No abstract

Summary1. Snakes often occur in species-rich assemblages, and sympatry is thought to be facilitated primarily by low diet overlap, not interspecific interactions. 2. We selected, a priori, three species pairs consisting of species that are morphologically and taxonomically similar and may therefore be likely to engage in interspecific, consumptive competition. We then examined a large-scale database of snake detection/nondetection data and used occupancy modelling to determine whether these species occur together more or less frequently than expected by chance while accounting for variation in detection probability among species and incorporating important habitat categories in the models. 3. For some snakes, we obtained evidence that the probabilities that habitat patches are used are influenced by the presence of potentially competing congeneric species. Specifically, timber rattlesnakes (Crotalus horridus) were less likely than expected by chance to use areas that also contained eastern diamond-backed rattlesnakes (Crotalus adamanteus) when the proportion of evergreen forest was relatively high. Otherwise, they occurred together more often than expected by chance. Complex relationships were revealed between habitat use, detection probabilities and occupancy probabilities of North American racers (Coluber constrictor) and coachwhips (Coluber flagellum) that indicated the probability of competitive exclusion increased with increasing area of grassland habitat, although there was some model uncertainty. Cornsnakes (Pantherophis guttatus or Pantherophis slowinskii) and ratsnakes (Pantherophis alleghaniensis, Pantherophis spiloides, or Pantherophis obsoletus) exhibited differences in habitat selection, but we obtained no evidence that patterns of use for this species pair were influenced by current interspecific interactions. 4. Overall, our results are consistent with the hypothesis that competitive interactions influence snake assemblage composition; the strength of these effects was affected by landscapescale habitat features. Furthermore, we suggest that current interspecific interactions may influence snake occupancy, challenging the paradigm that contemporary patterns of snake co-occurrence are largely a function of diet partitioning that arose over evolutionary time.

Habitat loss and degradation are thought to be the primary drivers of species extirpations, but for many species we have little information regarding specific habitats that influence occupancy. Snakes are of conservation concern throughout North America, but effective management and conservation are hindered by a lack of basic natural history information and the small number of large‐scale studies designed to assess general population trends. To address this information gap, we compiled detection/nondetection data for 13 large terrestrial species from 449 traps located across the southeastern United States, and we characterized the land cover surrounding each trap at multiple spatial scales (250‐, 500‐, and 1000‐m buffers). We used occupancy modeling, while accounting for heterogeneity in detection probability, to identify habitat variables that were influential in determining the presence of a particular species. We evaluated 12 competing models for each species, representing various hypotheses pertaining to important habitat features for terrestrial snakes. Overall, considerable interspecific variation existed in important habitat variables and relevant spatial scales. For example, kingsnakes (Lampropeltis getula) were negatively associated with evergreen forests, whereas Louisiana pinesnake (Pituophis ruthveni) occupancy increased with increasing coverage of this forest type. Some species were positively associated with grassland and scrub/shrub (e.g., Slowinski's cornsnake, Elaphe slowinskii) whereas others, (e.g., copperhead, Agkistrodon contortrix, and eastern diamond‐backed rattlesnake, Crotalus adamanteus) were positively associated with forested habitats. Although the species that we studied may persist in varied landscapes other than those we identified as important, our data were collected in relatively undeveloped areas. Thus, our findings may be relevant when generating conservation plans or restoration goals. Maintaining or restoring landscapes that are most consistent with the ancestral habitat preferences of terrestrial snake assemblages will require a diverse habitat matrix over large spatial scales.

Fire-maintained woodlands and savannas are important ecosystems for vertebrates in many regions of the world. These ecosystems are being restored by forest managers, but little information exists on herpetofaunal responses to this restoration in areas dominated by shortleaf pine (Pinus echinata). We compared habitat characteristics and herpetofaunal communities in restored pine woodlands to relatively unmanaged, second-growth forests in the Ouachita Mountains of western Arkansas, USA. We found woodland restoration with periodic burning affected species differently; some species benefited, some species appeared negatively affected, but most species did not respond clearly either way. Overall reptile captures were significantly (p = 0.041) greater in pine-woodlands than in unrestored forest; one species of snake and three species of lizards were captured more often in woodlands than unrestored forests. Among anurans, we found no significant difference in captures between woodlands and unrestored forests for any species. Among salamanders, we captured western slimy salamanders (Plethodon albagula) almost exclusively in unrestored forest, but captures of other species did not differ between the two treatments. Historically, the Ouachita region likely consisted of a mosaic that included both fire-maintained habitats (woodlands, savannas, and prairies) and areas of denser forest on mesic sites that were less likely to burn. Consequently, landscapes that retain both open woodlands and denser, less-intensely burned forest (in the form of unharvested greenbelts or separate stands) would likely promote and maintain a greater diversity of herpetofauna.

