Marc J. Mazerolle

Improving data analysis in herpetology: using Akaike's Information Criterion (AIC) to assess the strength of biological hypotheses

In ecology, researchers frequently use observational studies to explain a given pattern, such as the number of individuals in a habitat patch, with a large number of explanatory (i.e., independent) variables. To elucidate such relationships, ecologists have long relied on hypothesis testing to include or exclude variables in regression models, although the conclusions often depend on the approach used (e.g., forward, backward, stepwise selection). Though better tools have surfaced in the mid 1970s, they are still underutilized in certain fields, particularly in herpetology. This is the case of the Akaike information criterion (AIC) which is remarkably superior in model selection (i.e., variable selection) than hypothesis- based approaches. It is simple to compute and easy to understand, but more importantly, for a given data set, it provides a measure of the strength of evidence for each model that represents a plausible biological hypothesis relative to the entire set of models considered. Using this approach, one can then compute a weighted average of the estimate and standard error for any given variable of interest across all the models considered. This procedure, termed model-averaging or multimodel inference, yields precise and robust estimates. In this paper, I illustrate the use of the AIC in model selection and inference, as well as the interpretation of results analysed in this framework with two real herpetological data sets. The AIC and measures derived from it is should be routinely adopted by herpetologists.

Making Great Leaps Forward: Accounting for Detectability in Herpetological Field Studies

Abstract Detecting individuals of amphibian and reptile species can be a daunting task. Detection can be hindered by various factors such as cryptic behavior, color patterns, or observer experience. These factors complicate the estimation of state variables of interest (e.g., abundance, occupancy, species richness) as well as the vital rates that induce changes in these state variables (e.g., survival probabilities for abundance; extinction probabilities for occupancy). Although ad hoc methods (e.g., counts uncorrected for detection, return rates) typically perform poorly in the face of no detection, they continue to be used extensively in various fields, including herpetology. However, formal approaches that estimate and account for the probability of detection, such as capture-mark-recapture (CMR) methods and distance sampling, are available. In this paper, we present classical approaches and recent advances in methods accounting for detectability that are particularly pertinent for herpetological data sets. Through examples, we illustrate the use of several methods, discuss their performance compared to that of ad hoc methods, and we suggest available software to perform these analyses. The methods we discuss control for imperfect detection and reduce bias in estimates of demographic parameters such as population size, survival, or, at other levels of biological organization, species occurrence. Among these methods, recently developed approaches that no longer require marked or resighted individuals should be particularly of interest to field herpetologists. We hope that our effort will encourage practitioners to implement some of the estimation methods presented herein instead of relying on ad hoc methods that make more limiting assumptions.

AMPHIBIAN ROAD MORTALITY IN RESPONSE TO NIGHTLY VARIATIONS IN TRAFFIC INTENSITY

Pond-breeding amphibians frequently encounter roads during their movements across the landscape to reach their breeding, summering, or hibernation sites. Through night driving surveys conducted each summer from 1995–2002 on a 20-km stretch of secondary road within a national park of eastern Canada, I evaluated whether road traffic had cumulative effects on amphibian abundance over this period. I also investigated the effect of nightly variations in traffic intensity on the number of amphibians killed on the road. I recorded a total of 4643 amphibian crossing events during the 37 surveys. I did not detect any decreasing trend in abundance for amphibian roadside populations over the 8 years. The number of dead American toads (Bufo americanus) increased with increasing traffic intensity. The number of ranid frogs (Rana clamitans, R. pipiens, and R. sylvatica) dead on the road was greatest when many individuals were moving on the road and at moderate traffic intensities (approximately 10–18 vehicles/h). In contrast, spring peepers (Pseudacris crucifer) dead on the road increased with decreasing traffic intensity. The number of ambystomatid salamanders (Ambystoma laterale and A. maculatum) dead on the road, on the other hand, did not respond to traffic intensity. Nonetheless, results indicate that subtle variations in traffic intensity as those observed in this study (i.e., 5–26 vehicles/h) can increase mortality on the road for certain amphibian species. Future studies in landscape ecology should routinely consider measures associated with roads, such as the proximity and density of roads, when investigating amphibian abundance patterns in wetlands.

