Individual heterogeneity in recapture probability and survival estimates in cheetah

Accurate estimates of demographic parameters are key for understanding and predicting population dynamics and for providing insights for effective wildlife management. Up until recently, no suitable methodology has been available to estimate survival probabilities of species with asynchronous reproduction and a high level of individual variation in capture probabilities. The present work develops a capture-mark-recapture model for cheetahs in the Serengeti National Park, Tanzania, which (a) deals with continuous reproduction, (b) takes into account the high level of individual heterogeneity in capture probabilities and (c) is spatially explicit. Results show that (1) our approach, which is an extensive modification of the Cormack-Jolly-Seber model, provides a lower female adult survival estimate and a higher male adolescent survival estimate than previous approaches to estimate cheetah survival in the area, (2) using sighting location alone is not sufficient to capture the individual variation in resighting probabilities for both sexes, and (3) precision in estimated survival probabilities is generally increased. Species which are individually recognizable, wide-ranging and/or where individuals differ substantially in sightability are particularly appropriate to our modelling approach, and our methodology would thus be appropriate for a wide number of species to provide more accurate estimates of survival.     

Explaining long-distance dispersal: effects of dispersal distance on survival and growth in a stream salamander

Long-distance dispersal (LDD) may contribute disproportionately to range expansions, the creation of new evolutionary lineages, and species persistence in human-dominated landscapes. However, because data on the individual consequences of dispersal distance are extremely limited, we have little insight on how LDD is maintained in natural populations. I used six years of spatially explicit capture-mark-recapture (CMR) data to test the prediction that individual performance increases with dispersal distance in the stream salamander Gyrinophilus porphyriticus. Dispersal distance was total distance moved along the 1-km study stream, ranging from 0 to 565 m. To quantify individual performance, I used CMR estimates of survival and individual growth rates based on change in body length. Survival and growth rates increased significantly with dispersal distance. These relationships were not confounded by pre-dispersal body condition or by ecological gradients along the stream. Individual benefits of LDD were likely caused by an increase in the upper limit of settlement site quality with dispersal distance. My results do not support the view that the fitness consequences of LDD are unpredictable and instead suggest that consistent evolutionary mechanisms may explain the prevalence of LDD in nature. They also highlight the value of direct CMR data for understanding the individual consequences of variation in dispersal distance and how that variation is maintained in natural populations.     

The influence of disturbance events on survival and dispersal rates of Florida box turtles.

Disturbances have the potential to cause long-term effects to ecosystem structure and function, and they may affect individual species in different ways. Long-lived vertebrates such as turtles may be at risk from such events, inasmuch as their life histories preclude rapid recovery should extensive mortality occur. We applied capture-mark-recapture models to assess disturbance effects on a population of Florida box turtles (Terrapene carolina bauri) on Egmont Key, Florida, USA. Near the midpoint of the study, a series of physical disturbances affected the island, from salt water overwash associated with several tropical storms to extensive removal of nonindigenous vegetation. These disturbances allowed us to examine demographic responses of the turtle population and to determine if they affected dispersal throughout the island. Adult survival rates did not vary significantly either between sexes or among years of the study. Survival rates did not vary significantly between juvenile and adult turtles, or among years of the study. Furthermore, neither adult nor juvenile survival rates differed significantly between pre- and post-disturbance. However, dispersal rates varied significantly among the four major study sites, and dispersal rates were higher during the pre-disturbance sampling periods compared to post-disturbance. Our results suggest few long-term effects on the demography of the turtle population. Florida box turtles responded to tropical storms and vegetation control by moving to favorable habitats minimally affected by the disturbances and remaining there. As long as turtles and perhaps other long-lived vertebrates can disperse to non-disturbed habitat, and high levels of mortality do not occur in a population, a long life span may allow them to wait out the impact of disturbance with potentially little effect on long-term population processes.     

Abundance and survival rates of green turtles in an urban environment: coexistence of humans and an endangered species

Longitudinal capture-mark-recapture data were used to estimate abundance and survival rates for green turtles (Chelonia mydas) in San Diego Bay, California, USA. These turtles were closely associated with warm effluent from a power plant during winter months. The life stage distribution of green turtles in the bay ranged from post-pelagic juveniles to adults (44.0–110.4 cm straight carapace length). During 99 capture sessions between December 2, 1990, and March 25, 2009, 96 individual green turtles were caught. To estimate abundance and survival rates, robust-design mark-recapture models were fitted to capture-recapture histories using software MARK. The estimated annual survival rate was 0.861 (SE = 0.147, 95% CI = 0.356–0.986), whereas annual abundance ranged from 16 (SE = 6.3, 95% CI = 4–29) to 61 (SE = 13.2, 95% CI = 36–88). This study provides the first survival rate and abundance estimates for a green turtle foraging population in the highly industrialized San Diego Bay.     

