The estimation of population size remains one of the primary goals and challenges in ecology and provides a basis for debate and policy in wildlife management. Despite the development of efficient ...noninvasive sampling methods and robust statistical tools to estimate abundance, the maintenance of field sampling is still subject to economic and logistic constraints. These can result in intentional or unintentional interruptions in sampling and cause gaps in data time series, posing a challenge to abundance estimation, and ultimately conservation and management decisions. We applied an open population spatial capture–recapture (OPSCR) model to simulations and a real‐life case study to test the reliability of abundance inference to interruptions in data collection. Using individual detections occurring over consecutive sampling occasions, OPSCR models allow the estimation of abundance while accounting for lack of demographic and geographic closure between occasions. First, we simulated sampling data with interruptions in field sampling of different lengths and timing and checked the performance of an OPSCR model in deriving abundance for species with slow and intermediate life‐history strategies. Next, we introduced artificial sampling interruptions of various magnitudes and timing to a five‐year noninvasive monitoring data set of wolverines (Gulo gulo) in Norway and quantified the consequences for OPSCR model predictions. Inferences from OPSCR models were reliable even with temporal interruptions in monitoring. Interruption did not cause systematic bias, but increased uncertainty. Interruptions occurring at occasions near the beginning and the end of the sampling period caused higher uncertainty. The loss in precision was more severe for species with a faster life‐history strategy. OPSCR allows monitoring studies to provide contiguous abundance estimates to managers, stakeholders, and policy makers even when data are noncontiguous. OPSCR models do not only help cope with unintentional interruptions during sampling but also offer opportunities for using intentional sampling interruptions during the design of cost‐effective population surveys.
Group living is assumed to benefit prey by reducing predation risk due to dilution and detection effects. This increased safety could be exchanged against a reduced individual vigilance leading to ...increased foraging and fitness until costs of intra‐group competition offset this benefit. However, very few studies have been able to directly test the relationship between individual fitness and group size in non‐cooperative breeding species. Therefore, evidence that dilution and detection effects do indeed translate into increased fitness under predation risk remains mostly theoretical. Here, we aimed at empirically testing whether group size influences beneficial and costly behaviors, and ultimately the survival of individuals, in a prey species facing heavy predation pressure. We focused on plains zebra Equus quagga of Hwange National Park in Zimbabwe, a high lion density area. We collected behavioral data (i.e., individual vigilance and foraging, collective vigilance, and agonistic interactions) that we used as proxies to investigate the costs and benefits of grouping in this species. We then fitted multistate capture‐mark‐recapture and Cox proportional‐hazards models to estimate apparent survival of adults and young zebras, respectively, and assess whether a demographic signal of group size could be found. Our results show that collective vigilance increases with group size, a prerequisite of the detection effect, but individual vigilance does not consistently decrease with group size, which suggests that there is no group size effect on individual vigilance. Agonistic interactions were rare. These results suggest that there are no foraging costs of grouping here, and even some benefits such as a better collective vigilance. However, any benefits of being in larger groups did not translate into increased survival, as we found no effect of group size on the apparent survival of individuals. Since dilution and detection effects do not matter behaviorally and demographically in these plains zebras living under high predation risk, these results challenge our understanding of how predation shapes animal sociality: fitness benefits of dilution or detection should not be assumed.
This study describes two longitudinal serological surveys of European Bat Lyssavirus type 1 (EBLV-1) antibodies in serotine bat (Eptesicus serotinus) maternity colonies located in the North-East of ...France. This species is currently considered as the main EBLV-1 reservoir. Multievent capture-recapture models were used to determine the factors influencing bat rabies transmission as this method accounts for imperfect detection and uncertainty in disease states. Considering the period of study, analyses revealed that survival and recapture probabilities were not affected by the serological status of individuals, confirming the capacity of bats to be exposed to lyssaviruses without dying. Five bats have been found with EBLV-1 RNA in the saliva at the start of the study, suggesting they were caught during virus excretion period. Among these bats, one was interestingly recaptured one year later and harbored a seropositive status. Along the survey, some others bats have been observed to both seroconvert (i.e. move from a negative to a positive serological status) and serorevert (i.e. move from a positive to a negative serological status). Peak of seroprevalence reached 34% and 70% in site A and B respectively. On one of the 2 sites, global decrease of seroprevalence was observed all along the study period nuanced by oscillation intervals of approximately 2-3 years supporting the oscillation infection dynamics hypothesized during a previous EBLV-1 study in a Myotis myotis colony. Seroprevalence were affected by significantly higher seroprevalence in summer than in spring. The maximum time observed between successive positive serological statuses of a bat demonstrated the potential persistence of neutralizing antibodies for at least 4 years. At last, EBLV-1 serological status transitions have been shown driven by age category with higher seroreversion frequencies in adults than in juvenile. Juveniles and female adults seemed indeed acting as distinct drivers of the rabies virus dynamics, hypothesis have been addressed but their exact role in the EBLV-1 transmission still need to be specified.
