Partitioning Detectability Components in Populations Subject to Within-Season Temporary Emigration Using Binomial Mixture Models

Detectability of individual animals is highly variable and nearly always < 1; imperfect detection must be accounted for to reliably estimate population sizes and trends. Hierarchical models can simultaneously estimate abundance and effective detection probability, but there are several different mechanisms that cause variation in detectability. Neglecting temporary emigration can lead to biased population estimates because availability and conditional detection probability are confounded. In this study, we extend previous hierarchical binomial mixture models to account for multiple sources of variation in detectability. The state process of the hierarchical model describes ecological mechanisms that generate spatial and temporal patterns in abundance, while the observation model accounts for the imperfect nature of counting individuals due to temporary emigration and false absences. We illustrate our model’s potential advantages, including the allowance of temporary emigration between sampling periods, with a case study of southern red-backed salamanders Plethodon serratus. We fit our model and a standard binomial mixture model to counts of terrestrial salamanders surveyed at 40 sites during 3–5 surveys each spring and fall 2010–2012. Our models generated similar parameter estimates to standard binomial mixture models. Aspect was the best predictor of salamander abundance in our case study; abundance increased as aspect became more northeasterly. Increased time-since-rainfall strongly decreased salamander surface activity (i.e. availability for sampling), while higher amounts of woody cover objects and rocks increased conditional detection probability (i.e. probability of capture, given an animal is exposed to sampling). By explicitly accounting for both components of detectability, we increased congruence between our statistical modeling and our ecological understanding of the system. We stress the importance of choosing survey locations and protocols that maximize species availability and conditional detection probability to increase population parameter estimate reliability.     

Estimating survival of a streamside salamander: importance of temporary emigration,capture response,and location

Estimating survival for highly secretive aquatic animals, such as stream salamanders, presents numerous challenges. Salamanders often spend a considerable time in refugia where they are difficult to capture. Few studies have calculated vital rates for stream salamanders, yet the need is substantial as they are threatened by a wide range of land-use stressors, especially urban development. In this study, we used 34 months of continuous field samples collected at an urban and undisturbed stream and robust design mark-recapture analysis to evaluate the importance of temporary emigration, capture response, and location on survival estimates of the salamander Desmognathus fuscus. We constructed a set of candidate models incorporating combinations of time- and location-varying capture and recapture probabilities, capture responses, temporary emigration, and survival estimates and ranked models using Akaike’s Information Criterion. We found strong support for month-specific capture probabilities, recapture probabilities, temporary emigration and a negative behavioral response to capture in the majority of months. We found no support for variation in capture probabilities, recapture probabilities, and temporary emigration between locations. However, we found that location strongly influenced survival estimates. Specifically, survival estimates were significantly higher at the undisturbed site than at the urban site. Our results emphasize the importance of estimating capture probabilities, recapture probabilities, capture response, and temporary emigration when evaluating survival in highly secretive aquatic animals. Failure to account for these population parameters will likely yield biased estimates of survival in freshwater animal populations.     

Road effects on demographic traits of small mammal populations

Recent studies have highlighted the positive effects of road verges on the abundance of small mammals. However, most of these studies occurred in intensively grazed or cultivated areas, where verges were the last remnants of suitable habitats, which could mask the true effects of roads on population traits. We analysed the effects of roads on small mammal populations living in a well-preserved Mediterranean forest. We used the wood mouse (Apodemus sylvaticus) as a model of forest-dwelling small mammals that probably are among the species most affected by road clearings. Our study compared populations in similar habitat areas with and without road influence. We assessed abundance, survival and temporary emigration using extended Pollock’s robust design capture-recapture models. Moreover, we analysed population turnover, sex ratio, age structure and body condition. We found that wood mouse abundance and body condition were lower at the road bisected area, whereas the remaining population traits were similar. This suggests that the reduced habitat availability and quality due to the physical presence of the road and verge vegetation clearing are the main drivers of demographic differences in wood mouse populations between areas. Nevertheless, our results also suggest that in high-quality habitats surrounding national roads, wood mouse populations present similar dynamics to others living in undisturbed areas, despite the decrease in abundance and body condition. Overall, the often-reported increased small mammal abundance in road surroundings should not be generalized independently of habitat quality or to other population traits.     