The control of emigration and its consequences for the survival of populations

Dispersal is the key process enhancing the long-term persistence of metapopulations in heterogeneous and dynamic landscapes. However, any individual emigrating from a occupied patch also increases the risk of local population extinction. The consequences of this increase for metapopulation persistence likely depend on the control of emigration. In this paper, we present results of individual-based simulations to compare the consequences of density-independent (DIE) and density-dependent (DDE) emigration on the extinction risk of local populations and a two-patch metapopulation. (1) For completely isolated patches extinction risk increases linearly with realised emigration rates in the DIE scenario. (2) For the DDE scenario extinction risk is nearly insensitive to emigration as longs as emigration probabilities remain below ≈0.2. Survival chances are up to half an order of magnitude larger than for populations with DIE. (3) For low dispersal mortality both modes of emigration increase survival of a metapopulation by ca. one order of magnitude. (4) For high dispersal mortality only DDE can improve the global survival chances of the metapopulation. (5) With DDE individuals are only removed from a population at high population density and the risk of extinction due to demographic stochasticity is thus much smaller compared to the DIE scenario.With density-dependent emigration prospects of metapopulations survival may thus be much higher compared to a system with density-independent emigration. Consequently, the knowledge about the factors driving emigration may significantly affect our conclusions concerning the conservation status of species.     

Estimating species-specific survival and movement when species identification is uncertain

Incorporating uncertainty in the investigation of ecological studies has been the topic of an increasing body of research. In particular, mark-recapture methodology has shown that incorporating uncertainty in the probability of detecting individuals in populations enables accurate estimation of population-level processes such as survival, reproduction, and dispersal. Recent advances in mark-recapture methodology have included estimating population-level processes for biologically important groups despite the misassignment of individuals to those groups. Examples include estimating rates of apparent survival despite less than perfect accuracy when identifying individuals to gender or breeding state. Here we introduce a method for estimating apparent survival and dispersal in species that co-occur but that are difficult to distinguish. We use data from co-occurring populations of meadow voles (Microtus pennsylvanicus) and montane voles (M. montanus) in addition to simulated data to show that ignoring species uncertainty can lead to biased estimates of population processes. The incorporation of species uncertainty in mark-recapture studies should aid future research investigating ecological concepts such as interspecific competition, niche differentiation, and spatial population dynamics in sibling species.     

A multi-event model to study stage-dependent dispersal in radio-collared hares: when hunting promotes costly transience

Behavioral ecologists have often assumed that dispersal is costly mainly because of unfamiliarity with traversed habitats during dispersal and energy costs of the movement per se; thus, dispersers that have successfully settled should experience survival rates comparable to those of philopatric individuals. In this paper, we tested that hypothesis using 152 radio-collared European hares in a harvested population. We developed a multi-event capture recapture model, combining telemetry data and recoveries and separately modeling the foray probability, the settlement probability, and the permanent dispersal probability. The parameterization introduced here raises the possibility of separately testing effects on survival and dispersal probabilities at each stage of dispersal (departure, transience, and settlement). In accordance with our expectations, we reveal that dispersers incur higher mortality risks during transience and the early settlement period than philopatric individuals or settled dispersers. We also found that dispersers suffer from higher risks of being shot. Those results illustrate that unfamiliarity with the habitat during transience makes dispersal costly and that settled dispersers may enjoy survival rates comparable to those of philopatric individuals. Surprisingly, we also found that individuals have a higher probability of foraying during the hunting season. We suggest that hunting and related disturbances increase dispersal costs both by increasing mortality risk during transience and (perhaps) by increasing movement rates. We emphasize the need to take human pressures into account as factors that may drive the demographics of movements in populations.     

