Classical theory suggests that parasites will exhibit higher fitness in sympatric relative to allopatric host populations (local adaptation). However, evidence for local adaptation in natural host–parasite systems is often equivocal, emphasizing the need for infection experiments conducted over realistic geographic scales and comparisons among species with varied life history traits. Here, we used infection experiments to test how two trematode (flatworm) species ( Paralechriorchis syntomentera and Ribeiroia ondatrae ) with differing dispersal abilities varied in the strength of local adaptation to their amphibian hosts. Both parasites have complex life cycles involving sequential transmission among aquatic snails, larval amphibians and vertebrate definitive hosts that control dispersal across the landscape. By experimentally pairing 26 host-by-parasite population infection combinations from across the western USA with analyses of host and parasite spatial genetic structure, we found that increasing geographic distance—and corresponding increases in host population genetic distance—reduced infection success for P .  syntomentera , which is dispersed by snake definitive hosts. For the avian-dispersed R. ondatrae , in contrast, the geographic distance between the parasite and host populations had no influence on infection success. Differences in local adaptation corresponded to parasite genetic structure; although populations of P .  syntomentera exhibited ~10% mtDNA sequence divergence, those of R. ondatrae were nearly identical (<0.5%), even across a 900 km range. Taken together, these results offer empirical evidence that high levels of dispersal can limit opportunities for parasites to adapt to local host populations.

In the spring and summer of 2020, we found cysts of the nematode Eustrongylides spp. in Brown Trout (Salmo trutta), Rainbow Trout (Oncorhynchus mykiss), and Mountain Whitefish (Prosopium williamsoni) in the Deschutes River. The prevalence and maximum intensity of infection exceeded the range of published accounts for Eustrongylides spp. across a wide range of species. Because the 1st intermediate host is likely to be a tubificid worm, also known as a sludge worm or sewage worm, this exceptional prevalence of Eustrongylides spp. infection raises questions about conditions in the river that may be contributing to the high infection rate in fish.

Classical theory suggests that parasites will exhibit higher fitness in sympatric relative to allopatric host populations (local adaptation). However, evidence for local adaptation in natural host-parasite systems is often equivocal, emphasizing the need for infection experiments conducted over realistic geographic scales and comparisons among species with varied life history traits. Here, we used infection experiments to test how two trematode (flatworm) species (Paralechriorchis syntomentera and Ribeiroia ondatrae) with differing dispersal abilities varied in the strength of local adaptation to their amphibian hosts. Both parasites have complex life cycles involving sequential transmission among aquatic snails, larval amphibians and vertebrate definitive hosts that control dispersal across the landscape. By experimentally pairing 26 host-by-parasite population infection combinations from across the western USA with analyses of host and parasite spatial genetic structure, we found that increasing geographic distance-and corresponding increases in host population genetic distance-reduced infection success for P. syntomentera, which is dispersed by snake definitive hosts. For the avian-dispersed R. ondatrae, in contrast, the geographic distance between the parasite and host populations had no influence on infection success. Differences in local adaptation corresponded to parasite genetic structure; although populations of P. syntomentera exhibited similar to 10% mtDNA sequence divergence, those of R. ondatrae were nearly identical (<0.5%), even across a 900 km range. Taken together, these results offer empirical evidence that high levels of dispersal can limit opportunities for parasites to adapt to local host populations.

Abstract Published accounts describe the eggs of spotted frogs (Rana luteiventris and Rana pretiosa) as having either 1 or 2 jelly envelopes surrounding the ovum. Eggs of R. luteiventris are typically reported to have a single jelly envelope, whereas those of R. pretiosa are reported to have 2 envelopes. We found, however, that both species' eggs actually have 3 jelly envelopes. In this paper we describe the eggs of both species, and identify the origin of the discrepancy in the scientific literature.