Loglinear models for the robust design in mark-recapture experiments

The robust design is a method for implementing a mark-recapture experiment featuring a nested sampling structure. The first level consists of primary sampling sessions; the population experiences mortality and immigration between primary sessions so that open population models apply at this level. The second level of sampling has a short mark-recapture study within each primary session. Closed population models are used at this stage to estimate the animal abundance at each primary session. This article suggests a loglinear technique to fit the robust design. Loglinear models for the analysis of mark-recapture data from closed and open populations are first reviewed. These two types of models are then combined to analyze the data from a robust design. The proposed loglinear approach to the robust design allows incorporating parameters for a heterogeneity in the capture probabilities of the units within each primary session. Temporary emigration out of the study area can also be accounted for in the loglinear framework. The analysis is relatively simple; it relies on a large Poisson regression with the vector of frequencies of the capture histories as dependent variable. An example concerned with the estimation of abundance and survival of the red-back vole in an area of southeastern Québec is presented.     

Analysing the effect of movement on local survival: a new method with an application to a spatially structured population of the arboreal gecko Gehyra variegata

Mortality during movement between habitat patches is the most obvious cost of dispersal, but rarely it has been demonstrated empirically. An approach is presented, which uses capture–mark–recapture data of an arboreal gecko species to determine the effect of individual movement on local survival in a spatially structured population. Because capture–mark–recapture data are widely available for a range of animal species, it should be possible to extend their application to other species. The method is based on the assumption that the tendency to be a territorial animal or to be a floating animal is fixed during the study period. The advantage of our approach is that only one additional parameter has to be estimated for describing movement risks. We further tested the power of our approach to detect an association of movement and mortality with simulated capture histories. The study revealed a strong negative effect of movement on local survival. Hence, animals that moved more often between trees had a lower survival rate. Interestingly, the mean movement rate for males was significantly higher than for females, which should lead to a biased sex ratio towards females in the population. As there was an even sex ratio in the population, we discuss not mutually exclusive explanations for this finding like differences in emigration rates between sexes, differences in survival rates between sexes, or a skewed sex ratio in offspring.     

El Niño Southern Oscillation influences the abundance and movements of a marine top predator in coastal waters

Large‐scale climate modes such as El Niño Southern Oscillation (ENSO) influence population dynamics in many species, including marine top predators. However, few quantitative studies have investigated the influence of large‐scale variability on resident marine top predator populations. We examined the effect of climate variability on the abundance and temporary emigration of a resident bottlenose dolphin (Tursiops aduncus) population off Bunbury, Western Australia (WA). This population has been studied intensively over six consecutive years (2007–2013), yielding a robust dataset that captures seasonal variations in both abundance and movement patterns. In WA, ENSO affects the strength of the Leeuwin Current (LC), the dominant oceanographic feature in the region. The strength and variability of the LC affects marine ecosystems and distribution of top predator prey. We investigated the relationship between dolphin abundance and ENSO, Southern Annular Mode, austral season, rainfall, sea surface salinity and sea surface temperature (SST). Linear models indicated that dolphin abundance was significantly affected by ENSO, and that the magnitude of the effect was dependent upon season. Dolphin abundance was lowest during winter 2009, when dolphins had high temporary emigration rates out of the study area. This coincided with the single El Niño event that occurred throughout the study period. Coupled with this event, there was a negative anomaly in SST and an above average rainfall. These conditions may have affected the distribution of dolphin prey, resulting in the temporary emigration of dolphins out of the study area in search of adequate prey. This study demonstrated the local effects of large‐scale climatic variations on the short‐term response of a resident, coastal delphinid species. With a projected global increase in frequency and intensity of extreme climatic events, resident marine top predators may not only have to contend with increasing coastal anthropogenic activities, but also have to adapt to large‐scale climatic changes.     

