent status has not been determined. We developed Bayesian models of walrus population dynamics to assess the population by synthesizing information on population sizes, age structures, reproductive rates, and harvests for 1974–2015. Candidate models allowed for temporal variation in some or all vital rates, as well as density dependence or density independence in reproduction and calf survival. All selected models indicated that the population underwent a multidecade decline, which began moderating in the 1990s, and that annual reproductive rate and natural calf survival rates rose over time in a density‐dependent manner. However, selected models were equivocal regarding whether the natural juvenile survival rate was constant or decreasing over time. Depending on whether juvenile survival decreased after 1998, the population growth rate either increased during 1999–2015 or stabilized at a lesser level of decline than seen in the 1980s. The probability that the population was still declining in 2015 ranged from 45% to 87%.     

Are whooping cranes destined for extinction? Climate change imperils recruitment and population growth

Identifying climatic drivers of an animal population's vital rates and locating where they operate steers conservation efforts to optimize species recovery. The population growth of endangered whooping cranes (Grus americana) hinges on juvenile recruitment. Therefore, we identify climatic drivers (solar activity [sunspots] and weather) of whooping crane recruitment throughout the species’ life cycle (breeding, migration, wintering). Our method uses a repeated cross‐validated absolute shrinkage and selection operator approach to identify drivers of recruitment. We model effects of climate change on those drivers to predict whooping crane population growth given alternative scenarios of climate change and solar activity. Years with fewer sunspots indicated greater recruitment. Increased precipitation during autumn migration signified less recruitment. On the breeding grounds, fewer days below freezing during winter and more precipitation during breeding suggested less recruitment. We predicted whooping crane recruitment and population growth may fall below long‐term averages during all solar cycles when atmospheric CO₂ concentration increases, as expected, to 500 ppm by 2050. Species recovery during a typical solar cycle with 500 ppm may require eight times longer than conditions without climate change and the chance of population decline increases to 31%. Although this whooping crane population is growing and may appear secure, long‐term threats imposed by climate change and increased solar activity may jeopardize its persistence. Weather on the breeding grounds likely affects recruitment through hydrological processes and predation risk, whereas precipitation during autumn migration may influence juvenile mortality. Mitigating threats or abating climate change should occur within ≈30 years or this wild population of whooping cranes may begin declining.     

Harbor seal use of glacier ice and terrestrial haul‐outs in the Kenai Fjords,Alaska

Harbor seals, Phoca vitulina, use diverse haul‐out substrates including ice calved by tidewater glaciers. Numbers of seals at glacial and terrestrial haul‐outs on the southeastern Kenai Peninsula, Alaska, were assessed using aerial, vessel, and video surveys. Mean annual abundance at glacial and terrestrial haul‐outs differed temporally. From 2004 to 2011, numbers of seals counted during the molt increased 5.4%/yr at glacial haul‐outs and 9%/yr at terrestrial haul‐outs while numbers of pups increased 5.0%/yr at glacial sites and 1.5%/yr at terrestrial sites. Numbers of seals without pups counted during pupping increased 7.96%/yr at glacial sites and 5.1%/yr at terrestrial sites. Results indicate that pupping and molting locations are not equivalent and population monitoring during the molt does not necessarily reflect habitat association of pupping seals. Ratios of pups to total seals counted during pupping and the subsequent molt were used to contrast habitat use. Low proportions of pups at terrestrial haul‐outs, relative to most glacial haul‐outs, indicate an overall preference for pupping in glacial haul‐outs. High proportions of pups at most glacial sites (during pupping and molting) suggest reduced use of tidewater glacier habitats by nonbreeders and molting seals. Results suggest more seals associate with glacial haul‐outs than currently estimated.     

Viability analysis of a threatened amphibian population: modelling the past,present and future

Conservation actions that have maintained populations in the past may not necessarily do so in the future. Population viability analysis provides one tool for exploring the impact of management actions on large temporal scales. However, there are relatively few long‐term data sets that provide the demographic and environmental data demanded by such models. Using a 37‐yr data set, we used RAMAS Metapop to model the persistence of natterjack toads Epidalea (Bufo) calamita on a heathland in southern Britain. A retrospective analysis showed that the best fit between the predicted population trajectories and the real population was when the management carried out was modelled as an increase in K of 150 toads yr^?1. However, even if ongoing management continues to improve K by a further 40–60 toads yr^?1 over the next 50 yr, the population still has an extinction risk of at least 60% if other factors remain unchanged. Sensitivity analyses and simulated management scenarios indicated that the population was most sensitive to changes in the survival of juvenile (i.e. 1–2 yr old) toads. In addition, if the frequency and severity of pond desiccation increases, the risk of extinction was predicted to increase as a result of reduced recruitment. Low levels of extinction risk occurred irrespective of K when juvenile survival was enhanced in combination with low frequency and severity of pond desiccation. The models suggest that populations that are responding to management against a background of natural fluctuations may remain vulnerable to extinction for several decades. These extinction risks may increase if habitat management fails to offset reductions in recruitment and juvenile survival caused by environmental change.     

Demographic responses to weather fluctuations are context dependent in a long‐lived amphibian

Weather fluctuations have been demonstrated to affect demographic traits in many species. In long‐lived organisms, their impact on adult survival might be buffered by the evolution of traits that reduce variation in interannual adult survival. For example, skipping breeding is an effective behavioral mechanism that may limit yearly variation in adult survival when harsh weather conditions occur; however, this in turn would likely lead to strong variation in recruitment. Yet, only a few studies to date have examined the impact of weather variation on survival, recruitment and breeding probability simultaneously in different populations of the same species. To fill this gap, we studied the impact of spring temperatures and spring rainfall on survival, on reproductive skipping behavior and on recruitment in five populations of a long‐lived amphibian, the yellow‐bellied toad (Bombina variegata). Based on capture–recapture data, our findings demonstrate that survival depends on interactions between age, population and weather variation. Varying weather conditions in the spring result in strong variation in the survival of immature toads, whereas they have little effect on adult toads. Breeding probability depends on both the individual's previous reproductive status and on the weather conditions during the current breeding season, leading to high interannual variation in recruitment. Crucially, we found that the impact of weather variation on demographic traits is largely context dependent and may thus differ sharply between populations. Our results suggest that studies predicting the impact of climate change on population dynamics should be taken with caution when the relationship between climate and demographic traits is established using only one population or few populations. We therefore highly recommend further research that includes surveys replicated in a substantial number of populations to account for context‐dependent variation in demographic processes.