LANDSCAPE CHARACTERISTICS INFLUENCE POND OCCUPANCY BY FROGS AFTER ACCOUNTING FOR DETECTABILITY

2 Departement de phytologie, Pavillon Paul-Comtois, UniversiteLaval, Quebec, Quebec, G1K 7P4, Canada Abstract. Many investigators have hypothesized that landscape attributes such as the amount and proximity of habitat are important for amphibian spatial patterns. This has produced a number of studies focusing on the effects of landscape characteristics on am- phibian patterns of occurrence in patches or ponds, most of which conclude that the land- scape is important. We identified two concerns associated with these studies: one deals with their applicability to other landscape types, as most have been conducted in agricultural landscapes; the other highlights the need to account for the probability of detection. We tested the hypothesis that landscape characteristics influence spatial patterns of amphibian occurrence at ponds after accounting for the probability of detection in little-studied peatland landscapes undergoing peat mining. We also illustrated the costs of not accounting for the probability of detection by comparing our results to conventional logistic regression anal- yses. Results indicate that frog occurrence increased with the percent cover of ponds within 100, 250, and 1000 m, as well as the amount of forest cover within 1000 m. However, forest cover at 250 m had a negative influence on frog presence at ponds. Not accounting for the probability of detection resulted in underestimating the influence of most variables on frog occurrence, whereas a few were overestimated. Regardless, we show that conven- tional logistic regression can lead to different conclusions than analyses accounting for detectability. Our study is consistent with the hypothesis that landscape characteristics are important in determining the spatial patterns of frog occurrence at ponds. We strongly recommend estimating the probability of detection in field surveys, as this will increase the quality and conservation potential of models derived from such data.

BEHAVIOR OF AMPHIBIANS ON THE ROAD IN RESPONSE TO CAR TRAFFIC

Nocturnal car traffic often results in amphibian casualties, especially during rainy nights. The behavior of amphibians presumably influences their vulnerability to mortality on the road, but this hypothesis remains untested. We investigated the behavioral response of individuals of six species of amphibians on roads when confronted by an approaching vehicle. We first conducted a field study consisting of 50 night-driving surveys over 4 yr during which we recorded the behavior (i.e., moving or immobile) of frogs, toads, tree frogs, and salamanders encountered on a 20-km stretch of road. In an effort to tease apart the effects of headlights and the sound of motors on amphibian behavior, we carried out a field experiment on a test road where we exposed individuals to different car-associated stimuli. Here, we tested the hypothesis that simultaneous exposure to headlights and the sound of a car motor would elicit a stronger response than exposure to a single stimulus or a control. Based on the observations of the 2767 individuals in the field survey, immobility was the most common response to the approach of a car (mean probability of 0.82 of remaining immobile); the response differed across species but depended on the season of the survey (May–June vs. July–September). Similarly, the 91 individuals included in the field experiment were more likely to move during the control treatment than during any of the car-associated treatments. The combined stimuli elicited the strongest response, followed by the headlights-only and the motor-only treatments. Spring peepers (Pseudacris crucifer) tended to move more often than the other species we tested in the field experiment, which suggests they spend less time on the road and are less vulnerable to traffic mortality than other species. Both the field survey and field experiment consistently indicated that amphibians tend to remain immobile at the approach of a vehicle. This behavior highlights the vulnerability of amphibians to road traffic and should be considered in measures to mitigate road impacts.

Amphibian Activity, Movement Patterns, and Body Size in Fragmented Peat Bogs

I investigated the activity, direction of movement, and body size (snout-vent length) of am- phibians in both pristine and fragmented bogs of southeastern New Brunswick. I used drift-fences with pitfall traps to capture amphibians in six pristine bogs and six bogs undergoing peat mining (i.e, bog fragments) in 1997 and 1998. Results indicate that seasonal activity patterns of amphibians in bogs peak during August and correspond to movements of adults (following breeding) and juveniles (after metamor- phosis) from adjacent wetlands. A seasonal shift in species composition occurred, as most captures early in the season consisted almost exclusively of ranids, with an increase in salamander captures in late summer and fall. Climatic variables generally explained more of the variation in amphibian activity in fragments than in pristine bogs. Wood frog activity near fragment edges was more dependent on amount of precipi- tation than in pristine bogs. Wood frog and green frog movements were nonrandomly oriented relative to mined fragment edges. Orientation of leopard frog movements was strongly influenced by year. Wood frogs occurring in fragments were larger than those in pristine bogs. The size difference in green frogs was not significant but followed the same patterns as wood frogs. Leopard frogs within bog fragments were larger than those in pristine bogs but only in 1998. This study implies that peat mining influences amphibian activity and movement patterns in neighboring bog fragments. Larger individuals may be better suited for survival in disturbed environments, such as mined bogs, because they are less sensitive to desiccation than smaller ones.

Drainage Ditches Facilitate Frog Movements in a Hostile Landscape

Ditches are common in landscapes influenced by agricultural, forestry, and peat mining activities, and their value as corridors remains unassessed. Pond-breeding amphibians can encounter hostile environments when moving between breeding, summering, or hibernation sites, and are likely to benefit from the presence of ditches in the landscape. Within a system consisting of ditch networks in bogs mined for peat in eastern New Brunswick, Canada, I quantified the breeding, survival, and movements of green frogs (Rana clamitans melanota) in drainage ditches and also surveyed peat fields. Frogs rarely ventured on peat fields and most individuals frequented drainage ditches containing water, particularly in late summer. Though frogs did not breed in ditches, their survival rate in ditches was high (88%). Ditches did not hinder frog movements, as frogs moved independently of the current. Results indicate that drainage ditches containing water enable some movements between habitats isolated by peat mining, in contrast to peat surfaces, and suggest they function as amphibian movement corridors. Thus, such drainage ditches may mitigate the effects of peat extraction on amphibian populations. At the very least, these structures provide an alternative to hostile peat surfaces. This study highlights that small-scale corridors are potentially valuable in population dynamics.