Open removal models with temporary emigration and population dynamics to inform invasive animal management

Removal sampling data are the primary source of monitoring information for many populations (e.g., invasive species, fisheries). Population dynamics, temporary emigration, and imperfect detection are common sources of variation in monitoring data and are key parameters for informing management. We developed two open robust‐design removal models for simultaneously modeling population dynamics, temporary emigration, and imperfect detection: a random walk linear trend model (estimable without ancillary information), and a 2‐age class informed population model (InfoPM, closely related to integrated population models) that incorporated prior information for age‐structured vital rates and relative juvenile availability. We applied both models to multiyear, removal trapping time‐series of a large invasive lizard (Argentine black and white tegu, Salvator merianae) in three management areas of South Florida to evaluate the effectiveness of management programs. Although estimates of the two models were similar, the InfoPMs generally returned more precise estimates, partitioned dynamics into births, deaths, net migration, and provided a decision support tool to predict population dynamics under different effort scenarios while accounting for uncertainty. Trends in tegu superpopulation abundance estimates were increasing in two management areas despite generally high removal rates. However, tegu abundance appeared to decline in the Core management area, where trapping density was the highest and immigration the lowest. Finally, comparing abundance predictions of no‐removal scenarios to those estimated in each management area suggested significant population reductions due to management. These results suggest that local tegu population control via systematic trapping may be feasible with high enough trap density and limited immigration; and highlights the value of these trapping programs. We provided the first estimates of tegu abundance, capture probabilities, and population dynamics, which is critical for effective management. Furthermore, our models are applicable to a wide range of monitoring programs (e.g., carcass recovery or removal point‐counts).     

Use of the Robust Design to Estimate Seasonal Abundance and Demographic Parameters of a Coastal Bottlenose Dolphin (Tursiops aduncus) Population

As delphinid populations become increasingly exposed to human activities we rely on our capacity to produce accurate abundance estimates upon which to base management decisions. This study applied mark–recapture methods following the Robust Design to estimate abundance, demographic parameters, and temporary emigration rates of an Indo-Pacific bottlenose dolphin (Tursiops aduncus) population off Bunbury, Western Australia. Boat-based photo-identification surveys were conducted year-round over three consecutive years along pre-determined transect lines to create a consistent sampling effort throughout the study period and area. The best fitting capture–recapture model showed a population with a seasonal Markovian temporary emigration with time varying survival and capture probabilities. Abundance estimates were seasonally dependent with consistently lower numbers obtained during winter and higher during summer and autumn across the three-year study period. Specifically, abundance estimates for all adults and juveniles (combined) varied from a low of 63 (95% CI 59 to 73) in winter of 2007 to a high of 139 (95% CI 134 to148) in autumn of 2009. Temporary emigration rates (γ'') for animals absent in the previous period ranged from 0.34 to 0.97 (mean  =  0.54; ±SE 0.11) with a peak during spring. Temporary emigration rates for animals present during the previous period (γ'''') were lower, ranging from 0.00 to 0.29, with a mean of 0.16 (± SE 0.04). This model yielded a mean apparent survival estimate for juveniles and adults (combined) of 0.95 (± SE 0.02) and a capture probability from 0.07 to 0.51 with a mean of 0.30 (± SE 0.04). This study demonstrates the importance of incorporating temporary emigration to accurately estimate abundance of coastal delphinids. Temporary emigration rates were high in this study, despite the large area surveyed, indicating the challenges of sampling highly mobile animals which range over large spatial areas.     

Male-biased sex ratios in mature damselfly populations: real or artefact?

1. There is ongoing controversy about whether biased sex ratios in diploid insect populations are real or an artefact caused by different behaviours and/or different catchability of the sexes. This was tested by monitoring two field and three semi-natural populations of the damselfly Lestes sponsa. 2. Capture–mark–recapture data showed that population size estimates were about twice as large for males as for females at both field sites. Independent estimates of the sex ratios based on total numbers of males and females captured supported the male bias. 3. Males had higher recapture probabilities than females due to longer times between successive visits in females. Because the same pattern was found in the semi-natural populations, the longer intervals in females are no artefact due to their lower detectability. 4. Theoretical models show that the strong temporary emigration of females tends, if anything, to overestimate female population sizes and that the heterogeneity of recapture probabilities observed in males tends to underestimate male population sizes. Hence, behavioural differences between the sexes do not cause an artificially male-biased sex ratio. 5. Spatial data show that operational sex ratios are male biased at the pond but become female biased in the plots further away from the shoreline; however because of the decrease in densities away from the shoreline, this does not result in a global even sex ratio. 6. Spatial data, temporary emigration patterns, and independent estimates suggest strongly that the male-biased sex ratios in mature damselfly populations are real.     

