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Effects of fat reserves on annual apparent survival of blackbirds Turdus merula

1. Fat reserves are stored energy that may help birds survive periods of harsh winter weather. This hypothesis predicts that annual apparent survival is higher for birds with large fat reserves than for birds with few or no fat reserves in winter. 2. Blackbirds ( Turdus merula Linnaeus) were ringed in central Italy from 16 November to 20 February during 1990-2001. Fat scores were recorded for each bird. We used these capture-mark-recapture data for 1703 blackbirds to estimate the effect of large fat reserves on annual apparent survival , while controlling for transients, using computer programs SURVIV and MARK. Probability of birds retaining large fat reserves , or retaining few fat reserves , over 2 successive years was also estimated. 3. Birds with large fat reserves did not have higher estimated annual apparent survival than birds with few fat reserves (φ̂ large = φ̂ few = 0.595, SE = 0.043), inconsistent with our prediction. No effects of age, sex or year were detected on annual apparent survival . Birds with few fat reserves in any given year tended to have few fat reserves the following year (ψ̂ few→large = 0.332, SE = 0.052). Birds with large fat reserves in any given year were unlikely to have large fat reserves the next year (ψ̂ large→few = 0.585, SE = 0.080). 4. Large fat reserves may not increase annual survival of blackbirds wintering in central Italy. Winter weather in our study area may be too mild to effect survival . Alternatively, increased predation risk associated with large fat reserves may counteract any benefits of reduced starvation risk.

Journal of Animal Ecology

Factors affecting breeding dispersal of European ducks on Engure Marsh, Latvia

1. We used up to 35 years of capture-recapture data from nearly 3300 individual female ducks nesting on Engure Marsh , Latvia , and multistate modelling to test predictions about the influence of environmental, habitat and management factors on breeding dispersal probability within the marsh . 2. Analyses based on observed dispersal distances of common pochards and tufted ducks provided no evidence that breeding success in year t influenced dispersal distance between t and t + 1. 3. Breeding dispersal distances (year t to t + 1) of pochards and tufted ducks were associated with a delay in relative nest initiation dates in year t + 1. The delay was greater for pochards (c. 4 days) than for tufted ducks (c. 2 days) when females dispersed > 0.8 km. 4. Northern shovelers and tufted ducks moved from a large island to small islands at low water levels and from small islands to the large island at high water levels before the construction of elevated small islands (1960-82). Following this habitat management (1983-94), breeding fidelity was extremely high and not influenced by water level in the marsh for either species. 5. Because pochard nesting habitats in black-headed gull colonies were saturated during the entire study period, nesting females moved into and out of colonies with similar probabilities. Local survival probabilities and incubation body masses were higher for both yearlings (SY) and adults (ASY) nesting within gull colonies, suggesting that these females were of better quality than females nesting outside of the colonies. 6. Tufted ducks showed higher probabilities of moving from islands to emergent marshes when water levels were higher both before and after habitat management. However, rates of movement for a given water level were higher during the period before management than after. 7. Both pochards and tufted ducks exhibited asymmetric movement with respect to proximity to water, with higher movement probabilities to near-water nesting locations than away from these locations. 8. Multistate capture-recapture models provided analyses that were useful in investigating sources of variation in breeding dispersal probabilities.

Journal of Animal Ecology

Estimating natal dispersal movement rates of female European ducks with multistate modelling

