J. Hardin Waddle

Quantitative evidence for the effects of multiple drivers on continental-scale amphibian declines

Since amphibian declines were first proposed as a global phenomenon over a quarter century ago, the conservation community has made little progress in halting or reversing these trends. The early search for a “smoking gun” was replaced with the expectation that declines are caused by multiple drivers. While field observations and experiments have identified factors leading to increased local extinction risk, evidence for effects of these drivers is lacking at large spatial scales. Here, we use observations of 389 time-series of 83 species and complexes from 61 study areas across North America to test the effects of 4 of the major hypothesized drivers of declines. While we find that local amphibian populations are being lost from metapopulations at an average rate of 3.79% per year, these declines are not related to any particular threat at the continental scale; likewise the effect of each stressor is variable at regional scales. This result - that exposure to threats varies spatially, and populations vary in their response - provides little generality in the development of conservation strategies. Greater emphasis on local solutions to this globally shared phenomenon is needed.

A new parameterization for estimating co‐occurrence of interacting species

Models currently used to estimate patterns of species co-occurrence while accounting for errors in detection of species can be difficult to fit when the effects of covariates on species occurrence probabilities are included. The source of the estimation problems is the particular parameterization used to specify species co-occurrence probability. We develop a new parameterization for estimating patterns of co-occurrence of interacting species that allows the effects of covariates to be specified quite naturally without estimation problems. In our model, the occurrence of one species is assumed to depend on the occurrence of another, but the occurrence of the second species is not assumed to depend on the presence of the first species. This pattern of co-occurrence, wherein one species is dominant and the other is subordinate, can be produced by several types of ecological interactions (predator-prey, parasitism, and so on). A simulation study demonstrated that estimates of species occurrence probabilities were unbiased in samples of 50-100 locations and three surveys per location, provided species are easily detected (probability of detection > or = 0.5). Higher sample sizes (>200 locations) are needed to achieve unbiasedness when species are more difficult to detect. An analysis of data from treefrog surveys in southern Florida indicated that the occurrence of Cuban treefrogs, an invasive predator species, was highest near the point of its introduction and declined with distance from that location. Sites occupied by Cuban treefrogs were 9.0 times less likely to contain green treefrogs and 15.7 times less likely to contain squirrel treefrogs compared to sites without Cuban treefrogs. The detection probabilities of native treefrog species did not depend on the presence of Cuban treefrogs, suggesting that the native treefrog species are naive to the introduced species.

Modeling the Effect of Toe Clipping on Treefrog Survival: Beyond the Return Rate

Abstract Some studies have described a negative effect of toe clipping on return rates of marked anurans, but the return rate is limited in that it does not account for heterogeneity of capture probabilities. We used open population mark-recapture models to estimate both apparent survival (φ) and the recapture probability (p) of two treefrog species individually marked by clipping 2–4 toes. We used information-theoretic model selection to examine the effect of toe clipping on survival while accounting for variation in capture probability. The model selection results indicate strong support for an effect of toe clipping on survival of Green Treefrogs (Hyla cinerea) and only limited support for an effect of toe clipping on capture probability. We estimate there was a mean absolute decrease in survival of 5.02% and 11.16% for Green Treefrogs with three and four toes removed, respectively, compared to individuals with just two toes removed. Results for Squirrel Treefrogs (Hyla squirella) indicate little support for an effect of toe clipping on survival but may indicate some support for a negative effect on capture probability. We believe that the return rate alone should not be used to examine survival of marked animals because constant capture probability must be assumed, and our examples demonstrate how capture probability may vary over time and among groups. Mark-recapture models provide a method for estimating the effect of toe clipping on anuran survival in situations where unique marks are applied.

