Spatial Variation In Survival Research Articles

Estimating spatially variable and density-dependent survival using open-population spatial capture-recapture models.

Open-population spatial capture-recapture (OPSCR) models use the spatial information contained in individual detections collected over multiple consecutive occasions to estimate not only occasion-specific density, but also demographic parameters. OPSCR models can also estimate spatial variation in vital rates, but such models are neither widely used nor thoroughly tested. We developed a Bayesian OPSCR model that not only accounts for spatial variation in survival using spatial covariates but also estimates local density-dependent effects on survival within a unified framework. Using simulations, we show that OPSCR models provide sound inferences on the effect of spatial covariates on survival, including multiple competing sources of mortality, each with potentially different spatial determinants. Estimation of local density-dependent survival was possible but required more data due to the greater complexity of the model. Not accounting for spatial heterogeneity in survival led to up to 10% positive bias in abundance estimates. We provide an empirical demonstration of the model by estimating the effect of country and density on cause-specific mortality of female wolverines (Gulo gulo) in central Sweden and Norway. The ability to make population-level inferences on spatial variation in survival is an essential step toward a fully spatially explicit OPSCR model capable of disentangling the role of multiple spatial drivers of population dynamics.

Temporal variation and landcover influence survival in adult female mottled ducks

AbstractAdult survival is a key driver of waterfowl population growth and is subject to temporal and spatial variation. Mottled ducks (Anas fulvigula) are native to the Gulf Coast and peninsular Florida, USA, and have suffered population declines over the past decade, especially in Texas and Louisiana, USA. Although the cause of this decline is not well understood, previous research concluded variation in survival contributed to nearly a third of variation in the species' population growth rate. We used global positioning system‐groupe spécial mobile (GPS‐GSM) transmitters to study temporal and spatial variation in survival of adult female western Gulf Coast mottled ducks in southwestern Louisiana, 2017–2020. We evaluated weekly survival models parameterized with combinations of hunted and non‐hunted periods, biological seasons, and landcover types that were used by mottled ducks. There were 3 competitive survival models, and all contained 4 parameters that parsed the annual cycle into the non‐hunted period, first part of the general waterfowl season, and second part of the waterfowl season, and included the proportion of GPS locations in agricultural lands. Weekly survival was 0.979 during the first part of the general waterfowl hunting season, and 0.996 during the second part of the general waterfowl season. Daily survival rate increased with an increasing proportion of locations logged in agricultural lands. Annual survival rates were similar to other waterfowl that are not experiencing population declines, which suggests survival is not limiting population growth of mottled ducks along the western Gulf Coast. Managers should ensure the availability of refuge areas where hunting is prohibited during the first part of the general waterfowl season, when mottled ducks are at an increased risk of mortality, in addition to the targeted conservation of agricultural lands that provide cover and forage.

Estimation in the multinomial reencounter model – Where do migrating animals go and how do they survive in their destination area?

Spatial variation in survival has individual fitness consequences and influences population dynamics. Which space animals use during the annual cycle determines how they are affected by this spatial variability. Therefore, knowing spatial patterns of survival and space use is crucial to understand demography of migrating animals. Extracting information on survival and space use from observation data, in particular dead recovery data, requires explicitly identifying the observation process.We build a fully stochastic model for animals marked in populations of origin, which were found dead in spatially discrete destination areas. The model acts on the population level and includes parameters for use of space, survival and recovery probability. It is based on the division coefficient and the multinomial reencounter model. We use a likelihood-based approach, derive Restricted Maximum Likelihood-like estimates for all parameters and prove their existence and uniqueness.In a simulation study we demonstrate the performance of the model by using Bayesian estimators derived by the Markov chain Monte Carlo method. We obtain unbiased estimates for survival and recovery probability if the sample size is large enough. Moreover, we apply the model to real-world data of European robins Erithacus rubecula ringed at a stopover site. We obtain annual survival estimates for different spatially discrete non-breeding areas. Additionally, we can reproduce already known patterns of use of space for this species.