1. We used up to 34 years of capture-recapture data from about 22 100 new releases of day-old female ducklings and multistate modelling to test predictions about the influence of environmental, habitat and management factors on natal dispersal probability of three species of ducks within the Engure Marsh, Latvia. 2. The mean natal dispersal distances were very similar (c. 0.6-0.7 km) for all three species and were on average 2.7 times greater than breeding dispersal distances recorded within the same study system. 3. We were unable to confirm the kinship hypothesis and found no evidence that young first-nesting females nested closer to their relatives (either mother or sister) than to the natal nest. 4. Young female northern shovelers, like adults, moved from small islands to the large island when water level was high and vice versa when water level was low before the construction of elevated small islands. Movement probabilities between the two strata were much higher for young shovelers than adults, suggesting that young birds had not yet developed strong fidelity to the natal site. Movements of young female tufted ducks , unlike those of shovelers, were not dependent on water level fluctuations and reflected substantial flexibility in choice of first nesting sites. 5. Data for young birds supported our earlier conclusion that common pochard nesting habitats in black-headed gull colonies were saturated during the entire study period. Young females, like the two adult age groups, moved into and out of colonies with similar probability. Fidelity probability of female pochards to each stratum increased with age, being the lowest (0.62) for young (DK) females, intermediate (0.78) for yearlings (SY) and the highest (0.84) for adult (ASY) females. 6. Young female tufted ducks , like adults, showed higher probabilities of moving from islands to emergent marshes when water levels were higher both before and after habitat management. The relationship between the spring water levels and movement was much weaker for young females than for adults. 7. Young female diving ducks exhibited much stronger (compared to adults) asymmetric movement with respect to proximity to water, with higher movement probabilities to near-water locations than away from these locations. 8. Local survival of day-old ducklings during the first year of life was time-specific and very low (means for different strata/states 0.01-0.08) because of high rates of emigration and prefledging mortality.

Journal of Animal Ecology

Elk herbivory alters small mammal assemblages in high elevation drainages

Heavy herbivory by ungulates can substantially alter habitat, but the indirect consequences of habitat modification for animal assemblages that rely on that habitat are not well studied. This is a particularly important topic given that climate change can alter plant–herbivore interactions. We explored short-term responses of small mammal communities to recent exclusion of Rocky Mountain elk ( Cervus elaphus ) in high-elevation riparian drainages in northern Arizona, where elk impacts on vegetation have increased over the past quarter century associated with climate change. We used 10-ha elk exclosures paired with unfenced control drainages to examine how browsing influenced the habitat use, relative abundance, richness and diversity of a small mammal assemblage. We found that the small mammal assemblage changed significantly after 5 years of elk exclusion. Relative abundance of voles ( Microtus mexicanus ) increased in exclosure drainages, likely due to an increase in habitat quality. The relative abundances of woodrats ( Neotoma neomexicana ) and two species of mice ( Peromyscus maniculatus and P. boylii ) decreased in the controls, while remaining stable in exclosures. The decline of mice in control drainages was likely due to the decline in shrub cover that they use. Thus, elk exclusion may have maintained or improved habitat for mice inside the exclosures while habitat quality and mouse abundance both declined outside the fences. Finally, small mammal species richness increased in the exclosures relative to the controls while species diversity showed no significant trends. Together, our results show that relaxation of heavy herbivore pressure by a widespread native ungulate can lead to rapid changes in small mammal assemblages. Moreover, exclusion of large herbivores can yield rapid responses by vegetation that may enhance or maintain habitat quality for small mammal populations.