Estimating occupancy dynamics in an anuran assemblage from Louisiana, USA

ABSTRACT Effective monitoring programs are designed to track changes in the distribution, occurrence, and abundance of species. We developed an extension of Royle and Kérys (2007) single species model to estimate simultaneously temporal changes in probabilities of detection, occupancy, colonization, extinction, and species turnover using data on calling anuran amphibians, collected from 2002 to 2006 in the Lower Mississippi Alluvial Valley of Louisiana, USA. During our 5-year study, estimates of occurrence probabilities declined for all 12 species detected. These declines occurred primarily in conjunction with variation in estimates of local extinction probabilities (cajun chorus frog [Pseudacris fouquettei], spring peeper [P. crucifer], northern cricket frog [Acris crepitans], Copes gray treefrog [Hyla chrysoscelis], green treefrog [H. cinerea], squirrel treefrog [H. squirella], southern leopard frog [Lithobates sphenocephalus], bronze frog [L. clamitans], American bullfrog [L. catesbeianus], and Fowlers toad [Anaxyrus fowleri]). For 2 species (eastern narrow-mouthed toad [Gastrophryne carolinensis] and Gulf Coast toad [Incilius nebulifer]), declines in occupancy appeared to be a consequence of both increased local extinction and decreased colonization events. The eastern narrow-mouthed toad experienced a 2.5-fold increase in estimates of occupancy in 2004, possibly because of the high amount of rainfall received during that year, along with a decrease in extinction and increase in colonization of new sites between 2003 and 2004. Our model can be incorporated into monitoring programs to estimate simultaneously the occupancy dynamics for multiple species that show similar responses to ecological conditions. It will likely be an important asset for those monitoring programs that employ the same methods to sample assemblages of ecologically similar species, including those that are rare. By combining information from multiple species to decrease the variance on estimates of individual species, our results are advantageous compared to single-species models. This feature enables managers and researchers to use an entire community, rather than just one species, as an ecological indicator in monitoring programs.

Recovery of native treefrogs after removal of nonindigenous Cuban Treefrogs, Osteopilus septentrionalis

Abstract Florida is home to several introduced animal species, especially in the southern portion of the state. Most introduced species are restricted to the urban and suburban areas along the coasts, but some species, like the Cuban Treefrog (Osteopilus septentrionalis), are locally abundant in natural protected areas. Although Cuban Treefrogs are known predators of native treefrog species as both adults and larvae, no study has demonstrated a negative effect of Cuban Treefrogs on native treefrog survival, abundance, or occupancy rate. We monitored survival, capture probability, abundance, and proportion of sites occupied by Cuban Treefrogs and two native species, Green Treefrogs (Hyla cinerea) and Squirrel Treefrogs (Hyla squirella), at four sites in Everglades National Park in southern Florida with the use of capture–mark–recapture techniques. After at least 5 mo of monitoring all species at each site we began removing every Cuban Treefrog captured. We continued to estimate survival, abundance, and occupancy rates of native treefrogs for 1 yr after the commencement of Cuban Treefrog removal. Mark–recapture models that included the effect of Cuban Treefrog removal on native treefrog survival did not have considerable Akaikes Information Criterion (AIC) weight, although capture rates of native species were generally very low prior to Cuban Treefrog removal. Estimated abundance of native treefrogs did increase after commencement of Cuban Treefrog removal, but also varied with the season of the year. The best models of native treefrog occupancy included a Cuban Treefrog removal effect at sites with high initial densities of Cuban Treefrogs. This study demonstrates that an introduced predator can have population-level effects on similar native species.

Diet of the invasive Cuban Treefrog (Osteopilus septentrionalis) in pine rockland and mangrove habitats in South Florida

Abstract. Native to Cuba, the Bahamas and the Cayman Islands, the Cuban Treefrog (CTF) is an invasive species in Florida, with the ability to inflict serious ecological damage to invaded habitats. By examining the diet of the CTF, a known predator of native frogs, better predictions may be made of the impacts on native species and ecosystems. From 2002 – 2003, CTF diet was investigated in south Florida at four sites, two each within pine rockland and mangrove habitat. Within each habitat, one site exhibited a low density of CTFs and the other a high density of CTFs. CTFs were captured in PVC pipes attached to trees and stomach contents were examined after euthanasia. Beetles were the most numerous and widely consumed prey item among sites; roaches, orthopterans, spiders, ants, and caterpillars were also major dietary components. There were significant differences in the proportion of taxa consumed by CTFs between low and high density populations within each habitat, with the low density site in every instance having the higher proportion. Across habitats, ants comprised a significantly higher proportion of the diet in mangroves, whereas beetles, orthopterans, and snails comprised a significantly higher proportion of the diet in pine rocklands. Approximately 3.5% of all stomachs examined contained anuran remains. Though not significant, CTFs from low density sites consumed a higher proportion of frogs than those at high density sites. Corroborating previous research, the data show the CTF to be a generalist feeder, consuming a wide variety of invertebrate prey, with anurans playing only a minor role in the overall diet.