In stochastic environments, a change in a demographic parameter can influence the population growth rate directly or via a resulting impact on age structure. Stochastic elasticity of the long‐run ...stochastic growth rate λₛ to a demographic parameter offers a suitable way to measure the overall demographic response because it includes both the direct effect of changing the demographic parameter and its indirect effect through changes in the age structure. From 25 mammalian populations with contrasting life histories, we investigated how pace of life and population growth rate influence the demographic responses (measured as the relative contributions of the direct and indirect components of stochastic elasticity on λₛ). We found that in short‐lived species, the change in population structure resulting from an increase in yearling survival leads to an additional increase in λₛ, whereas in long‐lived species, the same change in population structure leads to a decrease. Short‐lived species thus display a boom‐bust life history strategy contrary to long‐lived species, for which the long lifespan dampens the demographic consequences of changing age structure. Irrespective of the species’ life history strategy, the change in population age structure resulting from an increase in adult survival leads to an additional increase in λₛ due to an increase of the proportion of mature individuals in the population. On the contrary, a change in population age structure resulting from an increase of reproductive performance leads to a decrease in λₛ that is due to the increase of the proportion of immature individuals in the population. Our comparative analysis of stochastic elasticity patterns in mammals shows the existence of different demographic responses to changes in age structure between short‐ and long‐lived species, which improves our understanding of population dynamics in variable environments in relation to the species‐specific pace of life.
1. The population dynamic and evolutionary effects of harvesting are receiving growing attention among biologists. Cause-specific estimates of mortality are necessary to determine and compare the ...magnitude and selectivity of hunting and other types of mortalities. In addition to the logistic and financial constraints on longitudinal studies, they are complicated by the fact that nonhunting mortality in managed populations usually consists of a mix of natural and human-caused factors. 2. We used multistate capture-recapture (MCR) models to estimate cause-specific survival of brown bears (Ursus arctos) in two subpopulations in Sweden over a 23-year period. In our analysis, we distinguished between legal hunting and other sources of mortality, such as intraspecific predation, accidents, poaching, and damage control removals. We also tested whether a strong increase in harvest quotas after 1997 in one of the subpopulations affected vulnerability to legal hunting. 3. Although only a fraction of mortalities other than legal hunting could be considered natural, this group of causes showed a general pattern of demographic selectivity expected from natural mortality regimes in populations of long-lived species, namely greater vulnerability of young animals. On the other hand, demographic effects on hunting vulnerability were weak and inconsistem. Out findings support the assumption that hunting and other mortalities were additive. 4. As expected, an increase in hunting pressure coincided with a correspondingly large increase in vulnerability to hunting in the affected subpopulation. Because even unbaised harvest can lead to selective pressures on life-history traits, such as size at primiparity, increasing harvest quotas may not only affect population growth directly, but could also alter optimal life-history strategies in brown bears and other carnivores. 5. Legal hunting is the most conveniently assessed and the most easily managed cause of mortality in many wild populations of large mammals. Although legal hunting is the single-most important cause of mortality for brown bears in Sweden, the combined mortality from other causes is of considerable magnitude and additionally shows greater selectivity in terms of sex and age than legal hunting. Therefore, its role in population dynamics and evolution should not be underestimated.
In species providing extended parental care, one or both parents care for altricial young over a period including more than one breeding season. We expect large parental investment and long‐term ...dependency within family units to cause high variability in life trajectories among individuals with complex consequences at the population level. So far, models for estimating demographic parameters in free‐ranging animal populations mostly ignore extended parental care, thereby limiting our understanding of its consequences on parents and offspring life histories.
We designed a capture–recapture multievent model for studying the demography of species providing extended parental care. It handles statistical multiple‐year dependency among individual demographic parameters grouped within family units, variable litter size, and uncertainty on the timing at offspring independence. It allows for the evaluation of trade‐offs among demographic parameters, the influence of past reproductive history on the caring parent's survival status, breeding probability, and litter size probability, while accounting for imperfect detection of family units. We assess the model performance using simulated data and illustrate its use with a long‐term dataset collected on the Svalbard polar bears (Ursus maritimus).
Our model performed well in terms of bias and mean square error and in estimating demographic parameters in all simulated scenarios, both when offspring departure probability from the family unit occurred at a constant rate or varied during the field season depending on the date of capture. For the polar bear case study, we provide estimates of adult and dependent offspring survival rates, breeding probability, and litter size probability. Results showed that the outcome of the previous reproduction influenced breeding probability.
Overall, our results show the importance of accounting for i) the multiple‐year statistical dependency within family units, ii) uncertainty on the timing at offspring independence, and iii) past reproductive history of the caring parent. If ignored, estimates obtained for breeding probability, litter size, and survival can be biased. This is of interest in terms of conservation because species providing extended parental care are often long‐living mammals vulnerable or threatened with extinction.