Arizona

Spatio-temporal dynamics of pneumonia in bighorn sheep

Bighorn sheep mortality related to pneumonia is a primary factor limiting population recovery across western North America, but management has been constrained by an incomplete understanding of the disease. We analysed patterns of pneumonia-caused mortality over 14 years in 16 interconnected bighorn sheep populations to gain insights into underlying disease processes. 2. We observed four age-structured classes of annual pneumonia mortality patterns: all-age, lamb-only, secondary all-age and adult-only. Although there was considerable variability within classes, overall they differed in persistence within and impact on populations. Years with pneumonia-induced mortality occurring simultaneously across age classes (i.e. all-age) appeared to be a consequence of pathogen invasion into a naïve population and resulted in immediate population declines. Subsequently, low recruitment due to frequent high mortality outbreaks in lambs, probably due to association with chronically infected ewes, posed a significant obstacle to population recovery. Secondary all-age events occurred in previously exposed populations when outbreaks in lambs were followed by lower rates of pneumonia-induced mortality in adults. Infrequent pneumonia events restricted to adults were usually of short duration with low mortality. 3. Acute pneumonia-induced mortality in adults was concentrated in fall and early winter around the breeding season when rams are more mobile and the sexes commingle. In contrast, mortality restricted to lambs peaked in summer when ewes and lambs were concentrated in nursery groups. 4. We detected weak synchrony in adult pneumonia between adjacent populations, but found no evidence for landscape-scale extrinsic variables as drivers of disease. 5. We demonstrate that there was a >60% probability of a disease event each year following pneumonia invasion into bighorn sheep populations. Healthy years also occurred periodically, and understanding the factors driving these apparent fade-out events may be the key to managing this disease. Our data and modelling indicate that pneumonia can have greater impacts on bighorn sheep populations than previously reported, and we present hypotheses about processes involved for testing in future investigations and management.

Journal of Animal Ecology

Reconciling resource utilization and resource selection functions

Summary: 1. Analyses based on utilization distributions (UDs) have been ubiquitous in animal space use studies, largely because they are computationally straightforward and relatively easy to employ. Conventional applications of resource utilization functions (RUFs) suggest that estimates of UDs can be used as response variables in a regression involving spatial covariates of interest. 2. It has been claimed that contemporary implementations of RUFs can yield inference about resource selection, although to our knowledge, an explicit connection has not been described. 3. We explore the relationships between RUFs and resource selection functions from a hueristic and simulation perspective. We investigate several sources of potential bias in the estimation of resource selection coefficients using RUFs (e.g. the spatial covariance modelling that is often used in RUF analyses). 4. Our findings illustrate that RUFs can, in fact, serve as approximations to RSFs and are capable of providing inference about resource selection, but only with some modification and under specific circumstances. 5. Using real telemetry data as an example, we provide guidance on which methods for estimating resource selection may be more appropriate and in which situations. In general, if telemetry data are assumed to arise as a point process, then RSF methods may be preferable to RUFs; however, modified RUFs may provide less biased parameter estimates when the data are subject to location error.

Journal of Animal Ecology

Estimating animal resource selection from telemetry data using point process models

Analyses of animal resource selection functions (RSF) using data collected from relocations of individuals via remote telemetry devices have become commonplace. Increasing technological advances, however, have produced statistical challenges in analysing such highly autocorrelated data. Weighted distribution methods have been proposed for analysing RSFs with telemetry data. However, they can be computationally challenging due to an intractable normalizing constant and cannot be aggregated (i.e. collapsed) over time to make space-only inference. In this study, we take a conceptually different approach to modelling animal telemetry data for making RSF inference. We consider the telemetry data to be a realization of a space–time point process. Under the point process paradigm, the times of the relocations are also considered to be random rather than fixed. We show the point process models we propose are a generalization of the weighted distribution telemetry models. By generalizing the weighted model, we can access several numerical techniques for evaluating point process likelihoods that make use of common statistical software. Thus, the analysis methods can be readily implemented by animal ecologists. In addition to ease of computation, the point process models can be aggregated over time by marginalizing over the temporal component of the model. This allows a full range of models to be constructed for RSF analysis at the individual movement level up to the study area level. To demonstrate the analysis of telemetry data with the point process approach, we analysed a data set of telemetry locations from northern fur seals (Callorhinus ursinus) in the Pribilof Islands, Alaska. Both a space–time and an aggregated space-only model were fitted. At the individual level, the space–time analysis showed little selection relative to the habitat covariates. However, at the study area level, the space-only model showed strong selection relative to the covariates.