Over a 13-year period we examined the mortality of cavity trees (n = 453) used by red-cockaded woodpeckers (Picoides borealis) on national forests in eastern Texas. Bark beetles (53%), wind snap (30%), and fire (7%) were the major causes of cavity tree mortality. Bark beetles were the major cause of mortality in loblolly (Pinus taeda) and shortleaf (P. echinata) pines, whereas fire was the major cause in longleaf pines (P. palustris). Cavity trees on the Angelina National Forest (NF) were dying at a higher rate than new, complete cavities were being excavated. Cavity enlargement by pileated woodpeckers (Dryocopus pileatus) on the Angelina NF was substantial, with 20% (49/ 249) of the cavity trees being enlarged over 7 years. To reduce cavity tree mortality, site disturbances in cluster areas (e.g., midstory control, prescribed burning, thinning) should be minimized during years when southern pine beetle (Dendroctonus frontalis) populations are elevated. Careful planning of timber cutting to avoid funneling wind into cluster areas might reduce wind damage to cavity trees.

No abstract

Red-cockaded Woodpeckers (P&ides borealis) nest and forage in pine-dominated forests. Research indicates that substantial hardwood midstory encroachment is detrimental to Red-cockaded Woodpecker populations, although the exact mechanisms are unknown. We examined foraging behavior in relation to midstory between August 1989 and February 1990. Red-cockaded Woodpeckers foraged at greater heights in areas of taller and denser midstory in the loblolly-shortleaf pine (Pirzus rueda and P. echimra, respectively) habitat, but not in longleaf pine (P. palustris) habitat with less-developed midstory vegetation than typical of loblolly-shortleaf pine habitat. In addition, Red-cockaded Woodpeckers concentrated foraging activities in or adjacent to forest stands or openings with reduced midstory vegetation. Overall, Red-cockaded Woodpeckers foraged disproportionately at heights and sites that minimized their exposure to dense midstory conditions. These results suggest that ecosystem management, preferably using prescribed fire, that reduces midstory vegetation will improve foraging habitat for Red-cockaded Woodpeckers.

We evaluated selection of nest sites by male Red-cockaded Woodpeckers (Picoides borealis) in Texas relative to the age of the cavity when only cavities excavated by the woodpeckers were available and when both naturally excavated cavities and artificial cavities were available. We also evaluated nest-cavity selection relative to the ability of naturally excavated cavity trees to produce resin, which is used by the woodpeckers to maintain a barrier against predation by rat snakes (Elaphe spp.). Longleaf pines (Pinus palustris) selected by breeding males as nest trees produced significantly greater resin yields at 2, 8, and 24 h post-wounding than cavity trees used for roosting by other group members. This preference was observed in loblolly pine (P. taeda) and shortleaf pine (P. echinata) cavity trees only at the 2-h resin-sampling period. When only naturally excavated cavities were available, Red-cockaded Woodpeckers in both longleaf pine and loblolly-shortleaf pine habitat selected the newest cavities available for their nest sites, possibly as a means to reduce parasite loads. When both naturally excavated and artificial cavity inserts were available, Red-cockaded Woodpeckers continued to select the newest cavity for nesting in loblolly-shortleaf pine habitat but not in longleaf pine habitat. Resin production in existing longleaf pine nest trees remained sufficient for continued use, whereas resin production in loblolly pine and shortleaf pine nest trees decreased through time, probably because of woodpecker activity at resin wells. For these latter tree species, breeding males switched to newer cavities and/or cavity trees with higher resin yields.