Population Dynamics of the Andean Lizard Anolis heterodermus: Fast‐slow Demographic Strategies in Fragmented Scrubland Landscapes

Habitat fragmentation and loss affect population stability and demographic processes, increasing the extinction risk of species. We studied Anolis heterodermus populations inhabiting large and small Andean scrubland patches in three fragmented landscapes in the Sabana de Bogotá (Colombia) to determine the effect of habitat fragmentation and loss on population dynamics. We used the capture‐mark‐recapture method and multistate models to estimate vital rates for each population. We estimated growth population rate and the most important processes that affect λ by elasticity analysis of vital rates. We tested the effects of habitat fragmentation and loss on vital rates of lizard populations. All six isolated populations showed a positive or an equilibrium growth rate (λ = 1), and the most important demographic process affecting λ was the growth to first reproduction. Populations from landscapes with less scrubland natural cover showed higher stasis of young adults. Populations in highly fragmented landscapes showed highest juvenile survival and growth population rates. Independent of the landscape's habitat configuration and connectivity, populations from larger scrubland patches showed low adult survivorship, but high transition rates. Populations varied from a slow strategy with low growth and delayed maturation in smaller patches to a fast strategy with high growth and early maturation in large patches. This variation was congruent with the fast‐slow continuum hypothesis and has serious implications for Andean lizard conservation and management strategies. We suggest that more stable lizard populations will be maintained if different management strategies are adopted according to patch area and habitat structure.     

Cooperative breeding behaviors in the Hawaiian Stilt (Himantopus mexicanus knudseni)

Cooperative breeding, which is commonly characterized by nonbreeding individuals that assist others with reproduction, is common in avian species. However, few accounts have been reported in Charadriiformes, particularly island‐nesting species. We present incidental observations of cooperative breeding behaviors in the Hawaiian Stilt (Himantopus mexicanus knudseni), an endangered subspecies of the Black‐necked Stilt (Himantopus mexicanus), during the 2012–2020 nesting seasons on the Hawaiian islands of O‘ahu and Moloka‘i. We describe two different behaviors that are indicative of cooperative breeding: (a) egg incubation by multiple adults; (b) helpers‐at‐the‐nest, whereby juveniles delay dispersal and reproduction to assist parents and siblings with reproduction. These observations are the first published accounts of cooperative breeding in this subspecies and merit further investigation, as cooperative breeding may improve population viability of the endangered, endemic Hawaiian Stilt.     

Joint Effects of Habitat Heterogeneity and Species’ Life-History Traits on Population Dynamics in Spatially Structured Landscapes

Both habitat heterogeneity and species’ life-history traits play important roles in driving population dynamics, yet there is little scientific consensus around the combined effect of these two factors on populations in complex landscapes. Using a spatially explicit agent-based model, we explored how interactions between habitat spatial structure (defined here as the scale of spatial autocorrelation in habitat quality) and species life-history strategies (defined here by species environmental tolerance and movement capacity) affect population dynamics in spatially heterogeneous landscapes. We compared the responses of four hypothetical species with different life-history traits to four landscape scenarios differing in the scale of spatial autocorrelation in habitat quality. The results showed that the population size of all hypothetical species exhibited a substantial increase as the scale of spatial autocorrelation in habitat quality increased, yet the pattern of population increase was shaped by species’ movement capacity. The increasing scale of spatial autocorrelation in habitat quality promoted the resource share of individuals, but had little effect on the mean mortality rate of individuals. Species’ movement capacity also determined the proportion of individuals in high-quality cells as well as the proportion of individuals experiencing competition in response to increased spatial autocorrelation in habitat quality. Positive correlations between the resource share of individuals and the proportion of individuals experiencing competition indicate that large-scale spatial autocorrelation in habitat quality may mask the density-dependent effect on populations through increasing the resource share of individuals, especially for species with low mobility. These findings suggest that low-mobility species may be more sensitive to habitat spatial heterogeneity in spatially structured landscapes. In addition, localized movement in combination with spatial autocorrelation may increase the population size, despite increased density effects.     

