Tracy A. G. Rittenhouse

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.

Effects of Timber Harvest on Amphibian Populations: Understanding Mechanisms from Forest Experiments

Harvesting timber is a common form of land use that has the potential to cause declines in amphibian populations. It is essential to understand the behavior and fate of individuals and the resulting consequences for vital rates (birth, death, immigration, emigration) under different forest management conditions. We report on experimental studies conducted in three regions of the United States to identify mechanisms of responses by pond-breeding amphibians to timber harvest treatments. Our studies demonstrate that life stages related to oviposition and larval performance in the aquatic stage are sometimes affected positively by clearcutting, whereas effects on juvenile and adult terrestrial stages are mostly negative. Partial harvest treatments produced both positive and weaker negative responses than clearcut treatments. Mitigating the detrimental effects of canopy removal, higher surface temperature, and loss of soil-litter moisture in terrestrial habitats surrounding breeding ponds is critical to maintaining viable amphibian populations in managed forested landscapes.

DISTRIBUTION OF AMPHIBIANS IN TERRESTRIAL HABITAT SURROUNDING WETLANDS

Estimating the distribution of amphibians in terrestrial habitats surrounding wetlands is essential for determining how much habitat is required to maintain viable amphibian populations and how much habitat may be allocated to other land use practices. We apply univariate kernel estimation in a new manner to determine the distribution of amphibians during the non-breeding season. We summarized data from 13 radio telemetry studies that reported net maximum distance traveled from the breeding site for each individual (n = 404 individuals), and calculated a univariate kernel density estimate for all data combined. Kernel density estimation provides a function for the probability of an amphibian being present at a given distance from the breeding site and bootstrap methods allow for error estimates of isopleth values. Amphibians generally occurred at a short distance from the wetland (50% isopleth was at 93 m) and declined at greater distances (95% isopleth was at 664 m); however, use of habitat immediately adjacent (e.g., < 30 m) to the breeding site was lower than the peak for all species. The shape of the distribution was consistent for frogs and salamanders; however, the 95% kernel isopleth for the salamander estimate (245 m) was less than half the distance of the frog estimate (703 m), indicating that frogs distributed themselves at much greater distances from the breeding site than salamanders. Kernel estimates for the two western species, Rana luteiventris and Bufo boreas, did not peak near the breeding site as in the other species, suggesting that non-breeding habitat for these species is not located near breeding sites. We were unable to detect a statistical difference between sexes, but females tend to use habitat at greater distances from the wetland than males. Our results revealed that amphibians are not uniformly distributed in terrestrial habitats surrounding wetlands.

Demographic Consequences of Terrestrial Habitat Loss for Pool‐Breeding Amphibians: Predicting Extinction Risks Associated with Inadequate Size of Buffer Zones

Much of the biodiversity associated with isolated wetlands requires aquatic and terrestrial habitat to maintain viable populations. Current federal wetland regulations in the United States do not protect isolated wetlands or extend protection to surrounding terrestrial habitat. Consequently, some land managers, city planners, and policy makers at the state and local levels are making an effort to protect these wetland and neighboring upland habitats. Balancing human land-use and habitat conservation is challenging, and well-informed land-use policy is hindered by a lack of knowledge of the specific risks of varying amounts of habitat loss. Using projections of wood frog (Rana sylvatica) and spotted salamander (Ambystoma maculatum) populations, we related the amount of high-quality terrestrial habitat surrounding isolated wetlands to the decline and risk of extinction of local amphibian populations. These simulations showed that current state-level wetland regulations protecting 30 m or less of surrounding terrestrial habitat are inadequate to support viable populations of pool-breeding amphibians. We also found that species with different life-history strategies responded differently to the loss and degradation of terrestrial habitat. The wood frog, with a short life span and high fecundity, was most sensitive to habitat loss and isolation, whereas the longer-lived spotted salamander with lower fecundity was most sensitive to habitat degradation that lowered adult survival rates. Our model results demonstrate that a high probability of local amphibian population persistence requires sufficient terrestrial habitat, the maintenance of habitat quality, and connectivity among local populations. Our results emphasize the essential role of adequate terrestrial habitat to the maintenance of wetland biodiversity and ecosystem function and offer a means of quantifying the risks associated with terrestrial habitat loss and degradation.

