Bayesian methods for analyzing movements in heterogeneous landscapes from mark-recapture data

Spatially referenced mark-recapture data are becoming increasingly available, but the analysis of such data has remained difficult for a variety of reasons. One of the fundamental problems is that it is difficult to disentangle inherent movement behavior from sampling artifacts. For example, in a typical study design, short distances are sampled more frequently than long distances. Here we present a modeling-based alternative that combines a diffusion-based process model with an observation model to infer the inherent movement behavior of the species from the data. The movement model is based on classifying the landscape into a number of habitat types, and assuming habitat-specific diffusion and mortality parameters, and habitat selection at edges between the habitat types. As the problem is computationally highly intensive, we provide software that implements adaptive Bayesian methods for effective sampling of the posterior distribution. We illustrate the modeling framework by analyzing individual mark-recapture data on the Glanville fritillary butterfly (Melitaea cinxia), and by comparing our results with earlier ones derived from the same data using a purely statistical approach. We use simulated data to perform an analysis of statistical power, examining how accuracy in parameter estimates depends on the amount of data and on the study design. Obtaining precise estimates for movement rates and habitat preferences turns out to be especially challenging, as these parameters can be highly correlated in the posterior density. We show that the parameter estimates can be considerably improved by alternative study designs, such as releasing some of the individuals into the unsuitable matrix, or spending part of the recapture effort in the matrix.     

Modelling dispersal with diffusion and habitat selection: Analytical results for highly fragmented landscapes

Quantifying dispersal is crucial both for understanding ecological population dynamics, and for gaining insight into factors that affect the genetic structure of populations. The role of dispersal becomes pronounced in highly fragmented landscapes inhabited by spatially structured populations. We consider a landscape consisting of a set of habitat patches surrounded by unsuitable matrix, and model dispersal by assuming that the individuals follow a random walk with parameters that may be specific to the habitat type. We allow for spatial variation in patch quality, and account for edge-mediated behavior, the latter meaning that the individuals bias their movement towards the patches when close to an edge between a patch and the matrix. We employ a diffusion approximation of the random walk model to derive analytical expressions for various characteristics of the dispersal process. For example, we derive formulae for the time that an individual is expected to spend in its current patch i, and for the time that it will spend in the matrix, both conditional on the individual hitting next a given patch j before hitting any of the other patches or dying. The analytical formulae are based on the assumptions that the landscape is infinitely large, that the patches are circularly shaped, and that the patches are small compared to interpatch distances. We evaluate the effect of these assumptions by comparing the analytical results to numerical results in a real patch network that violates all of the three assumptions. We then consider a landscape that fulfills the assumptions, and show that in this case the analytical results are in a very good agreement with the numerical results. The results obtained here allow the construction of computationally efficient dispersal models that can be used as components of metapopulation models.     

Age-dependent survival analyzed with Bayesian models of mark-recapture data

We describe a Bayesian random effects model of mark-recapture data that accounts for age-dependence in survival and individual heterogeneity in capture probabilities and survival. The model is applied to data on the Glanville fritillary butterfly (Melitaea cinxia) collected from a population enclosed in a large cage in the field. The cage population consisted of a mixture of butterflies originating from newly established and old populations in a large metapopulation in the Aland Islands in Finland. The explanatory variables in the model included the effects of temperature, sex, and population type (new vs. old) on capture probabilities, and the effects of age, sex, population type, and day vs. night on survival. We found that mortality rate increased with age, that mortality rate was much higher during the day than during the night, and that the life span of females originating from newly established populations was shorter than the life span of females from old populations. Capture probability decreased with increasing temperature and decreased with increasing mobility of individuals.     

Digit length ratio (2D/4D): comparing measurements from X-rays and photographs in field voles (<Emphasis Type="Italic">Microtus agrestis</Emphasis>)

The ratio of second-to-fourth digit length (2D/4D) has been suggested to be a useful adult age marker of intrauterine exposure to steroids because it should be sexually dimorphic and fixed already in utero. Numerous studies mainly on humans have supported this conclusion, but it is yet unclear how well this applies to other vertebrates. This information would be especially valuable to field biologists to whom measuring steroids in utero is often impossible. The non-human studies conducted so far have yielded inconsistent results, perhaps due to the variety of different methods employed in measuring 2D/4D. We examined the age and sex dependency and lateral asymmetry of 2D/4D in field voles (Microtus agrestis) and compared whether these effects differed between 2D/4D measurements taken from photographs and X-rays. Our results show that 2D/4D measurements from photos had a higher measurement error and gave consistently higher 2D/4D than those from X-rays. According to both measurement methods, the right paw showed higher 2D/4D values than the left paw (lateral asymmetry). Adult voles had a lower 2D/4D than juveniles when measured from X-rays, but not when measured from photographs. We found no evidence for a sex difference in 2D/4D using either of the measurement methods. Our findings thus suggest that X-rays, due to their greater accuracy, should be preferred over photographs for measuring digit ratios in rodents. Our results also indicate that in this species, 2D/4D is laterally asymmetric, but it may not be either fixed in utero or sexually dimorphic. In conclusion, 2D/4D appears to be a rather species- and method-specific measure and researchers should be careful when generalising its applicability to study early hormonal effects in vertebrates.