Using Energetic Models to Investigate the Survival and Reproduction of Beaked Whales (family Ziphiidae)

Mass stranding of several species of beaked whales (family Ziphiidae) associated with exposure to anthropogenic sounds has raised concern for the conservation of these species. However, little is known about the species’ life histories, prey or habitat requirements. Without this knowledge, it becomes difficult to assess the effects of anthropogenic sound, since there is no way to determine whether the disturbance is impacting the species’ physical or environmental requirements. Here we take a bioenergetics approach to address this gap in our knowledge, as the elusive, deep-diving nature of beaked whales has made it hard to study these effects directly. We develop a model for Ziphiidae linking feeding energetics to the species’ requirements for survival and reproduction, since these life history traits would be the most likely to be impacted by non-lethal disturbances. Our models suggest that beaked whale reproduction requires energy dense prey, and that poor resource availability would lead to an extension of the inter-calving interval. Further, given current information, it seems that some beaked whale species require relatively high quality habitat in order to meet their requirements for survival and reproduction. As a result, even a small non-lethal disturbance that results in displacement of whales from preferred habitats could potentially impact a population if a significant proportion of that population was affected. We explored the impact of varying ecological parameters and model assumptions on survival and reproduction, and find that calf and fetus survival appear more readily affected than the survival of adult females.     

Demographic Variables for Wild Asian Elephants Using Longitudinal Observations

Detailed demographic data on wild Asian elephants have been difficult to collect due to habitat characteristics of much of the species’ remaining range. Such data, however, are critical for understanding and modeling population processes in this endangered species. We present data from six years of an ongoing study of Asian elephants (Elephas maximus) in Uda Walawe National Park, Sri Lanka. This relatively undisturbed population numbering over one thousand elephants is individually monitored, providing cohort-based information on mortality and reproduction. Reproduction was seasonal, such that most births occurred during the long inter-monsoon dry season and peaked in May. During the study, the average age at first reproduction was 13.4 years and the 50^th percentile inter-birth interval was approximately 6 years. Birth sex ratios did not deviate significantly from parity. Fecundity was relatively stable throughout the observed reproductive life of an individual (ages 11–60), averaging between 0.13–0.17 female offspring per individual per year. Mortalities and injuries based on carcasses and disappearances showed that males were significantly more likely than females to be killed or injured through anthropogenic activity. Overall, however, most observed injuries did not appear to be fatal. This population exhibits higher fecundity and density relative to published estimates on other Asian elephant populations, possibly enhanced by present range constriction. Understanding the factors responsible for these demographic dynamics can shed insight on the future needs of this elephant population, with probable parallels to other populations in similar settings.     

The limits of demographic buffering in coping with environmental variation

Animal populations have developed multiple strategies to deal with environmental change. Among them, the demographic buffering strategy consists in constraining the temporal variation of the vital rate(s) that most affect(s) the overall performance of the population. Tortoises are known to buffer their temporal variation in adult survival, which typically has the highest contribution to the population growth rate λ, at the expense of a high variability on reproductive rates, which contribute far less to λ. To identify the effects of projected increases in droughts in its natural habitat, we use field data collected across 15 locations of Testudo graeca in southeast Spain over a decade. We analyse the effects of environmental variables on reproduction rates. In addition, we couple the demographic and environmental data to parameterise an integral projection model to simulate the effects of different scenarios of drought recurrence on λ under different degrees of intensity in the survival–reproduction tradeoff. We find that droughts negatively affect the probability of laying eggs; however, the overall effects on λ under the current drought recurrence (one/decade) are negligible when survival is constant (independent of the reduction of reproduction by drought events) and when survival increased as a tradeoff with the reduction of reproduction rates, with a threshold to population viability at three or more droughts/decade. Additionally, we show that, although some species may buffer current environmental regimes by carefully orchestrating how their vital rates vary through time, a demographic buffering strategy is insufficient to ensure population viability in extreme regimes. Our findings support the hypothesis that the demographic buffering strategy has a limit of effectiveness when adverse conditions occur frequently. Our methodological approach provides a framework for ecologists to determine how effective the management of environmental drivers can be for demographically buffering populations, and which scenarios may not provide long-term population persistence.     

