Organisms with complex life cycles may experience diverse stressors during their development. Stressors experienced in early life stages may influence the quantity and quality of individuals in later life stages. However, it is unclear if these effects persist later in life and how they may influence population dynamics. This dissertation uses two amphibian species, the Western spadefoot toad (Pelobates cultripes) and the spotted salamander (Ambystoma maculatum) to explore how biotic and abiotic factors experienced in aquatic and terrestrial environments influence phenotype and survival. We use a combination of field mesoscosm studies, laboratory studies and modeling to explore how early life history stressors persist in diverse environmental contexts. In Chapter 1, pond drying and larval density negatively influence multiple aspects of phenotype in the Western spadefoot toad. In Chapter 2, reduced body size due to larval stressors persisted in the first year of life in spotted salamanders in both high and low terrestrial resource environments. Additionally, there was no relationship between size at metamorphosis and post-metamorphic terrestrial survival. In Chapter 3, low terrestrial moisture levels reduced post-metamorphic growth rates by reducing food intake in juvenile spotted salamanders from both high and low larval density treatments. In Chapter 4, we scale up the effects of reduced body size in the Western spadefoot toad to explore how reduced body size due to pond drying may influence population densities using a stage specific matrix model. Reduced body size, by delaying maturity, may reduce adult densities in the Western spadefoot toad. This dissertation suggests that life stages are highly interconnected, as stressors experienced early in life stage may persist through their effects on phenotype in the absence of compensatory mechanisms. Variation in later life stages may also influence phenotype, but may not completely erase effects of early environments. Stressors experienced early in life may also translate to population level consequences, especially when organisms experience multiple stressors across life stages.

<div>Wetland-dwelling amphibians are of conservation interest for numerous reasons. They serve as biological indicators of water quality during their fully aquatic larval phase, and as carnivores that prey extensively on both terrestrial and aquatic invertebrates. These amphibians are an important link between terrestrial and aquatic food webs, and their wellbeing is an important factor when considering ecosystem health. Amphibians are facing global declines as their wetland habitats are being lost or degraded by human actions. There are efforts to restore wetland habitats, but it is far from certain which practices encourage amphibian occupancy.</div><div>I investigated which factors are important to the persistence of amphibians in restored and naturally formed wetlands to see if restored wetlands can accommodate similar species assemblages. Amphibians were surveyed over two years in a collection of 18 wetlands in Steuben and DeKalb counties, IN owned by The Nature Conservancy. Ambystomatid salamanders were surveyed using plastic minnow traps in springtime, and frogs were surveyed using call surveys in spring and summer. I used linear models to compare wetland plant dominance, wetland hydroperiod, restoration status, distance to nearest mature forest, adjacent forest age and basal area, and inter-wetland distance to amphibian species richness.</div><div>The species richness of Ambystoma salamanders was positively associated with larger wetlands, higher forest basal area, and central mudminnow presence; and negatively associated with older forests, distance to mature forests, and the presence of sunfishes. Ambystoma salamanders besides A. tigrinum were associated with ephemeral hydrology, naturally-formed wetlands, and a greater number of wetlands within one km; and negatively correlated with older forests.</div><div>Frog species richness was positively associated with larger wetland size, and negatively associated with seasonal wetlands, naturally-formed wetlands, distance to nearest mature forests, naturally formed wetlands, treatment for invasive plants, and number of other wetlands within 500m. Total amphibian species richness models did not perform well, but showed a preference for semi-permanent wetlands, smaller distance to mature forests, greater forest basal area, and greater distance between wetlands; and a preference against Scrub Shrub/Forest wetlands. Hylid frogs were negatively correlated with naturally formed wetlands. Ranid frogs were associated semi-permanent wetlands and negatively correlated with the number of other wetlands within 500 m.</div><div>Ambystomatid salamanders were found in restored wetlands, semi-permanent wetlands, and in wetlands containing central mudminnows. Frogs may dislike the disturbance from removing invasive grasses. Managers should factor the disparate habitat requirements of amphibian taxa into their plans for creating and managing restoration projects. Different amphibian groups appear to differ greatly in their habitat requirements, and diverse wetlands may enhance the species richness of an area. Skillfully restored wetlands appear to serve similar functions to original, naturally formed ones.</div>