We developed a capture–recapture model for studying the demography of species providing extended parental care. Our model handles statistical dependency among individual demographic parameters and life histories within family units until offspring independence, and include variable litter size, age at offspring independence and breeding frequency. It also includes the influence of past reproductive history on the caring parent's survival status and current reproduction, while accounting for imperfect detection of family units. We assess the model performances using simulated data, and illustrate its use with a long‐term dataset collected on the Svalbard polar bears (Ursus maritimus).
Addressing evolutionary questions in the wild remains a challenge. It is best done by monitoring organisms from birth to death, which is very difficult in part because individuals may or may not be ...resighted or recaptured. Although the issue of uncertain detection has long been acknowledged in ecology and conservation biology, in evolutionary studies of wild populations it is often assumed that detectability is perfect. We argue that this assumption may lead to flawed inference. We demonstrate that the form of natural selection acting on body mass of sociable weavers is altered and that the rate of senescence of roe deer is underestimated when not accounting for a value of detectability that is less than one. Because mark‐recapture models provide an explicit way to integrate and reliably model the detection process, we strongly recommend their use to address questions in evolutionary biology.
Data collected to inform time variations in natural population size are tainted by sampling error. Ignoring sampling error in population dynamics models induces bias in parameter estimators, e.g., ...density-dependence. In particular, when sampling errors are independent among populations, the classical estimator of the synchrony strength (zero-lag correlation) is biased downward. However, this bias is rarely taken into account in synchrony studies although it may lead to overemphasizing the role of intrinsic factors (e.g., dispersal) with respect to extrinsic factors (the Moran effect) in generating population synchrony as well as to underestimating the extinction risk of a metapopulation.
The aim of this paper was first to illustrate the extent of the bias that can be encountered in empirical studies when sampling error is neglected. Second, we presented a space-state modelling approach that explicitly accounts for sampling error when quantifying population synchrony. Third, we exemplify our approach with datasets for which sampling variance (i) has been previously estimated, and (ii) has to be jointly estimated with population synchrony. Finally, we compared our results to those of a standard approach neglecting sampling variance. We showed that ignoring sampling variance can mask a synchrony pattern whatever its true value and that the common practice of averaging few replicates of population size estimates poorly performed at decreasing the bias of the classical estimator of the synchrony strength.
The state-space model used in this study provides a flexible way of accurately quantifying the strength of synchrony patterns from most population size data encountered in field studies, including over-dispersed count data. We provided a user-friendly R-program and a tutorial example to encourage further studies aiming at quantifying the strength of population synchrony to account for uncertainty in population size estimates.
In an analysis of capture‐recapture data, the identification of a model that fits is a critical step. For the multisite (also called multistate) models used to analyze data gathered at several sites, ...no reliable test for assessing fit is currently available. We propose a test for the JMV model, a simple generalization of the Arnason‐Schwarz (AS) model, in the form of interpretable contingency tables. For the AS model, we suggest complementing the test for the JMV model with a likelihood ratio test of AS vs. JMV. The examination of an example leads us to propose further a partitioning that emphasizes the role of the memory model of Brownie et al. (1993 Biometrics49, 1173–1187) as a biologically more plausible alternative to the AS model.
Breeding dispersal is a key process of population structure and dynamics and is often triggered by an individual's breeding failure. In both colonial and territorial birds, reproductive success of ...conspecifics (RSc) can also lead individuals to change breeding sites after a failure on a site. Yet, few studies have simultaneously investigated the independent contribution of individual reproductive success (RSi) and of RSc on dispersal decision. Here, we develop a modeling framework to disentangle the effects of RSi and RSc on demographic parameters, while accounting for imperfect individual detection and other confounding factors such as age or dispersal behavior in the previous year. Using a 10‐year capture–recapture dataset composed of 1,595 banded tree swallows, we assessed the effects of nonmanipulated RSi and RSc on female breeding dispersal in this semicolonial passerine. Dispersal was strongly driven by RSi, but not by RSc. Unsuccessful females were 9.5–2.5 times more likely to disperse than successful ones, depending if they had dispersed or not in the previous year, respectively. Unsuccessful females were also three times less likely to be detected than successful ones. Contrary to theoretical and empirical studies, RSc did not drive the decision to disperse but influenced the selection of the following breeding site once dispersal had been initiated. Because detection of individuals was driven by RSi, which was positively correlated to RSc, assuming a perfect detection as in previous studies may have lead us to conclude that RSc affected dispersal patterns, yet our approach corrected for this bias. Overall, our results suggest that the value and use of RSc as public information to guide dispersal decisions are likely dictated by multiple ecological determinants, such as landscape structure and extent, if this cue is indeed used.
Currently, few studies have simultaneously investigated the contribution of individual reproductive success (RSi) and reproductive success of conspecifics (RSc). on dispersal decision, contrary to settlement decision after a dispersal event. Here, we develop a new analytical framework to disentangle the effects of RSi and RSc on demographic parameters, while accounting for imperfect individual detection and other confounding factors in tree swallow. Our results contrast with previous studies by the fact that RSc does not impact the decision to disperse and we discuss these results and the potential bias caused by the use of classical regression analyses.
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