Alaska

Location-only and use-availability data: analysis methods converge

This Special Feature arose from a session on a topic of the same name that took place during The Wildlife Society meeting in Kona, Hawaii, from 5 to 10 November, 2011. The purpose of that session and this Special Feature is to compare methods for predictive modelling of species geographical distributions and the modelling of habitat (resource) selection by animals. The predictive modelling of species geographical distributions and the modelling of habitat selection based on the environmental conditions at sites where animals are known to occur are essentially the same problem. Presence-only and used-available data both consist of a sample of locations with known presence of a species or an individual. A separate sample of locations from a study area, with unknown presence (pseudo-absence), is also assumed to exist. The probability or relative probability of presence of a species or individual is modelled and estimated across a certain time implicitly defined by the sampling mechanism, for example, by the time period during which museum specimens or radiotelemetry data were collected. A number of modelling methods have appeared in the literature over the last couple of decades. Many of these methods were made feasible by the availability of geographical information systems (GIS), global positioning system (GPS) radiotelemetry and public online data access initiatives (e.g. global biodiversity information facility). The papers in this Special Feature are intended to present the state of the methodological art in their subject area, with particular attention paid to contrasting the advantages and disadvantages of alternative methods of analysis for data.

Journal of Animal Ecology

Latitudinal and photic effects on diel foraging and predation risk in freshwater pelagic ecosystems

1. Clark & Levy ( American Naturalist , 131 , 1988, 271–290) described an antipredation window for smaller planktivorous fish during crepuscular periods when light permits feeding on zooplankton, but limits visual detection by piscivores. Yet, how the window is influenced by the interaction between light regime, turbidity and cloud cover over a broad latitudinal gradi- ent remains unexplored. 2. We evaluated how latitudinal and seasonal shifts in diel light regimes alter the foraging- risk environment for visually feeding planktivores and piscivores across a natural range of turbidities and cloud covers. Pairing a model of aquatic visual feeding with a model of sun and moon illuminance, we estimated foraging rates of an idealized planktivore and piscivore over depth and time across factorial combinations of latitude (0 – 70 ° ), turbidity (0  1 – 5 NTU) and cloud cover (clear to overcast skies) during the summer solstice and autumnal equinox. We evaluated the foraging-risk environment based on changes in the magnitude, duration and peak timing of the antipredation window. 3. The model scenarios generated up to 10-fold shifts in magnitude, 24-fold shifts in duration and 5  5-h shifts in timing of the peak antipredation window. The size of the window increased with latitude. This pattern was strongest during the solstice. In clear water at low turbidity (0  1 – 0  5 NTU), peaks in the magnitude and duration of the window formed at 57 – 60 ° latitude, before falling to near zero as surface waters became saturated with light under a midnight sun and clear skies at latitudes near 70 ° . Overcast dampened the midnight sun enough to allow larger windows to form in clear water at high latitudes. Conversely, at turbidities ≥ 2 NTU, greater reductions in the visual range of piscivores than planktivores created a window for long periods at high latitudes. Latitudinal dependencies were essentially lost during the equinox, indicating a progressive compression of the window from early summer into autumn. 4. Model results show that diel-seasonal foraging and predation risk in freshwater pelagic ecosystems changes considerably with latitude, turbidity and cloud cover. These changes alter the structure of pelagic predator – prey interactions, and in turn, the broader role of pelagic consumers in habitat coupling in lakes.