Detrimental effects of peat mining on amphibian abundance and species richness in bogs

Peat bogs of eastern North America are increasingly being mined for peat, which results in the extensive draining of these systems. First, I quantified the effects of peat mining on amphibian species richness and abundance in bog remnants adjacent to mining activity and unmined bogs. I then investigated amphibian occurrence patterns in bog remnants relative to the distance to the mined edge, after accounting for microhabitat and distance to landscape elements. The response of amphibians to peat mining varied across taxa. For instance, amphibian species richness, total amphibian captures (all spp. combined), as well as those of green frogs (Rana clamitans) were greater in unmined bogs than in bog remnants. Wood frogs (Rana sylvatica) were most abundant in unmined bogs in areas far from bog ponds, as well as in unmined bogs in areas near shrub patches. After accounting for microhabitat and landscape variables, American toads (Bufo americanus) in bog remnants were most abundant close to mined edges, whereas the other species did not respond to the proximity of mined edges. Peat mining negatively impacts amphibians occurring in bog remnants. The response of amphibians to peat mining is similar to the one of birds and plants, but differs with small mammals. Maintaining a complex mosaic of bog ponds, shrub and forest patches might help mitigate peat mining effects on amphibian patterns of abundance.

Tracking the sources of polybrominated diphenyl ethers in birds: foraging in waste management facilities results in higher DecaBDE exposure in males.

Differences in feeding ecology are now recognized as major determinants of inter-individual variations in contaminant profiles of free-ranging animals, but exceedingly little attention has been devoted to the role of habitat use. Marked inter-individual variations and high levels of polybrominated diphenyl ethers (PBDEs) (e.g., DecaBDE) have previously been documented in ring-billed gulls (Larus delawarensis) breeding in a colony near Montreal (QC, Canada). However, the environmental sources of these compounds, and thus the reasons causing these large inter-individual variations remain unidentified. In the present study, we used GPS-based telemetry (±5 to 10m precision) to track ring-billed gulls from this colony to reconstruct their movements at the landscape level. We related habitat use of individual gulls (n=76) to plasma concentrations (ng/g ww) and relative contributions (percentages) to Σ38PBDEs of major congeners in the internationally restricted PentaBDE and current-use DecaBDE mixtures. Male gulls that visited waste management facilities (WMFs; i.e., landfills, wastewater treatment plants and related facilities; 25% of all GPS-tracked males) exhibited greater DecaBDE (concentrations and percentages) and lower PentaBDE (percentages) relative to those that did not. In contrast, no such relationships were found in females. Moreover, in males, DecaBDE (concentrations and percentages) increased with percentages of time spent in WMFs (i.e., ~5% of total foraging time), while PentaBDE (percentages) decreased. No relationships between percentages of time spent in other habitats (i.e., urban areas, agriculture fields, and St. Lawrence River) were found in either sex. These findings suggest that animals breeding in the vicinity of WMFs as well as mobile species that only use these sites for short stopovers to forage, could be at risk of enhanced DecaBDE exposure.

Using dynamic N‐mixture models to test cavity limitation on northern flying squirrel demographic parameters using experimental nest box supplementation

Dynamic N-mixture models have been recently developed to estimate demographic parameters of unmarked individuals while accounting for imperfect detection. We propose an application of the Dail and Madsen (2011: Biometrics, 67, 577–587) dynamic N-mixture model in a manipulative experiment using a before-after control-impact design (BACI). Specifically, we tested the hypothesis of cavity limitation of a cavity specialist species, the northern flying squirrel, using nest box supplementation on half of 56 trapping sites. Our main purpose was to evaluate the impact of an increase in cavity availability on flying squirrel population dynamics in deciduous stands in northwestern Québec with the dynamic N-mixture model. We compared abundance estimates from this recent approach with those from classic capture–mark–recapture models and generalized linear models. We compared apparent survival estimates with those from Cormack–Jolly–Seber (CJS) models. Average recruitment rate was 6 individuals per site after 4 years. Nevertheless, we found no effect of cavity supplementation on apparent survival and recruitment rates of flying squirrels. Contrary to our expectations, initial abundance was not affected by conifer basal area (food availability) and was negatively affected by snag basal area (cavity availability). Northern flying squirrel population dynamics are not influenced by cavity availability at our deciduous sites. Consequently, we suggest that this species should not be considered an indicator of old forest attributes in our study area, especially in view of apparent wide population fluctuations across years. Abundance estimates from N-mixture models were similar to those from capture–mark–recapture models, although the latter had greater precision. Generalized linear mixed models produced lower abundance estimates, but revealed the same relationship between abundance and snag basal area. Apparent survival estimates from N-mixture models were higher and less precise than those from CJS models. However, N-mixture models can be particularly useful to evaluate management effects on animal populations, especially for species that are difficult to detect in situations where individuals cannot be uniquely identified. They also allow investigating the effects of covariates at the site level, when low recapture rates would require restricting classic CMR analyses to a subset of sites with the most captures.