Despite the important roles of freshwater gastropods in aquatic ecosystems, the taxonomic status of many taxa is unclear, which is compounded by a lack of information on species population genetic structuring, distribution, and dispersal patterns. The objective of this study was to address the biogeography of the freshwater snailPlanorbella trivolvis(Gastropoda: Planorbidae) in the western United States. We amplified two genetic markers (16S, COI) from individuals belonging to western USA populations and downloaded genetic data from GenBank. We utilized minimum spanning networks to assess the genetic patterns and performed Analysis of Molecular Variance and linear regression analyses to determine how geographic distance and watershed identity contributed to the observed genetic structuring. For both markers, we found that the majority of genetic variation was associated within and among populations, rather than among watersheds. Correspondingly, there was no significant effect of geographic distance on genetic distance, suggesting that long-distance dispersal was promoting gene flow between populations. The genetic similarity could reflect avian-mediated dispersal of snails along the Pacific Flyway, a major waterfowl migratory corridor. Further analysis of the population structuring across North America revealed East-West genetic structuring, suggesting that across longitudinal gradientsP. trivolvisexperiences significant genetic isolation.

Relatively few North American anurans overwinter in water and information is sparse on their movement from overwintering habitat to breeding sites. Oregon spotted frogs (Rana pretiosa) breed explosively in early spring and often overwinter submerged at sites that are distanced from breeding habitats. In montane parts of their range, wintering and breeding habitats can remain frozen for months. We investigated timing, duration, and potential cues for R. pretiosa migrations from a wintering lake near the Cascade Mountains in central Oregon, U.S.A. First and median migrant males moved slightly earlier than females. Onset of migration was as early as February 12 (males) and as late as April 4 (females) in years of mild and extended winters, respectively. Frogs were active at water temperatures below those associated with early breeding activities in one lowland R. pretiosa population. Higher proportions of frogs migrated before ice-out in years of prolonged winter conditions. Migrations were temporally compressed in years of later movement. This migration ‘rush’, along with the ability to move at cold temperatures and to vary timing of migrations likely helps montane R. pretiosa deal with colder and more variable spring conditions than lowland populations.

The Oregon Spotted Frog (Rana pretiosa) is endemic to the Pacific Northwest and was recently listed as threatened under the Endangered Species Act. We tested the hypothesis that reproductive or physiological stress and parasitic disease may be contributing to the decline of this species. We histologically examined gonads and kidneys of newly metamorphosed wild-caught R. pretiosa to confirm sex and search for evidence of abnormal reproductive development and parasites. A subset of these specimens were also cleared and stained for examination of their skeletal morphology to identify potential skeletal malformations. The sex ratio did not differ significantly from 1:1, and we found no skeletal abnormalities. Trematode metacercarial parasites were present in the kidneys of all Spotted Frogs examined. We also report, for the first time, oocytes developing in the testes of 5 out of the 11 newly metamorphosed male Spotted Frogs examined. Further study into gonadal development of this species is necessary to investigate the significance of testicular oocytes in developing R. pretiosa and to identify whether these gonadal abnormalities are related in any way to their decline.

Placobdella sophieae Oceguera-Figueroa et al., 2010 (Hirudinida: Glossiphoniidae) is reported from Oregon, California, and British Columbia for the first time. New hosts reported for P. sophieae include Taricha granulosa (rough-skinned newt), Rana pretiosa (Oregon spotted frog), and Anaxyrus boreas (western toad). Placobdella sophieae exhibits relatively low host specificity and all amphibians occurring in the Pacific Northwest are potential hosts.

Parasite infections often lead to dramatically different outcomes among host species. Although an emerging body of ecoimmunological research proposes that hosts experience a fundamental trade-off between pathogen defences and life-history activities, this line of inquiry has rarely been extended to the most essential outcomes of host-pathogen interactions: namely, infection and disease pathology. Using a comparative experimental approach involving 13 amphibian host species and a virulent parasite, we test the hypothesis that 'pace-of-life' predicts parasite infection and host pathology. Trematode exposure increased mortality and malformations in nine host species. After accounting for evolutionary history, species that developed quickly and metamorphosed smaller ('fast-species') were particularly prone to infection and pathology. This pattern likely resulted from both weaker host defences and greater adaptation by parasites to infect common hosts. Broader integration between life history theory and disease ecology can aid in identifying both reservoir hosts and species at risk of disease-driven declines.