Population regulation of territorial species: both site dependence and interference mechanisms matter

Spatial patterns of site occupancy are commonly driven by habitat heterogeneity and are thought to shape population dynamics through a site-dependent regulatory mechanism. When examining this, however, most studies have only focused on a single vital rate (reproduction), and little is known about how space effectively contributes to the regulation of population dynamics. We investigated the underlying mechanisms driving density-dependent processes in vital rates in a Mauritius kestrel population where almost every individual was monitored. Different mechanisms acted on different vital rates, with breeding success regulated by site dependence (differential use of space) and juvenile survival by interference (density-dependent competition for resources). Although territorial species are frequently assumed to be regulated through site dependence, we show that interference was the key regulatory mechanism in this population. Our integrated approach demonstrates that the presence of spatial processes regarding one trait does not mean that they necessarily play an important role in regulating population growth, and demonstrates the complexity of the regulatory process.     

Managing multiple vital rates to maximize greater sage-grouse population growth

Despite decades of field research on greater sage-grouse, range-wide demographic data have yet to be synthesized into a sensitivity analysis to guide management actions. We reviewed range-wide demographic rates for greater sage-grouse from 1938 to 2011 and used data from 50 studies to parameterize a 2-stage, female-based population matrix model. We conducted life-stage simulation analyses to determine the proportion of variation in population growth rate (λ) accounted for by each vital rate, and we calculated analytical sensitivity, elasticity, and variance-stabilized sensitivity to identify the contribution of each vital rate to λ. As expected for an upland game bird, greater sage-grouse showed marked annual and geographic variation in several vital rates. Three rates were demonstrably important for population growth: female survival, chick survival, and nest success. Female survival and chick survival, in that order, had the most influence on λ per unit change in vital rates. However, nest success explained more of the variation in λ than did the survival rates. In lieu of quantitative data on specific mortality factors driving local populations, we recommend that management efforts for greater sage-grouse first focus on increasing female survival by restoring large, intact sagebrush-steppe landscapes, reducing persistent sources of human-caused mortality, and eliminating anthropogenic habitat features that subsidize species that prey on juvenile, yearling, and adult females. Our analysis also supports efforts to increase chick survival and nest success by eliminating anthropogenic habitat features that subsidize chick and nest predators, and by managing shrub, forb, and grass cover, height, and composition to meet local brood-rearing and nesting habitat guidelines. We caution that habitat management to increase chick survival and nest success should not reduce the cover or height of sagebrush below that required for female survival in other seasons (e.g., fall, winter). The success or failure of management actions for sage-grouse should be assessed by measuring changes in vital rates over long time periods to avoid confounding with natural, annual variation. © 2011 The Wildlife Society.     

Unexpected Demography in the Recovery of an Endangered Primate Population

Assessments of the status of endangered species have focused on population sizes, often without knowledge of demographic and behavioral processes underlying population recovery. We analyzed demographic data from a 28-year study of a critically endangered primate, the northern muriqui, to investigate possible changes in demographic rates as this population recovered from near extirpation. As the population increased from 60 to nearly 300 individuals, its growth rate declined due to increased mortality and male-biased birth sex ratios; the increased mortality was not uniform across ages and sexes, and there has been a recent increase in mortality of prime-aged males. If not for a concurrent increase in fertility rates, the population would have stabilized at 200 individuals instead of continuing to grow. The unexpected increase in fertility rates and in adult male mortality can be attributed to the muriquis’ expansion of their habitat by spending more time on the ground. The demographic consequences of this behavioral shift must be incorporated into management tactics for this population and emphasize the importance of understanding demographic rates in the recovery of endangered species.     