Spatial Variation of Survival for Colorectal Cancer in Malaysia.

A patient’s survival may depend on several known and unknown factors and it may also vary spatially across a region. Socioeconomic status, accessibility to healthcare and other environmental factors are likely to contribute to survival rates. The aim of the study was to model the spatial variation in survival for colorectal cancer patients in Malaysia, accounting for individual and socioeconomic risk factors. We conducted a retrospective study of 4412 colorectal cancer (ICD-10, C18-C20) patients diagnosed from 2008 to 2013 to model survival in CRC patients. We used the data recorded in the database of the Malaysian National Cancer Patient Registry-Colorectal Cancer (NCPR-CRC). Spatial location was assigned based on the patients’ central district location, which involves 144 administrative districts of Malaysia. We fitted a parametric proportional hazards model in which the spatially correlated frailties were modelled by a log-Gaussian stochastic process to analyse the spatially referenced survival data, which is also known as a spatial survival model. After controlling for individual and area level characteristics, our findings indicate wide spatial variation in colorectal cancer survival across Malaysia. Better healthcare provision and higher socioeconomic index in the districts where patients live decreased the risk of death from colorectal cancer, but these associations were not statistically significant. Reliable measurement of environmental factors is needed to provide good insight into the effects of potential risk factors for the disease. For example, a better metric is needed to measure socioeconomic status and accessibility to healthcare in the country. The findings provide new information that might be of use to the Ministry of Health in identifying populations with an increased risk of poor survival, and for planning and providing cancer control services.

Does seasonality drive spatial patterns in demography? Variation in survival in African reed warblers Acrocephalus baeticatus across southern Africa does not reflect global patterns.

Among birds, northern temperate species generally have larger clutches, shorter development periods and lower adult survival than similarly-sized southern and tropical species. Even though this global pattern is well accepted, the driving mechanism is still not fully understood. The main theories are founded on the differing environmental seasonality of these zones (higher seasonality in the North). These patterns arise in cross-species comparisons, but we hypothesized that the same patterns should arise among populations within a species if different types of seasonality select for different life histories. Few studies have examined this. We estimated survival of an azonal habitat specialist, the African reed warbler, across the environmentally diverse African subcontinent, and related survival to latitude and to the seasonality of the different environments of their breeding habitats. Data (1998–2010) collected through a public ringing scheme were analyzed with hierarchical capture-mark-recapture models to determine resident adult survival and its spatial variance across sixteen vegetation units spread across four biomes. The models were defined as state-space multi-state models to account for transience and implemented in a Bayesian framework. We did not find a latitudinal trend in survival or a clear link between seasonality and survival. Spatial variation in survival was substantial across the sixteen sites (spatial standard deviation of the logit mean survival: 0.70, 95% credible interval (CRI): 0.33–1.27). Mean site survival ranged from 0.49 (95% CRI: 0.18–0.80) to 0.83 (95% CRI: 0.62–0.97) with an overall mean of 0.67 (95% CRI: 0.47–0.85). A hierarchical modeling approach enabled us to estimate spatial variation in survival of the African reed warbler across the African subcontinent from sparse data. Although we could not confirm the global pattern of higher survival in less seasonal environments, our findings from a poorly studied region contribute to the study of life-history strategies.

Accounting for individual behavioural variation in studies of habitat selection

A caribou wearing an animal-borne video camera (a) and animal-borne video footage taken from systems deployed on mule deer (Odocoileus hemionus) in north-central Washington state, USA (b-d). When paired with tracking technology, animal-borne video can reveal detailed information about behaviour and environmental features at each location: (b) feeding, (c) vigilant in the open, (d) vigilant in cover, (e) resting in the open. Accordingly, animal-borne video systems should allow for analyses of habitat selection by individuals in particular behavioural states. In Focus: DeCesare, N.J., Hebblewhite, M., Bradley, M., Hervieux, D., Neufeld, L. & Musiani, M. (2014) Linking habitat selection and predation risk to spatial variation in survival. Journal of Animal Ecology, 83, 343-352. Resource selection is often assumed to confer enhanced fitness, but this assumption is rarely examined. In a study involving woodland caribou subject to grey wolf predation, DeCesare et al. (2014) show that while patterns of selection by caribou did correspond with a fitness proxy (survival probability), individuals did not avoid wolf predation risk to the extent that would minimize mortality. Here, we use the results of this paper as a springboard for discussing the choice of fitness proxies and the need to account for individual behavioural variation in studies of resource selection.