A Comparison of Grizzly Bear Demographic Parameters Estimated from Non-Spatial and Spatial Open Population Capture-Recapture Models

Capture-recapture studies are frequently used to monitor the status and trends of wildlife populations. Detection histories from individual animals are used to estimate probability of detection and abundance or density. The accuracy of abundance and density estimates depends on the ability to model factors affecting detection probability. Non-spatial capture-recapture models have recently evolved into spatial capture-recapture models that directly include the effect of distances between an animal’s home range centre and trap locations on detection probability. Most studies comparing non-spatial and spatial capture-recapture biases focussed on single year models and no studies have compared the accuracy of demographic parameter estimates from open population models. We applied open population non-spatial and spatial capture-recapture models to three years of grizzly bear DNA-based data from Banff National Park and simulated data sets. The two models produced similar estimates of grizzly bear apparent survival, per capita recruitment, and population growth rates but the spatial capture-recapture models had better fit. Simulations showed that spatial capture-recapture models produced more accurate parameter estimates with better credible interval coverage than non-spatial capture-recapture models. Non-spatial capture-recapture models produced negatively biased estimates of apparent survival and positively biased estimates of per capita recruitment. The spatial capture-recapture grizzly bear population growth rates and 95% highest posterior density averaged across the three years were 0.925 (0.786–1.071) for females, 0.844 (0.703–0.975) for males, and 0.882 (0.779–0.981) for females and males combined. The non-spatial capture-recapture population growth rates were 0.894 (0.758–1.024) for females, 0.825 (0.700–0.948) for males, and 0.863 (0.771–0.957) for both sexes. The combination of low densities, low reproductive rates, and predominantly negative population growth rates suggest that Banff National Park’s population of grizzly bears requires continued conservation-oriented management actions.     

To catch or to sight? A comparison of demographic parameter estimates obtained from mark-recapture and mark-resight models

Accurate assessments of population parameters, such as survival and abundance, are critical for effective wildlife conservation. In order for wildlife managers to undertake long-term monitoring of populations, the data collection must be as cost-effective as possible. Two demographic modelling techniques commonly used are mark-recapture and mark-resight. Mark-resight can be used in conjunction with biotelemetry methods and offers a more cost effective alternative to the traditional mark-recapture models. However, there has been no empirical comparison of the demographic parameters obtained from the two modelling techniques. This study used photographs of natural markings to individually identify wobbegong sharks (Orectolobus maculatus) sighted during underwater surveys over a 2 year period, during eight distinct sampling periods, and analysed with Pollock’s robust design mark-recapture models. These estimates were then compared, using z tests, with Poisson-lognormal mark-resight models that used resightings of sharks previously tagged with telemetry transmitters, and the telemetry data to calculate the number of marked animals present in each sampling period. Sharks were categorised into four groups according to their sex and age-class (adult/juvenile). The results indicated that there was a high degree of transience in the population, with 62 % of sharks only being sighted in one sampling period. Based on normalized Akaike weights, there was no single ‘best’ model for either type of modelling technique and model averaging was used to determine the demographic estimates. Both models showed higher abundance of wobbegongs in the austral spring and summer seasons, however, the models produced statistically different results for five of the eight sampling periods. The mark-recapture model estimated apparent survival between 78 and 95 %, whereas the mark-resight models estimated it between 48 and 97 %. Crucially, there was no statistical difference between the survival estimates from corresponding sex/age-class. The temporary emigration parameters differed substantially between the model types. The mark-recapture model showed support for Markovian movement, whereas the mark-resight supported random emigration. The timing of the tagging events likely biased the abundance and temporary emigration parameters estimated by mark-resight models and must be taken into consideration when designing a mark-resight study. Despite this, this study shows that robust demographic estimates, that are comparable to labour intensive mark-recapture estimates, can still be obtained using mark-resight methods. Given the substantial increase in biotelemetry studies of medium and large sized vertebrates, mark-resight models may play an important future role in estimating demographic parameters.     