Evolution of allometries in the worker caste of Dorylus army ants

We investigated the effect of local environment on the demography and population dynamics of arctic ground squirrels ( Spermophilus parryii plesius ) by comparing reproduction, survival, and population trends of squirrels living in low elevation boreal forest and high elevation alpine tundra sites in southwestern Yukon Territory, Canada. Contrary to the trend for most birds and mammals, reproduction was significantly lower at the lower elevation and females living at higher elevation did not delay the age at which they first reproduced. Even though survival in the boreal forest was lower in summer than in the alpine, it was higher over winter so annual adult female survival was similar between sites.
Sensitivity analysis of model parameters revealed that in the forest, population growth rate (λ) was most sensitive to small changes in adult active season survival whereas for the alpine population, λ was most sensitive to changes in juvenile winter survival. In their respective habitats, these parameters also showed high year to year variation and thus contributed greatly to the population trends observed. Even though ground squirrels persisted in the boreal forest, the measured demographic rates indicate the forest was sink habitat (λ<1) and may have relied on nearby grassy meadows for immigrants. In contrast, the alpine habitat maintained a ground squirrel population in the absence of immigration (λ=1).
The variation in demographic rates between ground squirrels living at high and low elevation may arise from phenotypic responses of squirrels to different habitat structure. Arctic ground squirrels rely on sight to detect predators from a safe distance, and the boreal forest, with its lower visibility and higher predator density, appears to be suboptimal habitat.     

Reduced habitat quality increases intrinsic but not ecological costs of reproduction

Although the costs of reproduction are predicted to vary with the quality of the breeding habitat thereby affecting population dynamics and life‐history trade‐offs, empirical evidence for this pattern remains sparse and equivocal. Costs of reproduction can operate through immediate ecological mechanisms or through delayed intrinsic mechanisms. Ignoring these separate pathways might hinder the identification of costs and the understanding of their consequences. We experimentally investigated the survival costs of reproduction for adult little owls (Athene noctua) within a gradient of habitat quality. We supplemented food to nestlings, thereby relieving the parents’ effort for brood provisioning. We used radio‐tracking and Bayesian multistate modeling based on marked recapture and dead recovery to estimate survival rates of adult little owls across the year as a function of food supplementation and habitat characteristics. Food supplementation to nestlings during the breeding season increased parental survival not only during the breeding season but also during the rest of the year. Thus, the low survival of parents of unfed broods likely represents both, strong ecological and strong intrinsic costs of reproduction. However, while immediate ecological costs occurred also in high‐quality habitats, intrinsic costs carrying over to the post‐breeding period occurred only in low‐quality habitats. Our results suggest that immediate costs resulting from ecological mechanisms such as predation, are high also in territories of high habitat quality. Long‐term costs resulting from intrinsic trade‐offs, however, are only paid in low‐quality habitats. Consequently, differential effects of habitat quality on immediate ecological and delayed intrinsic mechanisms can mask the increase of costs of reproduction in low‐quality breeding habitats. Intrinsic costs may represent an underrated mechanism of habitat quality affecting adult survival rate thereby considerably accelerating population decline in degrading habitats. This study therefore highlights the need for a long‐term perspective to fully assess the costs of reproduction and the role of habitat quality in modifying these costs.     

Estimating Minimum Area of Suitable Habitat and Viable Population Size for the Northern Muriqui (Brachyteles hypoxanthus)

A population viability analysis (PVA) using the computer package VORTEX was conducted to assess the minimum viable population (MVP) of the Atlantic Forest endemic primate Brachyteles hypoxanthus. The objectives were: (1) to estimate demographic and genetic MVPs that could be used as quasi-extinction thresholds for future modeling, and (2) to estimate the minimum area of suitable habitat (MASH). The model predicted that populations of 40 and 700 individuals were necessary to achieve demographic and genetic stability, respectively. The model was more sensitive to changes in inbreeding depression, sex ratio and reproduction (percentage of breeding females). MASH estimated to contain genetically viable populations reached 11,570 ha. Muriquis have managed to persist despite severe habitat disturbance, but the results suggest that although most of the extant populations are not threatened by extinction, they are too small to be genetically viable in the long-run, and will loose most of their heterozygosity.     