Effects of native macroalgae and predators on survival, condition and growth of non-indigenous Pacific oysters (Crassostrea gigas)

Biological invasions by non-indigenous species are causing widespread environmental changes and can threaten biodiversity and ecosystem services. Biotic interactions can play a key role in promoting or inhibiting spread of non-indigenous species. The Pacific oyster (Crassostrea gigas Thunberg, 1793) is commonly found on intertidal shores with hard substrata, but is negatively associated with macroalgae and may be influenced by the macroalgae itself or by predators associated with it. The influence of macroalgae and predation on growth, condition and survival of juvenile Pacific oysters were tested in a field experiment at two intertidal boulder shores using hatchery-reared juvenile oysters deployed on ceramic tiles. After 10months, condition of oysters was not significantly decreased in the presence of macroalgal canopy. At one site growth was reduced by macroalgae and at the other, it tended to be enhanced, but the effect was not significant. An effect of predation could not be detected. Although spatial variation in survival and growth of juvenile oysters was found, the results suggest that pre-settlement processes and/or post-settlement processes acting upon juvenile oysters at sizes not tested in this study might better explain variation in abundance of Pacific oysters in intertidal habitats.

Linking habitat selection and predation risk to spatial variation in survival

1. A central assumption underlying the study of habitat selection is that selected habitats confer enhanced fitness. Unfortunately, this assumption is rarely tested, and in some systems, gradients of predation risk may more accurately characterize spatial variation in vital rates than gradients described by habitat selection studies.2. Here, we separately measured spatial patterns of both resource selection and predation risk and tested their relationships with a key demographic trait, adult female survival, for a threatened ungulate, woodland caribou (Rangifer tarandus caribou Gmelin). We also evaluated whether exposure to gradients in both predation risk and resource selection value was manifested temporally through instantaneous or seasonal effects on survival outcomes.3. We used Cox proportional hazards spatial survival modelling to assess the relative support for 5 selection- and risk-based definitions of habitat quality, as quantified by woodland caribou adult female survival. These hypotheses included scenarios in which selection ideally mirrored survival, risk entirely drove survival, non-ideal selection correlated with survival but with additive risk effects, an ecological trap with maladaptive selection and a non-spatial effect of annual variation in weather.4. Indeed, we found positive relationships between the predicted values of a resource selection function (RSF) and survival, yet subsequently incorporating an additional negative effect of predation risk greatly improved models further. This revealed a positive, but non-ideal relationship between selection and survival. Gradients in these covariates were also shown to affect individual survival probability at multiple temporal scales. Exposure to increased predation risk had a relatively instantaneous effect on survival outcomes, whereas variation in habitat suitability predicted by an RSF had both instantaneous and longer-term seasonal effects on survival.5. Predation risk was an additive source of hazard beyond that detected through selection alone, and woodland caribou selection thus was shown to be non-ideal. Furthermore, by combining spatial adult female survival models with herd-specific estimates of recruitment in matrix population models, we estimated a spatially explicit landscape of population growth predictions for this endangered species.