Journal of Animal Ecology

Life-stage-specific physiology defines invasion extent of a riverine fish

Many ecologists have called for mechanism-based investigations to identify the underlying controls on species distributions. Understanding these controls can be especially useful to construct robust predictions of how a species range may change in response to climate change or the extent to which a non-native species may spread in novel environments. Here, we link spatially intensive observations with mechanistic models to illustrate how physiology determines the upstream extent of the aquatic ectotherm smallmouth bass ( Micropterus dolomieu ) in two headwater rivers. Our results demonstrate that as temperatures become increasingly cold across a downstream to upstream gradient, food consumption in age 0 bass becomes increasingly constrained, and as a result, these fish become growth limited. Sufficient first summer growth of age 0 bass is essential for overwinter survival because young bass must persist from energy reserves accumulated during the summer, and those reserves are determined by body size. Our field data reveal the upstream extent of adult bass reproduction corresponds to a point in the downstream/upstream gradient where cold temperatures impair growth opportunities in young bass. This pattern was repeated in both study streams and explained why bass positioned nests twice as far upstream in the warm compared to the cold stream in the same basin. Placement of spawning nests by adult bass is likely subject to strong evolutionary selection in temperate systems: if bass spawn too far upstream, their young are unlikely to grow large enough to survive the winter. Consumption and growth in older bass (age 3–4) was far less sensitive to temperature. Based on these data, we suggest that temperature-sensitive age 0 bass constrain the upstream distribution limits of bass within temperate streams. In this study, we investigated how temperature-dependent physiology changed through the life history of a species and, in doing so, identified a climate-sensitive life-history stage that likely sets the distributional limits of all other life-history stages. We anticipate the framework developed here could be employed to identify how similar stage-specific environmental sensitivity determines distribution in many other ectothermic species.

Oregon

Context-dependent survival, fecundity and predicted population-level consequences of brucellosis in African buffalo

Chronic infections may have negative impacts on wildlife populations, yet their effects are difficult to detect in the absence of long-term population monitoring. Brucella abortus , the bacteria responsible for bovine brucellosis, causes chronic infections and abortions in wild and domestic ungulates, but its impact on population dynamics is not well understood. We report infection patterns and fitness correlates of bovine brucellosis in African buffalo based on (1) 7 years of cross-sectional disease surveys and (2) a 4-year longitudinal study in Kruger National Park (KNP), South Africa. We then used a matrix population model to translate these observed patterns into predicted population-level effects. Annual brucellosis seroprevalence ranged from 8&middot;7% (95% CI = 1&middot;8&ndash;15&middot;6) to 47&middot;6% (95% CI = 35&middot;1&ndash;60&middot;1) increased with age until adulthood (>6) and varied by location within KNP. Animals were on average in worse condition after testing positive for brucellosis ( F = &minus;5&middot;074, P < 0&middot;0001), and infection was associated with a 2&middot;0 (95% CI = 1&middot;1&ndash;3&middot;7) fold increase in mortality (&chi; 2 = 2&middot;039, P = 0&middot;036). Buffalo in low body condition were associated with lower reproductive success ( F = 2&middot;683, P = 0&middot;034), but there was no association between brucellosis and pregnancy or being observed with a calf. For the range of body condition scores observed in the population, the model-predicted growth rate was &lambda; = 1&middot;11 (95% CI = 1&middot;02&ndash;1&middot;21) in herds without brucellosis and &lambda; = 1&middot;00 (95% CI = 0&middot;85&ndash;1&middot;16) when brucellosis seroprevalence was 30%. Our results suggest that brucellosis infection can potentially result in reduced population growth rates, but because these effects varied with demographic and environmental conditions, they may remain unseen without intensive, longitudinal monitoring.

Kruger National Park

Host contact and shedding patterns clarify variation in pathogen exposure and transmission in threatened tortoise Gopherus agassizii : implications for disease modelling and management