Prioritizing Avian Species for Their Risk of Population-Level Consequences from Wind Energy Development

Recent growth in the wind energy industry has increased concerns about its impacts on wildlife populations. Direct impacts of wind energy include bird and bat collisions with turbines whereas indirect impacts include changes in wildlife habitat and behavior. Although many species may withstand these effects, species that are long-lived with low rates of reproduction, have specialized habitat preferences, or are attracted to turbines may be more prone to declines in population abundance. We developed a prioritization system to identify the avian species most likely to experience population declines from wind facilities based on their current conservation status and their expected risk from turbines. We developed 3 metrics of turbine risk that incorporate data on collision fatalities at wind facilities, population size, life history, species’ distributions relative to turbine locations, number of suitable habitat types, and species’ conservation status. We calculated at least 1 measure of turbine risk for 428 avian species that breed in the United States. We then simulated 100,000 random sets of cutoff criteria (i.e., the metric values used to assign species to different priority categories) for each turbine risk metric and for conservation status. For each set of criteria, we assigned each species a priority score and calculated the average priority score across all sets of criteria. Our prioritization system highlights both species that could potentially experience population decline caused by wind energy and species at low risk of population decline. For instance, several birds of prey, such as the long-eared owl, ferruginous hawk, Swainson’s hawk, and golden eagle, were at relatively high risk of population decline across a wide variety of cutoff values, whereas many passerines were at relatively low risk of decline. This prioritization system is a first step that will help researchers, conservationists, managers, and industry target future study and management activity.     

Demographic Outcomes and Ecosystem Implications of Giant Tortoise Reintroduction to Espa?ola Island,Galapagos

Restoration of extirpated species via captive breeding has typically relied on population viability as the primary criterion for evaluating success. This criterion is inadequate when species reintroduction is undertaken to restore ecological functions and interactions. Herein we report on the demographic and ecological outcomes of a five-decade-long population restoration program for a critically endangered species of “ecosystem engineer”: the endemic Española giant Galapagos tortoise (Chelonoidis hoodensis). Our analysis of complementary datasets on tortoise demography and movement, tortoise-plant interactions and Española Island’s vegetation history indicated that the repatriated tortoise population is secure from a strictly demographic perspective: about half of tortoises released on the island since 1975 were still alive in 2007, in situ reproduction is now significant, and future extinction risk is low with or without continued repatriation. Declining survival rates, somatic growth rates, and body condition of repatriates suggests, however, that resources for continued population growth are increasingly limited. Soil stable carbon isotope analyses indicated a pronounced shift toward woody plants in the recent history of the island’s plant community, likely a legacy of changes in competitive relations between woody and herbaceous plants induced by now-eradicated feral goats and prolonged absence of tortoises. Woody plants are of concern because they block tortoise movement and hinder recruitment of cactus–a critical resource for tortoises. Tortoises restrict themselves to remnant cactus patches and areas of low woody plant density in the center of the island despite an apparent capacity to colonize a far greater range, likely because of a lack of cactus elsewhere on the island. We conclude that ecosystem-level criteria for success of species reintroduction efforts take much longer to achieve than population-level criteria; moreover, reinstatement of endangered species as fully functioning ecosystem engineers may often require large-scale habitat restoration efforts in concert with population restoration.     