Combining Traits and Density to Model Recruitment of Sessile Organisms

We propose an integrative approach that explains patterns of recruitment to adult populations in sessile organisms by considering the numbers of individuals and their body size. A recruitment model, based on a small number of parameters, was developed for sessile organisms and tested using the barnacle Semibalanus balanoides, a marine invertebrate inhabiting North Atlantic intertidal shores. Incorporating barnacle body size improved model fit beyond that based on density alone, showing that growth played an important role in how resource limitation affected survival. Our approach uncovered the following: First, changes in the shape of the recruitment curve resulted from the balance between individual growth and mortality. Second, recruitment was limited by the least plastic trait used to characterise body size, operculum area. Basal area, a trait that responded to increases in barnacle density, did not contribute significantly to explain patterns of recruitment. Third, some temporal variation is explained by changes in the amount of space occupied by shells of dead barnacles: at high cover barnacles are densely packed and these shells remain long after death. Fourth, seasonal variation and spatial variation in survival can be separated from that resulting from resource limitation; survival was predicted for two different shores and four sampling times using a single recruitment model. We conclude that applying this integrative approach to recruitment will lead to a considerable advance in understanding patterns of mortality of early stages of sessile organisms.

Nonparametric spatial regression of survival probability: visualization of population sinks in Eurasian Woodcock

Both evolutionary ecologists and wildlife managers make inference based on how fitness and demography vary in space. Spatial variation in survival can be difficult to assess in the wild because (1) multisite study designs are not well suited to populations that are continuously distributed across a large area and (2) available statistical models accounting for detectability less than 1.0 do not easily cope with geographical coordinates. Here we use penalized splines within a Bayesian state-space modeling framework to estimate and visualize survival probability in two dimensions. The approach is flexible in that no parametric form for the relationship between survival and coordinates need be specified a priori. To illustrate our method, we study a game species, the Eurasian Woodcock Scolopax rusticola, based on band recovery data (5000 individuals) collected over a > 50 000-km2 area in west-central France with contrasted habitats and hunting pressures. We find that spatial variation in survival probability matches an index of hunting pressure and creates a mosaic of population sources and sinks. Such analyses could provide guidance concerning the spatial management of hunting intensity or could be used to identify pathways of spatial variation in fitness, for example, to study adaptation to changing landscape and climate.

Geographic variation in reproduction and survival of kelp gulls Larus dominicanus vetula in southern Africa

Different populations of a species tend to vary in survival and reproduction, but the extent and scale of such spatial variation are poorly known. We estimated survival and clutch size of kelp gulls Larus dominicanus vetula across their entire African range. At this large geographic scale, we found no evidence for spatial variation in survival, and there was no variation in clutch size. However, there was considerable variation in clutch size among colonies within regions. Over the whole study, mean annual survival of juvenile and adult birds was 0.44 and 0.84, and mean clutch size was 2.2 eggs. A matrix population model showed that population growth was least sensitive to variation in clutch size, and the observed variation in clutch size could not fully account for the observed variation in population growth among colonies and regions. Our results thus suggest that dispersal and/or variation in survival (including egg/nestling survival) at a small spatial scale are also important for the spatial pattern of kelp gull population dynamics. These results are consistent with a metapopulation approach to spatial population dynamics.

Modeling Spatial Variation in Leukemia Survival Data

In this article we combine ideas from spatial statistics with lifetime data analysis techniques to investigate possible spatial variation in survival of adult acute myeloid leukemia patients in northwest England. Exploratory analysis suggests both clinically and statistically significant variation in survival rates across the region. A multivariate gamma frailty model incorporating spatial dependence is proposed and applied, with results confirming the dependence of hazard on location.

Life-History Implications of Large-Scale Spatial Variation in Adult Survival of Black Brant (Branta Bernicla Nigricans)

Abstract We used capture–recapture methods to estimate adult survival rates for adult female Black Brant (Branta bernicla nigricans; hereafter “brant”) from three colonies in Alaska, two on the Yukon-Kuskokwim Delta, and one on Alaska's Arctic coast. Costs of migration and reproductive effort varied among those colonies, enabling us to examine variation in survival in relation to variation in these other variables. We used the Barker model in program MARK to estimate true annual survival for brant from the three colonies. Models allowing for spatial variation in survival were among the most parsimonious models but were indistinguishable from a model with no spatial variation. Point estimates of annual survival were slightly higher for brant from the Arctic (0.90 ± 0.036) than for brant from either Tutakoke River (0.85 ± 0.004) or Kokechik Bay (0.86 ± 0.011). Thus, our survival estimates do not support a hypothesis that the cost of longer migrations or harvest experienced by brant from the Arctic reduced their annual survival relative to brant from the Yukon-Kuskokwim Delta. Spatial variation in survival provides weak support for life-history theory because brant from the region with lower reproductive investment had slightly higher survival.