AN ABSTRACT OF THE DISSERTATION OF GRETCHEN ELIZABETH PADGETT-FLOHR for the Doctor of Philosophy degree in Zoology, presented on March 5, 2009 at Southern Illinois University of Carbondale. TITLE: BATRACHOCHYTRIUM DENDROBATIDIS IN CENTRAL CALIFORNIA AMPHIBIANS MAJOR PROFESSOR: Eric Schauber Amphibian chytridiomycosis has been identified as a disease responsible for the decline and extinction of many amphibian taxa world wide, but little research has been conducted on the disease in Mediterranean climates. To address this gap in the data I studied the amphibian assemblage present across a ~6,475 ha site in central California and investigated the occurrence of the etiological agent, Batrachochytrium dendrobatidis (BD) from organismal, community, landscape and historical perspectives. I initially tested the accuracy and reliability of a proposed diagnostic screening test for BD in four larval species that occur on the site. The screening test proposed by Fellers et al. (2001) and Vredenburg and Summers (2001) consisted of examining larval amphibian mouthparts for abnormalities and or defects, based on their hypothesis that mouthpart defects are clinical signs of BD infection. Sensitivity and specificity of the diagnostic screening test were 76% and 58%, respectively, indicating that the proposed screening test was not a reliable diagnostic test for BD infection for the four species I examined. I conducted controlled laboratory experiments to examine the consequences of BD infection in the two threatened California species that occur on my study site: Rana draytonii and Ambystoma californiense. Both species were susceptible to infection, but all infected animals survived the 18-month study. Infected A. californiense sloughed skin at three times the rate of uninfected salamanders, a pattern that may have long-term energetic costs potentially leading to population-level consequences of sublethal infection by BD. I conducted a retrospective survey of the California Academy of Sciences' (San Francisco, California, USA) amphibian collection, testing for BD in four amphibian species collected from central California between 1897 and 2005 to assess whether the pathogen is novel versus endemic. The earliest detection of BD was in two Rana catesbeiana collected in 1961, and the data support the hypothesis that BD was a novel pathogen introduced into central California prior to 1961 that spread geographically and taxonomically from at least one point of introduction and is now endemic throughout most of central California. I analyzed how environmental factors, amphibian community composition, land use practices, and landscape structure affect the dynamics of the pathogen's distribution on my study site in central California. The distribution of BD in ponds within the landscape varied markedly between years and increases were associated with precipitation, mean minimum and maximum temperatures, and presence of particular species. Pseudacris regilla infection patterns were highly indicative of overall patterns of pond BD status. Fourteen ponds were identified as BD hotpots (BD-positive three of four years). Occurrences of the pathogen within the landscape were spatially autocorrelated and ponds in close proximity to BD hotspots were more likely to test positive. Local land use, (presence/absence of grazing or recreational activity and developed lands), apparently did not influence BD status of a pond. My studies show that BD was likely a novel pathogen introduced into California ca. 1961 that has since become established as an endemic pathogen throughout most of central California. The listed amphibian species that occur in central California can be infected with BD but appear to be resistant to manifesting amphibian chytridiomycosis, and the data from the studies herein could support one of two hypotheses: that natural selection acting over the past 48 years has selected for those individuals that were resistant to the disease; or that the species on my site have always been resistant to BD. The research I conducted further supports the hypothesis that BD is locally vectored by native amphibians (e.g. P. regilla) moving between ponds and that local ecological constraints likely limit vectoring of BD by non-native species. These findings contribute substantially to elucidating and understanding the responses of amphibian populations to disease/pathogen introduction and lay groundwork for future investigations into the host-pathogen-environment relationship as it relates to declining amphibian populations.

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Spotted Salamanders, Ambystoma maculatum, are a widely distributed pond-breeding amphibian that spends an estimated 95% of its life in woodlands surrounding breeding ponds. Their terrestrial ecology remains poorly understood. Few studies have tracked the movement patterns of Spotted Salamanders, and they have given limited information on fine scale patterns. Studies of distribution around vernal pool environments have brought increased attention to their terrestrial ecology. To this end I tracked a sample of Passive Integrated Transponder (PIT) tagged animals with a Radio Frequency Identification Device (RFID). Quadrat-variance methods and a Poisson distribution were used to analyze spatial distribution patterns over 2 years. The home range size of 10 Spotted Salamanders was calculated using the revised minimum area method. A clumped distribution was found at larger spatial scales, except in the first year of the study. Home range size was shown to be limited and highly variable among 10 individuals (Mean=4.29m2, Standard Deviation=6.25).

Because of growing concern of habitat fragmentation and its adverse effects on salamander communities in Appalachian region, sympatric populations of ambystomatid salamanders A. opacum and A. maculatum were studied in Northeast Tennessee to address a number of questions: i) the extent of sexual size dimorphism (SSD) in both species, ii) what traits influence the dorsal skin pigmentation and how, iii) whether gender differences in developmental stability occur, and iv) the extent of phenotypic variation within each species. The findings of this study revealed SSD in both species of salamanders. The most parsimonious statistical model was developed that explained the influence of body mass, dorsal body area, and sex on development of dorsal white pigmentation in marbled salamanders. Data on asymmetry indicate that females are under more stress than males in marbled salamanders, while for spotted salamanders nonsignificant asymmetry is indicative of similar level of stress in both sexes. Data on coefficient of variation (CV) suggest stabilizing selection on optimal body size and mass in female marbled salamanders compared to males; however, for spotted salamanders CV indicates relatively similar selection pressure for body size and mass for both sexes.

Salamanders are one of the most at-risk taxa in the world due to habitat destruction, pollution, climate change, invasive diseases, and more. This has led to a need for conservation breeding programs that are often associated with zoos and aquaria. Salamanders can be difficult to breed in captivity, though, due to their dependence on specific environmental cues and other unknown factors that stimulate sperm and egg production. To overcome these challenges, assisted reproductive technologies (ART) such as exogenous hormone administration, sperm cryopreservation, and in-vitro fertilization have been developed to increase offspring propagation and maintain genetic diversity. If genetically robust populations of salamander species can be sustainably managed ex situ into the future, then their species can be protected in situ through practices such as reintroductions into native habitats. Given the importance of salamanders to healthy, functioning ecosystems, my doctoral research focused on methods to advance caudate conservation efforts through ART. This dissertation addresses four key areas targeting salamander reproduction for conservation: 1) Novel, non-invasive hormone administration routes; 2) Sperm extender toxicity; 3) Novel sperm cryoprotectants; and 4) Application of ART to target salamander species. The objectives were to: 1) Compare nasal, oral, and intramuscular delivery routes of gonadotropin-releasing hormone on spermic response; 2) Compare sperm extenders at varying osmolalities for maintaining sperm quality over time; 3) Test dimethyl sulfoxide versus dimethylformamide as cryoprotectants to increase frozen-thawed sperm viability, motility, and fertilization capability in the eastern tiger salamander (Ambystoma tigrinum); and 4) Transfer ART protocols developed from the eastern tiger salamander to the eastern hellbender (Cryptobranchus alleganiensis) and spotted salamander (Ambystoma maculatum). This work answers critical questions that should help advance salamander ART research into the future and lead to more sustainably managed caudate populations.

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