基于标记-重捕模型开展野生动物红外相机种群监测的方法及案例

红外相机技术的广泛应用推动了动物种群生态学研究方法的发展和革新, 特别是基于标记-重捕模型框架通过非损伤取样方式对物种数量和密度等种群参数的可靠估计, 为保护濒危物种和评估保护成效提供了有力的科学依据。对于身体上具有独特天然标记的动物(如多数猫科动物), 可依据红外相机拍摄身体上的独特斑点或条纹鉴别个体, 再运用标记-重捕模型, 估计动物种群数量、密度等参数。本文概述了标记-重捕模型的基本原理、特点以及国内外的应用, 特别是近年来发展出的空间标记-重捕模型。总结了从相机布设到数据分析的具体流程、操作原则, 并以青城山家猫为实例, 展示了应用红外相机数据通过空间标记-重捕模型估计种群密度和数量的基本步骤。最后展望了该模型在种群动态、景观廊道设计、资源选择等方面的应用和发展趋势。     

Abundance Estimation With a Transient Model Under the Robust Design

Abstract: A common situation in capture-mark-recapture (CMR) studies on birds and other organisms is to capture individuals not belonging to the studied population only present during the short time of the capture session. Presence of such transient individuals affects demographic parameter estimation from CMR data. Methods exist to reduce biases on survival estimates in the presence of transients and have been shown to be particularly efficient within the Robust Design framework (several secondary capture sessions within a short time interval during which the studied population can be assumed closed). We present a new model to estimate population size accounting for transients. We first used simulated data to show that the method reduces positive biases due to transients. In a second step, we applied the method to a real CMR dataset on a reed warbler (Acrocephalus scirpaceus) population. Population size estimates are reduced by up to 50% when correcting for the presence of transients. Many field studies on managed animal populations use capture-recapture methodology to obtain crucial parameters of the focal population demography. The resulting data sets are used either to estimate population size ignoring the presence of transients, or to estimate vital rates, accounting for transients but overlooking abundance estimation. Our method conciliates these 2 approaches.     

Seasonal abundance and adult survival of bottlenose dolphins (Tursiops truncatus) in a community that cooperatively forages with fishermen in southern Brazil

A subgroup of a population of Tursiops truncatus in southern Brazil is known for a cooperative behavior with artisanal fishermen whereby the dolphins shoal fish towards net‐casting fishermen. Combining photo‐identification data collected between September 2007 and 2009 with mark‐recapture and Pollock's robust design models, we assessed abundance within seasons and survival and temporary emigration rates of dolphins between seasons. We also reanalyzed a previous data set collected during 1989–1991, and Cormack‐Jolly‐Seber models were applied to estimate survival rates for each of the study periods. The abundance of marked “cooperative” dolphins varied between seasons from 18 (CI: 17–24) to 21 (CI: 20–24). The total abundance varied from 59 in the winter of 2008 (CI: 49–72) to 50 in the autumn of 2009 (CI: 40–62). The annual adult survival was estimated to be 0.917 (CI: 0.876–0.961), close to that estimated from data collected in the 1990s (0.941; CI: 0.888–0.998). The emigration probability was low (0.031; CI: 0.011–0.084) and different capture probabilities between the “cooperative” and “noncooperative” dolphins indicated a degree of behavioral segregation. The precision of our estimates is likely to provide sufficient power to detect population change, but we recommend a precautionary management approach to protect this vulnerable dolphin community and its unique cooperative feeding tradition.     

Quantitative genetic study on sexual difference in emigration behavior of Drosophila melanogaster in a natural population.

A quantitative genetic analysis was conducted on emigration response behavior using 140 second chromosome lines of Drosophila melanogaster. Fourteen sets of 5 x 5 partial diallel cross experiments were made in the parental generation. The emigration activity per batch of 50 male and 50 female F1 progeny was scored with Sakai's population system. Sexual difference did not appear in the emigration activity in these experiments. A significant genotype x sex x set interaction was detected. The genetic variance components of emigration activity differed between sexes: In males, additive genetic variance of emigration activity was 0.0497 +/- 0.0092 and dominance variance, 0.0018 +/- 0.0046; in females, additive, 0.0373 +/- 0.0076 and dominance, 0.0169 +/- 0.0044. Additive genetic correlation between sexes for the emigration activity was 0.685 +/- 0.150, deviating significantly from unity. These results suggested that the genes affecting emigration activity would operate differently between sexes of D. melanogaster in natural populations.     