Grassland songbird survival and recruitment in agricultural landscapes: implications for source-sink demography

Population growth and decline are particularly sensitive to changes in three key life-history parameters: annual productivity, juvenile survival, and adult survival. However, for many species these parameters remain unknown. For example, although grassland songbirds are imperiled throughout North America, for this guild, only a small number of studies have assessed these parameters. From 2002 to 2006, in the agricultural landscape of the Champlain Valley of Vermont and New York, USA, we studied Savannah Sparrow (Passerculus sandwichensis) and Bobolink (Dolichonyx oryzivorus) demography on four grassland treatments: (1) early-hayed fields cut before 11 June and again in early- to mid-July; (2) middle-hayed fields cut once between 21 June and 10 July; (3) late-hayed fields cut after 1 August; and (4) rotationally grazed pastures. We assessed whether these treatments affected adult apparent survival (phi) and recruitment (f), how sensitive these parameters were to the presence of nonbreeders and local dispersal, and the populations' ability to persist in these four habitats. On average, birds using late-hayed fields had > 25% higher apparent survival than those on the more intensively managed early-hayed, middle-hayed, and grazed fields. Overall male phi was 35% higher than female phi, and Savannah Sparrow phi was 44% higher than Bobolink phi. Across all analyses and treatments, apparent survival estimates were 0.58-0.85 for male and 0.48-0.71 for female Savannah Sparrows, and 0.52-0.70 for male and 0.19-0.55 for female Bobolinks. For males of both species, potential nonbreeders decreased the precision of and lowered apparent survival estimates by 25%; female estimates showed little variation with the inclusion of nonbreeders. Inclusion of local dispersal observations increased apparent survival estimates and, in many cases, increased precision, though the effect was stronger for Savannah Sparrows than for Bobolinks, and also stronger for males than for females. High Savannah Sparrow apparent survival rates resulted in stable or near stable populations (lambda approximately 1), particularly in late-hayed and grazed fields, while low Bobolink apparent survival rates resulted in strongly declining populations (lambda < 1) in all treatments.     

Spatiotemporal variation in survival rates: implications for population dynamics of yellow-bellied marmots

Spatiotemporal variation in age-specific survival rates can profoundly influence population dynamics, but few studies of vertebrates have thoroughly investigated both spatial and temporal variability in age-specific survival rates. We used 28 years (1976-2003) of capture-mark-recapture (CMR) data from 17 locations to parameterize an age-structured Cormack-Jolly-Seber model, and investigated spatial and temporal variation in age-specific annual survival rates of yellow-bellied marmots (Marmota flaviventris). Survival rates varied both spatially and temporally, with survival of younger animals exhibiting the highest degree of variation. Juvenile survival rates varied from 0.52 +/- 0.05 to 0.78 +/- 0.10 among sites and from 0.15 +/- 0.14 to 0.89 +/- 0.06 over time. Adult survival rates varied from 0.62 +/- 0.09 to 0.80 +/- 0.03 among sites, but did not vary significantly over time. We used reverse-time CMR models to estimate the realized population growth rate (lamda), and to investigate the influence of the observed variation in age-specific survival rates on lamda. The realized growth rate of the population closely covaried with, and was significantly influenced by, spatiotemporal variation in juvenile survival rate. High variability in juvenile survival rates over space and time clearly influenced the dynamics of our study population and is also likely to be an important determinant of the spatiotemporal variation in the population dynamics of other mammals with similar life history characteristics.     

Strategic Rat Control for Restoring Populations of Native Species in Forest Fragments

Forest fragments have biodiversity value that may be enhanced through management such as control of non‐native predators. However, such efforts may be ineffective, and research is needed to ensure that predator control is done strategically. We used Bayesian hierarchical modeling to estimate fragment‐specific effects of experimental rat control on a native species targeted for recovery in a New Zealand pastoral landscape. The experiment was a modified BACI (before‐after‐control‐impact) design conducted over 6 years in 19 forest fragments with low‐density subpopulations of North Island Robins (Petroica longipes). The aim was to identify individual fragments that not only showed clear benefits of rat control, but also would have a high probability of subpopulation growth even if they were the only fragment managed. We collected data on fecundity, adult and juvenile survival, and juvenile emigration, and modeled the data in an integrated framework to estimate the expected annual growth rate (λ) of each subpopulation with and without rat control. Without emigration, subpopulation growth was estimated as marginal (λ = 0.95–1.05) or negative (λ = 0.74–0.90) without rat control, but it was estimated as positive in all fragments (λ = 1.4–2.1) if rats were controlled. This reflected a 150% average increase in fecundity and 45% average increase in adult female survival. The probability of a juvenile remaining in its natal fragment was 0.37 on average, but varied with fragment connectivity. With juvenile emigration added, 6 fragments were estimated to have a high (>0.8) probability of being self‐sustaining (λ > 1) with rat control. The key factors affecting subpopulation growth rates under rat control were low connectivity and stock fencing because these factors were associated with lower juvenile emigration and higher fecundity, respectively. However, there was also substantial random variation in adult survival among fragments, illustrating the importance of hierarchical modeling for fragmentation studies. Control Estratégico de Ratas para Restaurar Poblaciones de Especies Nativas en Fragmentos de Bosque     