Summary Most directly transmitted infections require some form of close contact between infectious and susceptible hosts to spread. Often disease models assume contacts are equal and use mean field estimates of transmission probability for all interactions with infectious hosts. Such methods may inaccurately describe transmission when interactions differ substantially in their ability to cause infection. Understanding this variation in transmission risk may be critical to properly model and manage some infectious diseases. In this study, we investigate how varying exposure and transmission may be key to understanding disease dynamics in the threatened desert tortoise Gopherus agassizii . We created heterogeneity in Mycoplasma agassizii exposure (the putative bacterial agent of a respiratory disease) by varying the duration of interactions between naturally infected and uninfected captive desert tortoises. Using qPCR, we identified new infections and compared models of transmission probability as a function of contact duration and pathogen load. We then examined the contact patterns of a wild tortoise population using proximity loggers to identify heterogeneity in contact duration. The top-ranked model predicting M. agassizii transmission included a dose term defined as the product of the number of days in proximity to an infected host and the infection level of that host. Models predicted low transmission probability for short interactions, unless the infectious host had a high load of M. agassizii : such hosts were predicted to transmit infection at higher rates with any amount of contact. We observed predominantly short-lived interactions in a free-ranging tortoise population and thus, expect transmission patterns in this population to vary considerably with the frequency and duration of high infection levels. Mean field models may misrepresent natural transmission patterns in this and other populations depending on the distribution of high-risk contact and shedding events. Rapid outbreaks in generally solitary species may result from changes to their naturally low-risk contact patterns or due to increases in the frequency of severe infections or super-shedding events &ndash; population characteristics that should be further investigated to develop effective management strategies.

Nevada

Piscivorous fish exhibit temperature-influenced binge feeding during an annual prey pulse

Understanding the limits of consumption is important for determining trophic influences on ecosystems and predator adaptations to inconsistent prey availability. Fishes have been observed to consume beyond what is sustainable (i.e. digested on a daily basis), but this phenomenon of hyperphagia (or binge-feeding) is largely overlooked. We expect hyperphagia to be a short-term (1-day) event that is facilitated by gut volume providing capacity to store consumed food during periods of high prey availability to be later digested. We define how temperature, body size and food availability influence the degree of binge-feeding by comparing field observations with laboratory experiments of bull trout ( Salvelinus confluentus ), a large freshwater piscivore that experiences highly variable prey pulses. We also simulated bull trout consumption and growth during salmon smolt outmigrations under two scenarios: 1) daily consumption being dependent upon bioenergetically sustainable rates and 2) daily consumption being dependent upon available gut volume (i.e. consumption is equal to gut volume when empty and otherwise ‘topping off’ based on sustainable digestion rates). One-day consumption by laboratory-held bull trout during the first day of feeding experiments after fasting exceeded bioenergetically sustainable rates by 12- to 87-fold at low temperatures (3 °C) and by ˜1·3-fold at 20 °C. The degree of binge-feeding by bull trout in the field was slightly reduced but largely in agreement with laboratory estimates, especially when prey availability was extremely high [during a sockeye salmon ( Oncorhynchus nerka ) smolt outmigration and at a counting fence where smolts are funnelled into high densities]. Consumption by bull trout at other settings were lower and more variable, but still regularly hyperphagic. Simulations demonstrated the ability to binge-feed increased cumulative consumption (16–32%) and cumulative growth (19–110%) relative to only feeding at bioenergetically sustainable rates during the ˜1-month smolt outmigration period. Our results indicate the ability for predators to maximize short-term consumption when prey are available can be extreme and is limited primarily by gut volume, then mediated by temperature; thus, predator–prey relationships may be more dependent upon prey availability than traditional bioenergetic models suggest. Binge-feeding has important implications for energy budgets of consumers as well as acute predation impacts on prey.

Journal of Animal Ecology

Benefits of the destinations, not costs of the journeys, shape partial migration patterns

1. The reasons that lead some animals to seasonally migrate, and others to remain in the same area year-round, are poorly understood. Associations between traits, such as body size, and migration provide clues. For example, larger species and individuals are more likely to migrate. 2. One explanation for this size bias in migration is that larger animals are capable of moving faster (movement hypothesis). However, body size is linked to many other biological processes. For instance, the energetic balances of larger animals are generally more sensitive to variation in food density because of body size effects on foraging and metabolism and this sensitivity could drive migratory decisions (forage hypothesis). 3. Identifying the primary selective forces that drive migration ultimately requires quantifying fitness impacts over the full annual migratory cycle. Here, we develop a full annual migratory cycle model from metabolic and foraging theory to compare the importance of the forage and movement hypotheses. We parameterize the model for Galapagos tortoises, which were recently discovered to be size-dependent altitudinal migrants. 4. The model predicts phenomena not included in model development including maximum body sizes, the body size at which individuals begin to migrate, and the seasonal timing of migration and these predictions generally agree with available data. Scenarios strongly support the forage hypothesis over the movement hypothesis. Furthermore, male Galapagos tortoises on Santa Cruz Island would be unable to grow to their enormous sizes without access to both highlands and lowlands. 5. Whereas recent research has focused on links between traits and the migratory phases of the migratory cycle, we find that effects of body size on the non-migratory phases are far more important determinants of the propensity to migrate. Larger animals are more sensitive to changing forage conditions than smaller animals with implications for maintenance of migration and body size in the face of environmental change.