Spatially explicit maximum likelihood methods for capture-recapture studies

Live-trapping capture-recapture studies of animal populations with fixed trap locations inevitably have a spatial component: animals close to traps are more likely to be caught than those far away. This is not addressed in conventional closed-population estimates of abundance and without the spatial component, rigorous estimates of density cannot be obtained. We propose new, flexible capture-recapture models that use the capture locations to estimate animal locations and spatially referenced capture probability. The models are likelihood-based and hence allow use of Akaike's information criterion or other likelihood-based methods of model selection. Density is an explicit parameter, and the evaluation of its dependence on spatial or temporal covariates is therefore straightforward. Additional (nonspatial) variation in capture probability may be modeled as in conventional capture-recapture. The method is tested by simulation, using a model in which capture probability depends only on location relative to traps. Point estimators are found to be unbiased and standard error estimators almost unbiased. The method is used to estimate the density of Red-eyed Vireos (Vireo olivaceus) from mist-netting data from the Patuxent Research Refuge, Maryland, U.S.A. Estimates agree well with those from an existing spatially explicit method based on inverse prediction. A variety of additional spatially explicit models are fitted; these include models with temporal stratification, behavioral response, and heterogeneous animal home ranges.     

Evaluating hypotheses about dispersal in a vulnerable butterfly

Sound management of species requires reliable estimates of dispersal. Indeed, dispersal of individuals among local populations is a key factor in the biology and persistence of local populations and metapopulations. Here, the small-scale dispersal pattern of a vulnerable species, the endemic Sardinian chalk hill blue butterfly, was studied by applying capture–recapture multistate models and a model selection based on AIC values. Model parameters were survival, capture and movement probabilities. The model selection showed that (a) survival probability of individuals varied between sexes, (b) capture probability varied between sexes and among patches, and (c) movement probability varied with direction. The probability of movement among adjacent local populations was generally low and ranged from 0.009 to 0.212. Movement probabilities were subsequently modeled using data on interpatch distance and donor patch population size or area. The ultrastructural biology-based models turned out to be the most appropriate models for inference, showing that dispersal decreases with increasing interpatch distance and increasing donor patch population size or area, and suggesting that butterfly dispersal is affected by patch geometry and the presence of conspecifics. The application of multistate models, the model selection approach, and ultrastructural modeling allowed testing the validity of some general hypotheses related to dispersal in metapopulations and helped elucidate the butterfly small-scale dispersal pattern.     

Ecological and methodological factors affecting detectability and population estimation in elusive species

Although mark-recapture methods are among the most powerful tools for monitoring wildlife populations, the secretive nature of some species requires a comprehensive understanding of the factors that affect capture probability to maximize accuracy and precision of population parameter estimates (e.g., population size and survivorship). Here, we used aquatic snakes as a case study in applying rigorous mark-recapture methods to estimate population parameters for secretive species. Specifically, we used intensive field sampling and robust design mark-recapture analyses in Program MARK to test specific hypotheses about ecological and methodological factors influencing detectability of two species of secretive aquatic snakes, the banded watersnake (Nerodia fasciata), and the black swamp snake (Seminatrix pygaea). We constructed a candidate set of a priori mark-recapture models incorporating various combinations of time- and sex-varying capture and recapture probabilities, behavioral responses to traps (i.e., trap-happiness or trap-shyness), and temporary emigration, and we ranked models for each species using Akaike's Information Criterion. For both banded watersnakes and black swamp snakes we found strong support for time-varying capture and recapture probabilities and strong trap-happy responses, factors that can bias population estimation if not accommodated in the models. We also found evidence of sex-dependent temporary emigration in black swamp snakes. Our study is among the first comprehensive assessments of factors affecting detectability in snakes and provides a framework for studies aimed at monitoring populations of other secretive species. © 2010 The Wildlife Society.     

Open capture-recapture models with heterogeneity: I. Cormack-Jolly-Seber model

In open population capture-recapture studies, it is usually assumed that similar animals (e.g., of the same sex and age group) have similar survival rates and capture probabilities. These assumptions are generally perceived to be an oversimplification, and they can lead to incorrect model selection and biased parameter estimates. Allowing for individual variability in survival and capture probabilities among apparently similar animals is now becoming possible, due to advances in closed population models and improved computing power. This article presents a flexible framework of likelihood-based models which allow for individual heterogeneity in survival and capture rates. Heterogeneity is modeled using finite mixtures, which have enough flexibility of distribution shape to accommodate a wide variety of different patterns of individual variation. The models condition on the first capture of each animal, and include as a special case the Cormack-Jolly-Seber model. Model selection is done either using Akaike's information criterion or by likelihood ratio tests, making available checks of different influences on survival rates. Bias in parameter estimates is reduced by including individual heterogeneity. Model selection and bias reduction are important in population studies and for making informed management decisions.     

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

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