Negative Effects of Habitat Loss on Survival of Migrant Warblers in a Forest Mosaic

Abstract: Habitat loss and fragmentation in forested landscapes often negatively affect animal abundance; however, whether these factors also affect fitness is not well known. We hypothesized that observed decreases in bird occurrence and abundance in landscapes with harvested forests are associated with reduced apparent survival of adults. We defined apparent survival as an estimate of survival that accounts for an imperfect resighting probability, but not permanent emigration (i.e., dispersal). We examined the association between spatially extensive habitat loss and apparent survival of males of 2 Neotropical migrant species, Blackburnian Warbler (Dendroica fusca) and Black‐Throated Green Warbler (D. virens), over 7 years in the Greater Fundy Ecosystem, New Brunswick, Canada. We estimated apparent survival among and within breeding seasons. We quantified amount of habitat in the context of individual species. In this landscape, boundaries between land‐cover types are gradual rather than clearly identifiable and abrupt. Estimated apparent within‐season survival of both species decreased as a function of amount of habitat within a 2000‐m radius; survival was approximately 12 times (95% CI 3.43–14) greater in landscapes with 85% habitat than in landscapes with 10% habitat. Apparent annual survival also decreased as a function of amount of habitat within a 100‐m radius. Over the range of habitat amount, apparent annual survival decreased 15% (95% CI 7–29%) as the amount of habitat decreased. Our results suggest that reduced species occurrence in landscapes with low proportions of habitat is due partly to lower apparent survival at these sites. This mechanism operates both directly (i.e., via effects on mortality or dispersal during breeding) and possibly through indirect effects during the nonbreeding season. Habitat loss was associated not only with a lower number of individuals, but also with lower survival of those individuals.     

Information processing in models for density-dependent emigration: A comparison

Density-dependent emigration has been recognized as a fitness enhancing strategy. Yet, especially in the modelling literature there is no consensus about how density-dependent emigration should quantitatively be incorporated into metapopulation models. In this paper we compare the performance of five different dispersal strategies (defined by the functional link between density and emigration probability). Four of these strategies are based on published functional relationships between local population density and emigration probability, one assumes density-independent dispersal. We use individual-based simulations of time-discrete metapopulation dynamics and conduct evolution experiments for a broad range of values for dispersal mortality and environmental stochasticity. For each set of these conditions we analyze the evolution of emigration rates in ‘monoculture experiments’ (with only one type of dispersal strategy used by all individuals in the metapopulation) as well as in selection experiments that allow a pair-wise comparison of the performance of each functional type. We find that a single-parameter ‘asymptotic threshold’ strategy - derived from the marginal value theorem - with a decelerating increase of emigration rate with increasing population density, out-competes any other strategy, i.e. density-independent emigration, a ‘linear threshold’ strategy and a flexible three-parameter strategy. Only when environmental conditions select for extremely high emigration probabilities (close to one), strategies may perform approximately equally. A simple threshold strategy derived for the case of continuous population growth performs even worse than the density-independent strategy. As the functional type of the dispersal function implemented in metapopulation models may severely affect predictions concerning the survival of populations, range expansion, or community changes we clearly recommend to carefully select adequate functions to model density-dependent dispersal.     

Roles of survival and dispersal in reintroduction success of Griffon vulture (Gyps fulvus).