LIFE-HISTORY IMPLICATIONS OF LARGE-SCALE SPATIAL VARIATION IN ADULT SURVIVAL OF BLACK BRANT (BRANTA BERNICLA NIGRICANS)

Abstract We used capture–recapture methods to estimate adult survival rates for adult female Black Brant (Branta bernicla nigricans; hereafter “brant”) from three colonies in Alaska, two on the Yukon-Kuskokwim Delta, and one on Alaska's Arctic coast. Costs of migration and reproductive effort varied among those colonies, enabling us to examine variation in survival in relation to variation in these other variables. We used the Barker model in program MARK to estimate true annual survival for brant from the three colonies. Models allowing for spatial variation in survival were among the most parsimonious models but were indistinguishable from a model with no spatial variation. Point estimates of annual survival were slightly higher for brant from the Arctic (0.90 ± 0.036) than for brant from either Tutakoke River (0.85 ± 0.004) or Kokechik Bay (0.86 ± 0.011). Thus, our survival estimates do not support a hypothesis that the cost of longer migrations or harvest experienced by brant from the Arctic reduced t...

Analysis of spatial and yearly variation in winter survival of winter wheat

Four years of winter survival data for winter wheat (Triticum aestivum L.) were collected on a loam soil located on the Central Experimental Farm at Ottawa, Ontario (45° 23′N, 75° 43′W). The site was low-lying and subject to frequent winter flooding and ice sheet formation. It appeared level although there was microtopographic variation with a range in elevation of approximately 0.15 m. The objective of the study was to gain insight into factors which might affect winter survival. Two varieties, a soft white and a hard red winter wheat, were planted in September. Crop establishment was measured in late fall and the percentage survival was measured in April of the following year. We assumed the large spatial differences in survival were not totally random, but rather were affected by spatial variation in environmental factors such as snow and ice depth, soil moisture and temperature. Hourly measurements of soil temperature at a depth of 0.05 m were recorded throughout the fall, winter and spring. Fall and spring soil moisture at the same depth were measured on the plot, as well as snow and ice depth at selected times throughout the winter. Measurements were taken on a grid covering the plot to help explain spatial variation in survival. In addition, detailed measurements of elevation were taken on a grid. Meteorological data were available from the nearby weather site. While soil temperatures were never low enough to kill plants according to the CERES model, the spatial variation in winter survival was associated with differences in elevation and the resulting surface drainage patterns.

Temporal and Spatial Variation in Survival Rates of the Tropical Lizard Anolis limifrons

We evaluated survival of the lizard, Anolis limifrons at two sites, AVA and Lutz, from 1976-1979 and during two periods at Lutz site, 1971-1976 vs 1976-1979, at Barro Colorado Island, Panama. Survival of adult females and males did not differ nor did survival of juveniles and adults. In contrast, survival was significantly higher at Lutz site during 1971-1976 than during 1976-1979 and survival was significantly higher at AVA than at Lutz site during 1976-1979. On an annual basis, mean survival rates were 0.042, 0.013, 0.055 for Lutz 1971-1976, Lutz 1976-1979, and AVA 1976-1979, respectively. These rates are in accord with reports of annual population turnover for this and other small mainland Anolis. Temporal and spatial variation in survival was not associated with habitat, season, year of observation, or numbers and abundances of avian predators. Survival patterns of mainland Anolis are contrasted with those of West Indian species in terms of life history evolution.