Journal of Animal Ecology

Effects of breeder turnover and harvest on group composition and recruitment in a social carnivore

Breeder turnover can influence population growth in social carnivores through changes to group size, composition and recruitment. Studies that possess detailed group composition data that can provide insights about the effects of breeder turnover on groups have generally been conducted on species that are not subject to recurrent annual human harvest. We wanted to know how breeder turnover affects group composition and how harvest, in turn, affects breeder turnover in cooperatively breeding grey wolves ( Canis lupus Linnaeus 1758). We used noninvasive genetic sampling at wolf rendezvous sites to construct pedigrees and estimate recruitment in groups of wolves before and after harvest in Idaho, USA. Turnover of breeding females increased polygamy and potential recruits per group by providing breeding opportunities for subordinates although resultant group size was unaffected 1 year after the turnover. Breeder turnover had no effect on the number of nonbreeding helpers per group. After breeding male turnover, fewer female pups were recruited in the new males’ litters. Harvest had no effect on the frequency of breeder turnover. We found that breeder turnover led to shifts in the reproductive hierarchies within groups and the resulting changes to group composition were quite variable and depended on the sex of the breeder lost. We hypothesize that nonbreeding females direct help away from non-kin female pups to preserve future breeding opportunities for themselves. Breeder turnover had marked effects on the breeding opportunities of subordinates and the number and sex ratios of subsequent litters of pups. Seemingly subtle changes to groups, such as the loss of one individual, can greatly affect group composition, genetic content, and short-term population growth when the individual lost is a breeder.

Idaho

Life histories and conservation of long-lived reptiles, an illustration with the American crocodile (Crocodylus acutus)

Successful species conservation is dependent on adequate estimates of population dynamics, but age-specific demographics are generally lacking for many long-lived iteroparous species such as large reptiles. Accurate demographic information allows estimation of population growth rate, as well as projection of future population sizes and quantitative analyses of fitness trade-offs involved in the evolution of life-history strategies. Here, a long-term capture–recapture study was conducted from 1978 to 2014 on the American crocodile ( Crocodylus acutus ) in southern Florida. Over the study period, 7,427 hatchlings were marked and 380 individuals were recaptured for as many as 25 years. We estimated survival to be strongly age dependent with hatchlings having the lowest survival rates (16%) but increasing to nearly 90% at adulthood based on mark–recapture models. More than 5% of the female population were predicted to be reproductive by age 8 years; the age-specific proportion of reproductive females steadily increased until age 18 when more than 95% of females were predicted to be reproductive. Population growth rate, estimated from a Leslie–Lefkovitch stage-class model, showed a positive annual growth rate of 4% over the study period. Using a prospective sensitivity analysis, we revealed that the adult stage, as expected, was the most critical stage for population growth rate; however, the survival of younger crocodiles before they became reproductive also had a surprisingly high elasticity. We found that variation in age-specific fecundity has very limited impact on population growth rate in American crocodiles. We used a comparative approach to show that the original life-history strategy of American crocodiles is actually shared by other large, long-lived reptiles: while adult survival rates always have a large impact on population growth, this decreases with declining increasing growth rates, in favour of a higher elasticity of the juvenile stage. Crocodiles, as a long-lived and highly fecund species, deviate from the usual association of life histories of “slow” species. Current management practices are focused on nests and hatchling survival; however, protection efforts that extend to juvenile crocodiles would be most effective for conservation of the species, especially in an ever-developing landscape.