The success of reintroduction programs greatly depends on the amount of mortality and dispersal of the released individuals. Although local environmental pressures are likely to play an important role in these processes, they have rarely been investigated because of the lack of spatial replicates of reintroduction. In the present study, we analyzed a 25-year data set encompassing 272 individuals released in five reintroduction programs of Griffon Vultures (Gyps fulvus) in France to examine the respective roles of survival and dispersal in program successes and failures. We use recent developments in multi-strata capture-recapture models to take into account tag loss in survival estimates and to consider and estimate dispersal among release areas. We also examined the effects of sex, age, time, area, and release status on survival, and we tested whether dispersal patterns among release areas were consistent with habitat selection theories. Results indicated that the survival of released adults was reduced during the first year after release, with no difference between sexes. Taking into account local observations only, we found that early survival rates varied across sites. However when we distinguished dispersal from mortality, early survival rates became equal across release sites. It thus appears that among reintroduction programs difference in failure and success was due to differential dispersal among release sites. We revealed asymmetrical patterns of dispersal due to conspecific attraction: dispersers selected the closest and the largest population. We showed that mortality can be homogeneous from one program to another while, on the contrary, dispersal is highly dependent on the matrix of established populations. Dispersal behavior is thus of major interest for metapopulation restoration and should be taken into account in planning reintroduction designs.     

An integrated approach for predicting fates of reintroductions with demographic data from multiple populations

We devised a novel approach to model reintroduced populations whereby demographic data collected from multiple sites are integrated into a Bayesian hierarchical model. Integrating data from multiple reintroductions allows more precise population-growth projections to be made, especially for populations for which data are sparse, and allows projections that account for random site-to-site variation to be made before new reintroductions are attempted. We used data from reintroductions of the North Island Robin (Petroica longipes), an endemic New Zealand passerine, to 10 sites where non-native mammalian predators are controlled. A comparison of candidate models that we based on deviance information criterion showed that rat-tracking rate (an index of rat density) was a useful predictor of robin fecundity and adult female survival, that landscape connectivity and a binary measure of whether sites were on a peninsula were useful predictors of apparent juvenile survival (probably due to differential dispersal away from reintroduction sites), and that there was unexplained random variation among sites in all demographic rates. We used the two best supported models to estimate the finite rate of increase (λ) for populations at each of the 10 sites, and for a proposed reintroduction site, under different levels of rat control. Only three of the reintroduction sites had λ distributions completely >1 for either model. At two sites, λ was expected to be >1 if rat-tracking rates were <5%. At the other five reintroduction sites, λ was predicted to be close to 1, and it was unclear whether growth was expected. Predictions of λ for the proposed reintroduction site were less precise than for other sites because distributions incorporated the full range of site-to-site random variation in vital rates. Our methods can be applied to any species for which postrelease data on demographic rates are available and potentially can be extended to model multiple species simultaneously.     

Habitat-mediated foraging limitations drive survival bottlenecks for juvenile salmon

Realistic population models and effective conservation strategies require a thorough understanding of mechanisms driving stage-specific mortality. Mortality bottlenecks for many species occur in the juvenile stage and are thought to result from limitation on food or foraging habitat during a "critical period" for growth and survival. Without a way to account for maternal effects or to measure integrated consumption rates in the field, it has been virtually impossible to test these relationships directly. Hence uncertainties about mechanisms underlying such bottlenecks remain. In this study we randomize maternal effects across sites and apply a new method for measuring consumption integrated over weeks to months to test the hypothesis that food limitation drives early-season juvenile mortality bottlenecks in Atlantic salmon (Salmo salar). Using natural signatures of geologically derived cesium (133Cs), we estimated consumption rates of >400 fry stocked into six streams. Two to four weeks after stocking, consumption was extremely low across sites (0.005 g x g(-1) x d(-1)) and was predicted to be below maintenance rations (i.e., yielding negative energy balances) for the majority of individuals from five of six sites. However, consumption during this time was positively correlated with growth rates and survival (measured at the end of the growing season). In contrast, consumption rates increased in mid- (0.030 g x g(-1) x d(-1)) and late (0.035 g x g(-1) x d(-1)) seasons, but juvenile survival and consumption were not correlated, and correlations between growth and consumption were weak. These findings are consistent with predictions of a habitat-based bioenergetic model constructed using the actual stream positions of the individual fish in the present study, which indicates that habitat-based models capture important environmental determinants of juvenile growth and survival. Hence, by combining approaches, reducing maternal effects and controlling initial conditions, we offer a general framework for linking foraging with juvenile survival and present the first direct consumption-based evidence for the early season bottleneck hypothesis.     

