Accounting for geographic variation in species-habitat associations during habitat suitability modeling.

Range-wide species conservation efforts are facilitated by spatially explicit estimates of habitat suitability. However, species-environment relationships often vary geographically and models assuming geographically constant relationships may result in misleading inferences. We present the first range-wide habitat suitability model (HSM) for the federally threatened eastern indigo snake (Drymarchon couperi) as a case study illustrating an approach to account for known latitudinal variation in habitat associations. Specifically, we modeled habitat suitability using interactive relationships between minimum winter temperature and several a priori environmental covariates and compared our results to those from models assuming geographically constant relationships. We found that multi-scale models including interactive effects with winter temperature outperformed single-scale models and models not including interactive effects with winter temperature. Our top-ranked model had suitable range-wide predictive performance and identified numerous large (i.e., ≥1000 ha) potential habitat patches throughout the indigo snake range. Predictive performance was greatest in southern Georgia and northern Florida likely reflecting more restrictive indigo snake habitat associations in these regions. This study illustrates how modeling interactive effects between temperature and environmental covariates can improve the performance of HSMs across geographically varying environmental gradients.

Multiscale assessment of functional connectivity: Landscape genetics of eastern indigo snakes in an anthropogenically fragmented landscape in central Florida.

Landscape features can strongly influence gene flow and the strength and direction of these effects may vary across spatial scales. However, few studies have evaluated methodological approaches for selecting spatial scales in landscape genetics analyses, in part because of computational challenges associated with optimizing landscape resistance surfaces (LRS). We used the federally threatened eastern indigo snake (Drymarchon couperi) in central Florida as a case study with which to compare the importance of landscape features and their scales of effect in influencing gene flow. We used genetic algorithms (ResistanceGA) to empirically optimize LRS using categorical land cover surfaces, multiscale resource selection surfaces (RSS), and four combinations of landscape covariates measured at multiple spatial scales (multisurface multiscale LRS). We compared LRS where scale was selected using pseudo- and full optimization. Multisurface multiscale LRS received more empirical support than LRS optimized from categorical land cover surfaces or RSS. Multiscale LRS with scale selected using full optimization generally outperformed those with scale selected using pseudo-optimization. Multiscale LRS with large spatial scales (1200-1800 m) received the most empirical support. Our results highlight the importance of considering landscape features across multiple spatial scales in landscape genetic analyses, particularly broad scales relative to species movement potential. Different effects of scale on home range-level movements and dispersal could explain weak associations between habitat suitability and gene flow in other studies. Our results also demonstrate the importance of large tracts of undeveloped upland habitat with heterogenous vegetation communities and low urbanization for promoting indigo snake connectivity.

Relationships of catch-per-unit-effort metrics with abundance vary depending on sampling method and population trajectory.

Catch-per-unit-effort (CPUE) is often used to monitor wildlife populations and to develop statistical population models. Animals caught and released are often not included in CPUE metrics and their inclusion may create more accurate indices of abundance. We used 21 years of detailed harvest records for bobcat (Lynx rufus) in Wisconsin, U.S.A., to calculate CPUE and 'actual CPUE' (ACPUE; including animals caught and released) from bobcat hunters and trappers. We calibrated these metrics to an independent estimate of bobcat abundance and attempted to create simple but effective models to estimate CPUE and ACPUE using harvest success data (i.e., bobcats harvested/available permits). CPUE showed virtually no relationship with bobcat abundance across all years, but both CPUE and ACPUE had stronger, non-linear, and negative relationships with abundance during the periods when the population was decreasing. Annual harvest success strongly predicted composite ACPUE and CPUE from hunters and trappers and hunter ACPUE and CPUE but was a poorer predictor of trapper ACPUE and CPUE. The non-linear, and sometimes weak, relationships with bobcat abundance likely reflect the increasing selectivity of bobcat hunters for trophy animals. Studies calibrating per-unit-effort metrics against abundance should account for population trajectories and different harvest methods (e.g., hunting and trapping). Our results also highlight the potential for estimating per-unit-effort metrics from relatively simple and inexpensive data sources and we encourage additional research into the use of per-unit-effort metrics for population estimation.

Taxonomic and conservation implications of population genetic admixture, mito-nuclear discordance, and male-biased dispersal of a large endangered snake, Drymarchon couperi.

Accurate species delimitation and description are necessary to guide effective conservation of imperiled species, and this synergy is maximized when multiple data sources are used to delimit species. We illustrate this point by examining Drymarchon couperi (Eastern Indigo Snake), a large, federally-protected species in North America that was recently divided into two species based on gene sequence data from three loci and heuristic morphological assessment. Here, we re-evaluate the two-species hypothesis for D. couperi by evaluating both population genetic and gene sequence data. Our analyses of 14 microsatellite markers revealed 6-8 genetic population clusters with significant admixture, particularly across the contact zone between the two hypothesized species. Phylogenetic analyses of gene sequence data with maximum-likelihood methods suggested discordance between mitochondrial and nuclear markers and provided phylogenetic support for one species rather than two. For these reasons, we place Drymarchon kolpobasileus into synonymy with D. couperi. We suggest inconsistent patterns between mitochondrial and nuclear DNA are driven by high dispersal of males relative to females. We advocate for species delimitation exercises that evaluate admixture and gene flow in addition to phylogenetic analyses, particularly when the latter reveal monophyletic lineages. This is particularly important for taxa, such as squamates, that exhibit strong sex-biased dispersal. Problems associated with over-delimitation of species richness can become particularly acute for threatened and endangered species, because of high costs to conservation when taxonomy demands protection of more individual species than are supported by accumulating data.

The Influence of Sex and Season on Conspecific Spatial Overlap in a Large, Actively-Foraging Colubrid Snake.

Understanding the factors influencing the degree of spatial overlap among conspecifics is important for understanding multiple ecological processes. Compared to terrestrial carnivores, relatively little is known about the factors influencing conspecific spatial overlap in snakes, although across snake taxa there appears to be substantial variation in conspecific spatial overlap. In this study, we described conspecific spatial overlap of eastern indigo snakes (Drymarchon couperi) in peninsular Florida and examined how conspecific spatial overlap varied by sex and season (breeding season vs. non-breeding season). We calculated multiple indices of spatial overlap using 6- and 3-month utilization distributions (UD) of dyads of simultaneously adjacent telemetered snakes. We also measured conspecific UD density values at each telemetry fix and modeled the distribution of those values as a function of overlap type, sex, and season using generalized Pareto distributions. Home range overlap between males and females was significantly greater than overlap between individuals of the same sex and male home ranges often completely contained female home ranges. Male home ranges overlapped little during both seasons, whereas females had higher levels of overlap during the non-breeding season. The spatial patterns observed in our study are consistent with those seen in many mammalian carnivores, in which low male-male overlap and high inter-sexual overlap provides males with greater access to females. We encourage additional research on the influence of prey availability on conspecific spatial overlap in snakes as well as the behavioral mechanisms responsible for maintaining the low levels of overlap we observed.