Abundance decline in the avifauna of the European Union reveals cross‐continental similarities in biodiversity change

Although global assessments provide evidence of biodiversity decline, some have questioned the strength of the evidence, with local assemblage studies often showing a more balanced picture of biodiversity change. The multifaceted nature of biodiversity and imperfect monitoring datasets may partially explain these findings. Here, using an extensive dataset, we find significant biodiversity loss in the native avifauna of the European Union (EU). We estimate a decline of 17–19% in the overall breeding bird abundance since 1980: a loss of 560–620 million individual birds. Both total and proportional declines in bird numbers are high among species associated with agricultural land. The distribution of species’ population growth rates (ln) is centered close to zero, with numerical decline driven by substantial losses in abundant species. Our work supports previous assessments indicating substantial recent biodiversity loss and calls to reduce the threat of extinctions and restore species’ abundances, for the sake of nature and people.     

Effects of climate variability on the demography of wild geladas

Nonhuman primates are an essential part of tropical biodiversity and play key roles in many ecosystem functions, processes, and services. However, the impact of climate variability on nonhuman primates, whether anthropogenic or otherwise, remains poorly understood. In this study, we utilized age‐structured matrix population models to assess the population viability and demographic variability of a population of geladas (Theropithecus gelada) in the Simien Mountains, Ethiopia with the aim of revealing any underlying climatic influences. Using data from 2008 to 2019 we calculated annual, time‐averaged, and stochastic population growth rates (λ) and investigated relationships between vital rate variability and monthly cumulative rainfall and mean temperature. Our results showed that under the prevailing environmental conditions, the population will increase (λ _s = 1.021). Significant effects from rainfall and/or temperature variability were widely detected across vital rates; only the first year of infant survival and the individual years of juvenile survival were definitively unaffected. Generally, the higher temperature in the hot‐dry season led to lower survival and higher fecundity, while higher rainfall in the hot‐dry season led to increased survival and fecundity. Overall, these results provide evidence of greater effects of climate variability across a wider range of vital rates than those found in previous primate demography studies. This highlights that although primates have often shown substantial resilience to the direct effects of climate change, their vulnerability may vary with habitat type and across populations.     

Population Dynamics of the Endangered, Long-Lived Perennial Species, Ligularia sibirica

Ligularia sibirica is a relict wetland perennial plant species of the Czech and Slovak Republic. Explaining variation in population growth rate and identifying the causes of that variation is important for effective protection of such an endangered species. Matrix models based on four years of data of 11 populations were used to identify the pattern of variation in the demographic vital rates of this species, and to examine the causes of the variation such as population size and habitat type. Further, the matrix model was used to determine the population growth rate, longevity and risk of extinction of each population and to identify the specific vital rates that most affect population growth rate. The results showed that population growth rates were significantly different between years and populations. Temporal variation was mostly due to variable survival of adult individuals, while spatial variation was mainly driven by fertility of one small currently expanding population. Further, most studied populations of L. sibirica are performing well and only those growing in nitrogen-rich habitats have a high extinction risk. The results also indicate that all populations have low adult mortality, long-lived individuals (61.3?years on average) and some populations also show features of remnant populations (i.e., the persistence of populations in severe conditions in spite of no reproduction). Our results imply that detailed demographic data are needed to understand the long-term prospects of these populations. These data may serve as an early warning system for this species long before an obvious decline occurs in the populations.     

Using Multiscale Spatial Models to Assess Potential Surrogate Habitat for an Imperiled Reptile

In evaluating conservation and management options for species, practitioners might consider surrogate habitats at multiple scales when estimating available habitat or modeling species’ potential distributions based on suitable habitats, especially when native environments are rare. Species’ dependence on surrogates likely increases as optimal habitat is degraded and lost due to anthropogenic landscape change, and thus surrogate habitats may be vital for an imperiled species’ survival in highly modified landscapes. We used spatial habitat models to examine a potential surrogate habitat for an imperiled ambush predator (eastern diamondback rattlesnake, Crotalus adamanteus; EDB) at two scales. The EDB is an apex predator indigenous to imperiled longleaf pine ecosystems (Pinus palustris) of the southeastern United States. Loss of native open-canopy pine savannas and woodlands has been suggested as the principal cause of the species’ extensive decline. We examined EDB habitat selection in the Coastal Plain tidewater region to evaluate the role of marsh as a potential surrogate habitat and to further quantify the species’ habitat requirements at two scales: home range (HR) and within the home range (WHR). We studied EDBs using radiotelemetry and employed an information-theoretic approach and logistic regression to model habitat selection as use vs. availability. We failed to detect a positive association with marsh as a surrogate habitat at the HR scale; rather, EDBs exhibited significantly negative associations with all landscape patches except pine savanna. Within home range selection was characterized by a negative association with forest and a positive association with ground cover, which suggests that EDBs may use surrogate habitats of similar structure, including marsh, within their home ranges. While our HR analysis did not support tidal marsh as a surrogate habitat, marsh may still provide resources for EDBs at smaller scales.     