Reproductive Phenology of Three Lizard Species in Costa Rica, with Comments on Seasonal Reproduction of Neotropical Lizards

Abstract We quantified reproductive phenology for three lizard species collected in northeastern Costa Rica over a 14-month sampling period from November 1973 to December 1974. Two species, a gecko (Lepidoblepharis xanthostigma) and a skink (Sphenomorphus cherriei), reproduced continuously at this site, whereas an anole (Norops limifrons) showed a slight decline in reproduction during the dry season. The decrease in reproduction of N. limifrons may reflect a response to moisture stress during the driest part of the year. When seasonal patterns of reproduction among 32 populations of Neotropical lizards with small clutch sizes were analyzed with the effect of phylogenetic relatedness removed, we found that seasonality of reproduction was strongly related to mean annual rainfall but not to number of dry season months.

Changes in Abundance of Gopher Tortoise Burrows at Cape Sable, Florida

Abstract The southernmost population of Gopherus polyphemus (Gopher Tortoises) occurs at Cape Sable, FL. The burrows of this population were surveyed in 1979 using strip transects to estimate Gopher Tortoise burrow density. We present data from a 1990 strip-transect survey and a 2001 line-transect survey of burrows for comparison. We found a significantly lower density of active burrows and a significantly higher density of abandoned burrows in 2001 compared to 1979 and 1990, but the overall density of burrows did not differ significantly over time. The distribution of burrow widths in 2001 was not significantly different from the 1982 distribution, and the bimodal pattern suggests that reproduction has occurred at the site. The 76% decline of active burrows at the site suggests that Gopher Tortoises have been subject to mortality. Reduction of habitat quality and tropical storms are possible explanations for the apparent decline in the Gopher Tortoise population at Cape Sable, but more monitoring is required to understand the long-term trends in this coastal population.

Testing hypotheses on distribution shifts and changes in phenology of imperfectly detectable species

U.S. Geological Survey-Amphibian Research and Monitoring Initiative (ARMI). Grant Number: 505; Autonomous Province of Trento

Anuran site occupancy and species richness as tools for evaluating restoration of a hydrologically-modified landscape

A fundamental goal of wetland restoration is to reinstate pre-disturbance hydrological conditions to degraded landscapes, facilitating recolonization by native species and the production of resilient, functional ecosystems. To evaluate restoration success, baseline conditions need to be determined and a reference target needs to be established that will serve as an ecological blueprint in the restoration process. During the summer wet seasons of 2010 and 2011, we used automated recording units to monitor a community of calling anuran amphibians in the Picayune Strand State Forest of Southwest Florida, USA. This area is undergoing hydrological restoration as part of the Comprehensive Everglades Restoration Plan. We compared occurrence of anurans at sites in the restoration area, to nearby locations in relatively undisturbed habitat (reference sites). We assessed the utility of the latter as restoration targets, using a hierarchical model of community species occupancy to estimate the probability of occurrence of anurans in restoration and reference locations. We detected 14 species, 13 of which were significantly more likely to occur in reference areas. All 14 species were estimated by our model to occur at these sites but, across both years, only 8–13 species were estimated to occur at restoration sites. The composition and structure of these habitats within and adjacent to the Picayune Strand State Forest indicate that they are suitable targets for habitat restoration, as measured by amphibian occurrence and species richness. These areas are important sources for recolonization of anuran amphibians as the hydrologically degraded Picayune Strand undergoes restoration to mitigate the effects of overdrainage and habitat loss.