Forced dispersal of juvenile guanacos (<Emphasis Type="Italic">Lama guanicoe</Emphasis>): causes,variation, and fates of individuals dispersing at different times

We examined adult-juvenile conflict in the guanaco (Lama guanicoe). During spring, territorial males become increasingly aggressive toward all juveniles born the previous year and begin expelling them from family groups. In an apparent effort to reduce aggression, juveniles display submissive crouches when being observed, approached, or attacked by the territorial male. Therefore, we assessed the influence of juvenile submissive behavior on the timing of dispersal and also examined if dispersal time was related to survival and reproductive performance as adults. We also evaluated hypotheses regarding the evolution of juvenile mammalian dispersal in the context of if and how each may favor the forced dispersal of juvenile guanacos by territorial males. Juveniles generally dispersed in late spring and early summer, and a nearly equal proportion of females (n=46; 48%) and males (n=49; 52%) dispersed. More-submissive animals generally dispersed later than less-submissive animals. Juvenile sex and dispersal time were not related to survival. In contrast, juvenile sex and dispersal time were related to reproductive performance. The probability of reproducing was highest when juveniles dispersed early and decreased with increasing time in family groups prior to dispersal. The largest proportion of juveniles was forced to disperse during a 2-week interval following the peak of the breeding season. Competition for food resources is likely very intense at this juncture and territorial males may force older juveniles to disperse in order to divert food resources to younger neonates. Additionally, juveniles may be forced to disperse after territorial males mate their mothers to prevent lost mating opportunities, because females leave territories when their offspring disperse and possibly prior to mating with males. We conclude that the forced dispersal of juvenile guanacos by territorial males is ultimately driven by competition for food resources on territories. The timing of dispersal, however, may be tempered by the chronology of matings between territorial males and particular adult females, and/or genetic relatedness between territorial males and juveniles.     

Habitat fragmentation lowers survival of a tropical forest bird.

Population ecology research has long been focused on linking environmental features with the viability of populations. The majority of this work has largely been carried out in temperate systems and, until recently, has examined the effects of habitat fragmentation on survival. In contrast, we looked at the effect of forest fragmentation on apparent survival of individuals of the White-ruffed Manakin (Corapipo altera) in southern Costa Rica. Survival and recapture rates were estimated using mark-recapture analyses, based on capture histories from 1993 to 2006. We sampled four forest patches ranging in size from 0.9 to 25 ha, and four sites in the larger 227-ha Las Cruces Biological Station Forest Reserve (LCBSFR). We found a significant difference in annual adult apparent survival rates for individuals marked and recaptured in forest fragments vs. individuals marked and recaptured in the larger LCBSFR. Contrary to our expectation, survival and recapture probabilities did not differ between male and female manakins. Also, there was no support for the existence of annual variation in survival within each study site. Our results suggest that forest fragmentation is likely having an effect on population dynamics for the White-ruffed Manakin in this landscape. Therefore, populations that appear to be persisting in fragmented landscapes might still be at risk of local extinction, and conservation action for tropical birds should be aimed at identifying and reducing sources of adult mortality. Future studies in fragmentation effects on reproductive success and survival, across broad geographical scales, will be needed before it is possible to achieve a clear understanding of the effects of habitat fragmentation on populations for both tropical and temperate regions.     

Dispersal depression with habitat fragmentation in the bog fritillary butterfly

Habitat fragmentation is expected to impose strong selective pressures on dispersal rates. However, evolutionary responses of dispersal are not self-evident, since various selection pressures act in opposite directions. Here we disentangled the components of dispersal behavior in a metapopulation context using the Virtual Migration model, and we linked their variation to habitat fragmentation in the specialist butterfly Proclossiana eunomia. Our study provided a nearly unique opportunity to study how habitat fragmentation modifies dispersal at the landscape scale, as opposed to microlandscapes or simulation studies. Indeed, we studied the same species in four landscapes with various habitat fragmentation levels, in which large amounts of field data were collected and analyzed using similar methodologies. We showed the existence of quantitative variations in dispersal behavior correlated with increased fragmentation. Dispersal propensity from habitat patches (for a given patch size), and mortality during dispersal (for a given patch connectivity) were lower in more fragmented landscapes. We suggest that these were the consequences of two different evolutionary responses of dispersal behavior at the individual level: (1) when fragmentation increased, the reluctance of individuals to cross habitat patch boundaries also increased; (2) when individuals dispersed, they flew straighter in the matrix, which is the best strategy to improve dispersal success. Such evolutionary responses could generate complex nonlinear patterns of dispersal changes at the metapopulation level according to habitat fragmentation. Due to the small size and increased isolation of habitat patches in fragmented landscapes, overall emigration rate and mortality during dispersal remained high. As a consequence, successful dispersal at the metapopulation scale remained limited. Therefore, to what extent the selection of individuals with a lower dispersal propensity and a higher survival during dispersal is able to limit detrimental effects of habitat fragmentation on dispersal success is unknown, and any conclusion that metapopulations would compensate for them is flawed.     