The Role of Microhabitats in the Desiccation and Survival of Anurans in Recently Harvested Oak–Hickory Forest

Abstract Juvenile survival is an important life history feature, because recent modeling efforts suggest that modest changes in juvenile survival rates due to habitat change may greatly affect population growth rates. We compared water loss and survival rates of recently metamorphosed American Toad (Anaxyrus americanus), Green Frog (Lithobates clamitans), and Wood Frog (Lithobates sylvaticus) juveniles contained within four microhabitats, two of which occur in uncut control forest (i.e., forest ridgetop, forest drainage) and two within recently harvested forest (i.e., clearcut open, clearcut brushpile). Survival was higher in forest drainage than forest ridgetop, indicating that microhabitats within continuous forest are not equally suitable. Brushpiles of coarse woody debris reduced desiccation risks in clearcuts as indicated by survival differences between clearcut open and clearcut brushpile and survival in clearcut open was very low for all species in both years. We found species differences in survival as well as a species by microhabitat interaction in water loss rates. These results are best explained by observed behavioral differences as opposed to physiological differences among species. We conclude that desiccation can be a major source of mortality for juveniles entering terrestrial habitats, especially habitat altered by anthropogenic land-use. Desiccation risks are greatest in areas with low soil moisture conditions, which for our study included microhabitats within clearcuts without coarse woody debris, forested ridgetops, and years with below average rainfall.

EFFECTS OF TIMBER HARVESTING ON POND-BREEDING AMPHIBIAN PERSISTENCE: TESTING THE EVACUATION HYPOTHESIS

Numerous studies have documented the decline of amphibians following timber harvest. However, direct evidence concerning the mechanisms of population decline is lacking and hinders attempts to develop conservation or recovery plans and solutions for forest species. We summarized the mechanisms by which abundance of amphibians may initially decline following timber harvest into three testable hypotheses: (1) mortality, (2) retreat, and (3) evacuation. Here, we tested the evacuation hypothesis within a large-scale, replicated experiment. We used drift fences with pitfall traps to capture pond-breeding amphibians moving out of experimental clearcut quadrants and into control quadrants at four replicate arrays located within the Daniel Boone Conservation Area on the upper Ozark Plateau in Warren County, Missouri, USA. During the preharvest year of 2004, only 51.6% of the 312 individuals captured were moving out of pre-clearcut quadrants, and movement did not differ from random. In contrast, during both postharvest years of 2005 and 2006, the number of captures along the quadrant edge increased, and a higher proportion of individuals (59.9% and 56.6%, respectively, by year) were moving out of clearcut quadrants than entering. Salamanders moved out of clearcuts in large percentages (Ambystoma annulatum, 78.2% in 2005, 78.2% in 2006; A. maculatum, 64.0% in 2005, 57.1% in 2006). Frogs and toads also moved out of clearcut quadrants, but in lower percentages (Bufo americanus, 59.6% in 2005, 53.3% in 2006; Rana clamitans, 52.7% in 2006). Salamanders moved out of clearcuts with low-wood treatments more than out of clearcuts with high-wood treatments. Movement of salamanders out of clearcuts was independent of sex. Estimated movement out of clearcuts represented between 8.7% and 35.0% of the total breeding adults captured for two species of salamanders. Although we recognize that some portion of the amphibian population may retreat underground for short periods and others may not survive the effects of timber harvest, these data are the first direct evidence showing that individuals are capable of leaving clearcuts and shifting habitat use.

Survival costs associated with wood frog breeding migrations: effects of timber harvest and drought

Migration presents a trade-off for individuals between the potential fitness benefits of reaching high-quality habitat vs. the potential costs of migration. Within an information-theoretic framework, we examined the costs of migration for adult wood frogs (Rana sylvatica) in response to timber harvest and annual weather conditions using Cox proportional-hazard estimates of survival. In 2004 prior to timber harvest, survival did not differ between the inside (0.75, SE = 0.078) and outside (0.73, SE = 0.235) of the circular timber harvest arrays (each 164 m radius). Following timber harvest, survival inside arrays in both 2005 and 2006 (0.22, SE = 0.065; 0.42, SE = 0.139) was lower than survival outside of the arrays and prior to harvest. Sources of mortality included predation in all years and desiccation in the drought year of 2005. The most-supported models for explaining both predation and desiccation risks reflected behaviors as opposed to timber harvest or weather conditions. Both predation and desiccation risks increased when frogs made frequent movements or were located near breeding ponds. Optimal behaviors for reducing predation and desiccation risks were the same before and after timber harvest; however, the survival consequences for not adopting these behaviors were more severe following harvest. Our results provide empirical evidence for (1) the ecological pressures that influence migratory behavior and (2) differential survival in relation to migratory behavior which reveals why frogs move relatively long distances away from breeding sites.