Global changes explain the long-term demographic trend of the Eurasian common lizard (Squamata: Lacertidae)

The demographic trend of a species depends on the dynamics of its local populations, which can be compromised by local or by global phenomena. However, the relevance of local and global phenomena has rarely been investigated simultaneously. Here, we tested whether local phenomena compromised a species’ demographic trend using the Eurasian common lizard Zootoca vivipara, the terrestrial reptile exhibiting the widest geographic distribution, as a model species. We analyzed the species’ ancient demographic trend using genetic data from its 6 allopatric genetic clades and tested whether its demographic trend mainly depended on single clades or on global phenomena. Zootoca vivipara’s effective population size increased since 2.3 million years ago and started to increase steeply and continuously from 0.531 million years ago. Population growth rate exhibited 2 maxima, both occurring during global climatic changes and important vegetation changes on the northern hemisphere. Effective population size and growth rate were negatively correlated with global surface temperatures, in line with global parameters driving long-term demographic trends. Zootoca vivipara’s ancient demography was neither driven by a single clade, nor by the 2 clades that colonized huge geographic areas after the last glaciation. The low importance of local phenomena, suggests that the experimentally demonstrated high sensitivity of this species to short-term ecological changes is a response in order to cope with short-term and local changes. This suggests that what affected its long-term demographic trend the most, were not these local changes/responses, but rather the important and prolonged global climatic changes and important vegetation changes on the northern hemisphere, including the opening up of the forest by humans.     

Evaluating the demographic buffering hypothesis with vital rates estimated for Weddell seals from 30 years of mark-recapture data

1. Life-history theory predicts that those vital rates that make larger contributions to population growth rate ought to be more strongly buffered against environmental variability than are those that are less important. Despite the importance of the theory for predicting demographic responses to changes in the environment, it is not yet known how pervasive demographic buffering is in animal populations because the validity of most existing studies has been called into question because of methodological deficiencies. 2. We tested for demographic buffering in the southern-most breeding mammal population in the world using data collected from 5558 known-age female Weddell seals over 30 years. We first estimated all vital rates simultaneously with mark-recapture analysis and then estimated process variance and covariance in those rates using a hierarchical Bayesian approach. We next calculated the population growth rate's sensitivity to changes in each of the vital rates and tested for evidence of demographic buffering by comparing properly scaled values of sensitivity and process variance in vital rates. 3. We found evidence of positive process covariance between vital rates, which indicates that all vital rates are affected in the same direction by changes in annual environment. Despite the positive correlations, we found strong evidence that demographic buffering occurred through reductions in variation in the vital rates to which population growth rate was most sensitive. Process variation in vital rates was inversely related to sensitivity measures such that variation was greatest in breeding probabilities, intermediate for survival rates of young animals and lowest for survival rates of older animals. 4. Our work contributes to a small but growing set of studies that have used rigorous methods on long-term, detailed data to investigate demographic responses to environmental variation. The information from these studies improves our understanding of life-history evolution in stochastic environments and provides useful information for predicting population responses to future environmental change. Our results for an Antarctic apex predator also provide useful baselines from a marine ecosystem when its top- and middle-trophic levels were not substantially impacted by human activity.