Assessing tiger population dynamics using photographic capture-recapture sampling

Although wide-ranging, elusive, large carnivore species, such as the tiger, are of scientific and conservation interest, rigorous inferences about their population dynamics are scarce because of methodological problems of sampling populations at the required spatial and temporal scales. We report the application of a rigorous, noninvasive method for assessing tiger population dynamics to test model-based predictions about population viability. We obtained photographic capture histories for 74 individual tigers during a nine-year study involving 5725 trap-nights of effort. These data were modeled under a likelihood-based, "robust design" capture-recapture analytic framework. We explicitly modeled and estimated ecological parameters such as time-specific abundance, density, survival, recruitment, temporary emigration, and transience, using models that incorporated effects of factors such as individual heterogeneity, trap-response, and time on probabilities of photo-capturing tigers. The model estimated a random temporary emigration parameter of gamma" = gamma' = 0.10 +/- 0.069 (values are estimated mean +/- SE). When scaled to an annual basis, tiger survival rates were estimated at S = 0.77 +/- 0.051, and the estimated probability that a newly caught animal was a transient was tau = 0.18 +/- 0.11. During the period when the sampled area was of constant size, the estimated population size N(t) varied from 17 +/- 1.7 to 31 +/- 2.1 tigers, with a geometric mean rate of annual population change estimated as lambda = 1.03 +/- 0.020, representing a 3% annual increase. The estimated recruitment of new animals, B(t), varied from 0 +/- 3.0 to 14 +/- 2.9 tigers. Population density estimates, D, ranged from 7.33 +/- 0.8 tigers/100 km2 to 21.73 +/- 1.7 tigers/100 km2 during the study. Thus, despite substantial annual losses and temporal variation in recruitment, the tiger density remained at relatively high levels in Nagarahole. Our results are consistent with the hypothesis that protected wild tiger populations can remain healthy despite heavy mortalities because of their inherently high reproductive potential. The ability to model the entire photographic capture history data set and incorporate reduced-parameter models led to estimates of mean annual population change that were sufficiently precise to be useful. This efficient, noninvasive sampling approach can be used to rigorously investigate the population dynamics of tigers and other elusive, rare, wide-ranging animal species in which individuals can be identified from photographs or other means.     

Survival probability estimates for the endangered loggerhead sea turtle resident in southern Great Barrier Reef waters

Sex- and age-class-specific survival of a loggerhead turtle population resident in southern Great Barrier Reef waters was estimated using a long-term capture-mark-recapture (CMR) study and the Cormack-Jolly-Seber modelling approach. The CMR history profiles for 271 loggerheads tagged over 9 years (1984-1992) were classified into two age classes (adult, immature) based on somatic growth and reproductive traits. The sex and maturity status of each turtle was determined from visual examination of reproductive organs using laparoscopy. A reduced-parameter model accounting for constant survival with sex- and time-specific recapture was adequate for estimating age-class-specific survival probabilities, but inclusion of time-specific transient behaviour was informative for the immature age class. The annual fluctuations in the estimated proportion of transient immatures was not a function of sampling effort, but could be due to anomalous oceanographic conditions affecting dispersal of the immature class. There was no sex-specific difference in survival probabilities for either age class, but females were more likely to be recaptured than males, which might be related to behavioural differences such as sex-biased dispersal. The expected annual survival probability for adults was 0.875 (95% CI: 0.84-0.91). The expected annual survival probability for immatures was 0.859 (95% CI: 0.83-0.89), but when the transients were accounted for, the expected annual survival for the resident immature loggerheads was 0.918 (95% CI: 0.88-0.96). These are the first substantive estimates of annual survival probabilities for any loggerhead sea-turtle stock and provide a basis for developing a better understanding of loggerhead population dynamics.