Leaf litter input mediates tadpole performance across forest canopy treatments.

Understanding the mechanisms limiting the distributions of organisms is necessary for predicting changes in community composition along habitat gradients. In many areas of the USA, land originally cleared for agriculture has been undergoing a process of reforestation, creating a gradient of canopy cover. For small temporary wetlands, this gradient can alter abiotic conditions and influence the resource base of wetland food webs by affecting litter inputs. As distributions of amphibians and many other temporary wetland taxa correlate with canopy cover, we experimentally manipulated shade levels and litter types in pond mesocosms to explore mechanisms limiting species performance in wetlands with canopy cover. Most differences between ponds were mediated by litter type rather than direct effects of shading. Although all three amphibian species tested are open-canopy specialists, spring peepers were the only species to show decreased survival in shaded ponds. Pond litter type generally had strong effects on growth and development rates, with tadpoles of two species in grass litter ponds growing to twice the size of, and metamorphosing 7 days earlier than, those in leaf litter ponds. Contrary to our initial hypothesis, shade level and litter type showed very few significant interactions. Our results indicate that the effects of shading cannot be considered in isolation of vegetation changes in pond basins when evaluating the effects of forest succession on temporary pond communities.

Postbreeding Habitat Use of Wood Frogs in a Missouri Oak-Hickory Forest

Abstract Fitness benefits to individuals from using a particular habitat during the non-breeding season are likely species- and habitat-specific. Our goal was to define the postbreeding habitat use of adult Wood Frogs (Rana sylvatica) within continuous oak-hickory forest in Missouri. We used radio-telemetry to determine whether adult Wood Frogs are evenly spaced throughout this forest type or clumped at a particular resource. In addition, we determined microhabitat selection using conditional logistic regression that compares the microhabitat at frog locations to paired points located 2 m from the frog. Adult frogs migrated from breeding sites located on ridgetops into ephemeral, rocky ravines. Use of drainages by Wood Frogs depended on the distance between the breeding site and drainage, and the orientation of drainages relative to the pond edge influenced whether migratory paths of frogs are funneled or spaced apart. The most supported model of microhabitat selection indicated that frogs selected locations with increased leaf litter depth and air temperature and with decreased humidity and light compared to paired points. Persistence of Wood Frog populations along the southwestern edge of their range requires successful annual migrations between breeding sites and forested drainages, which are important nonbreeding habitat for Wood Frogs in a Missouri oak-hickory forest.

Breeding and Recruitment Phenology of Amphibians in Missouri Oak-Hickory Forests

ABSTRACT Knowing the phenologies of species in a region helps guide management and conservation activities in breeding ponds and surrounding terrestrial habitats. We examined the phenology of pond-breeding amphibians in central Missouri oak-hickory forests. Two ponds were monitored for 4 y from 2000–2003 and five other ponds were monitored for 1 y during 2004 using drift fences with pitfall traps. We found 15 species of pond-breeding amphibians, nine of which we captured in sufficient abundance to evaluate breeding phenology. Among the nine species, breeding migrations occur from Feb. to Nov., while subsequent metamorph emigration occurred primarily from May to Oct.. Our ponds were nearly permanent, resulting in salamander-dominated communities in these oak-hickory forests. Pond-use was partitioned by species that differed in their timing of reproduction and post-metamorphic emergence. For example, breeding in the fall gives larval ringed salamanders a potential size advantage over the spring-breeding spotted salamander larvae. However, the fall breeding strategy requires ponds with long hydroperiods that hold water continuously from Aug. through May. Green frogs and central newts also required long hydroperiods for their larval stage (>160 d). American toads, however, are adapted to exploit ponds with shorter, less predictable hydroperiods as they only required ponds to hold water for as little as 60 d. Management for multiple species of pond-breeding amphibians in a landscape benefits from the inclusion of wetlands with a variety of hydroperiods.