Journal of Animal Ecology

Spatiotemporal heterogeneity in prey abundance and vulnerability shapes the foraging tactics of an omnivore

Prey abundance and prey vulnerability vary across space and time, but we know little about how they mediate predator–prey interactions and predator foraging tactics. To evaluate the interplay between prey abundance, prey vulnerability and predator space use, we examined patterns of black bear ( Ursus americanus ) predation of caribou ( Rangifer tarandus ) neonates in Newfoundland, Canada using data from 317 collared individuals (9 bears, 34 adult female caribou, 274 caribou calves). During the caribou calving season, we predicted that landscape features would influence calf vulnerability to bear predation, and that bears would actively hunt calves by selecting areas associated with increased calf vulnerability. Further, we hypothesized that bears would dynamically adjust their foraging tactics in response to spatiotemporal changes in calf abundance and vulnerability (collectively, calf availability). Accordingly, we expected bears to actively hunt calves when they were most abundant and vulnerable, but switch to foraging on other resources as calf availability declined. As predicted, landscape heterogeneity influenced risk of mortality, and bears displayed the strongest selection for areas where they were most likely to kill calves, which suggested they were actively hunting caribou. Initially, the per‐capita rate at which bears killed calves followed a type‐I functional response, but as the calving season progressed and calf vulnerability declined, kill rates dissociated from calf abundance. In support of our hypothesis, bears adjusted their foraging tactics when they were less efficient at catching calves, highlighting the influence that predation phenology may have on predator space use. Contrary to our expectations, however, bears appeared to continue to hunt caribou as calf availability declined, but switched from a tactic of selecting areas of increased calf vulnerability to a tactic that maximized encounter rates with calves. Our results reveal that generalist predators can dynamically adjust their foraging tactics over short time‐scales in response to changing prey abundance and vulnerability. Further, they demonstrate the utility of integrating temporal dynamics of prey availability into investigations of predator–prey interactions, and move towards a mechanistic understanding of the dynamic foraging tactics of a large omnivore.

Journal of Animal Ecology

Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition

1. Although interspecific competition plays a principle role in shaping species behaviour and demography, little is known about the population-level outcomes of competition between large carnivores, and the mechanisms that facilitate coexistence. 2. We conducted a multi-landscape analysis of two widely distributed, threatened large carnivore competitors to offer insight into coexistence strategies and assist with species-level conservation. 3. We evaluated how interference competition affects occupancy, temporal activity and population density of a dominant competitor, the lion (Panthera leo), and its subordinate competitor, the leopard (Panthera pardus). We collected camera-trap data over three years in ten study sites covering 5,070 km2. We used multispecies occupancy modelling to assess spatial responses in varying environmental and prey conditions and competitor presence, and examined temporal overlap and the relationship between lion and leopard densities across sites and years. 4. Results showed that both lion and leopard occupancy was independent of – rather than conditional on – their competitor’s presence across all environmental covariates. Marginal occupancy probability for leopard was higher in areas with more bushy, ‘hideable’ habitat, human (tourist) activity and topographic ruggedness, whereas lion occupancy decreased with increasing hideable habitat and increased with higher abundance of very large prey. Temporal overlap was high between carnivores and there was no significant relationship between species densities. 4. Lions pose a threat to the survival of individual leopards, but they exerted no tractable influence on leopard spatial or temporal dynamics. Furthermore, lions did not appear to suppress leopard populations, suggesting that intraguild competitors can coexist in the same areas without population decline. Aligned conservation strategies that promote functioning ecosystems, rather than target individual species, are therefore suggested to achieve cost- and space-effective conservation.

Journal of Animal Ecology