Summary 1. An emerging framework in animal disease ecology seeks to ‘decompose’ a host’s response to disease into resistance, or its ability to resist infection following exposure, and tolerance, or its ability to limit the damage associated with infection. How these processes vary over the life history of a host, however, and whether developmental changes in resistance and tolerance account for ‘critical windows’ of disease vulnerability remain open questions. 2. Critical developmental windows are particularly important for infections that alter host development. Recently, increased observations of amphibians with severe limb malformations have stimulated debate over the causes responsible and whether malformation types can be used to infer the agent responsible. The trematode parasite Ribeiroia ondatrae, for example, is often implicated in accounts of extra-legged frogs, but is believed to be unimportant in explaining missing legged animals. Here, we test the influence of host developmental stage, from eggs to post-metamorphosis, on the risk of mortality and the types of malformations produced in Pacific chorus frogs (Pseudacris regilla) following exposure to trematode infection. 3. Consistent with a critical window of vulnerability, host mortality and malformations were greatest among animals exposed during pre-limb and early limb development (15–90%) and decreased to <5% with progressive development. Early stage animals also exhibited a higher frequency of missing limbs, whereas extra limbs and limb elements developed predominantly among tadpoles exposed after limb development was initiated. Hosts infected later in limb development were normal or exhibited only minor outgrowths and abnormal skin webbings. 4. Increases in host tolerance rather than host resistance largely explained the observed changes in pathology. Prior to host metamorphosis, parasites exhibited comparable success invading host tissue, but the amount of resulting damage differed significantly as a function of host size and developmental stage. Following metamorphosis hosts were significantly more resistant to infections, however. 5. These findings highlight the importance of critical developmental windows for infectious diseases and underscore the role of developmental changes in host tolerance in controlling this process. Forecasted changes in climate, for example, have enormous potential to influence both the timing and intensity of host–parasite interactions in nature.

minima, egg mass, oviposition More than half of the known remaining populations of the Oregon Spotted Frog (Rana pretiosa) are at elevations .1200 m along the Cascade Range and its eastern flank in Oregon (Pearl and Hayes 2005). Rana pretiosa in this region typically breed soon after thaw and often lay eggs in water ,20 cm deep (Pearl and others 2009). Egg masses in shallow microhabitats experience broad temperature fluctuations, and ice formation on the surface is common (Bull and Shepherd 2003). At .20 oviposition sites around Sunriver in central Oregon (elevation 1270 m), we have observed high survival of R. pretiosa embryos after exposure to subfreezing air temperatures and ice cover for up to several days. These observations and the characteristics of oviposition sites across much of the species’ range along the Cascade Range suggest that R. pretiosa eggs in this region may be more tolerant of low temperatures than previously reported. Information on thermal limits of R. pretiosa embryos derives from studies of 1 population in the Fraser River Valley of southwestern British Columbia. The low elevation of this site (approximately 50 m above sea level) and its proximity to the Strait of Georgia and Pacific Ocean make its climate more moderate than much of the extant range of R. pretiosa. Licht (1971) reported a lethal minimum, the temperature at which egg survival is ,50%, near 66C for eggs from that site at a range of stable temperature treatments in the laboratory. A subset of embryos survived lower temperatures for up to 8 h, after which trials were terminated (Licht 1971), but data were not presented on presence or timing of develop

We used genetic methods to estimate the effective number of breeders (N b) in a population of Rana pretiosa, an imperiled amphibian in western North America. Microsatellite data was gathered from large samples of adults, eggs, and juveniles collected in 2006. We wished to determine where in the life cycle the greatest reductions in N b occur, and to compare genetic estimates of N b to an egg mass count estimate of the number of breeding adults. We predicted that N b estimated at the metamorph stage would be reduced by increased variance in family size due to egg mass mortality. Contrary to our prediction, estimates of N b at the egg and metamorph stages were similar. Thus, we found no evidence of inflated variance in family size between the two stages. If our results for this population are typical for R. pretiosa, then increased variance in family size during the egg to metamorph stage may not be a strong factor in reducing the effective population sizes (N e) relative to the census sizes (N) in this species.

Renewed controversy has emerged over the likely causes and consequences of deformed amphibians, particularly those with missing limbs. The results of a series of experiments by Ballengée and Sessions (2009) implicate aquatic predators (i.e. dragonfly larvae) in causing such abnormalities. Skelly and Benard (2010), however, argued that the small scale of these experiments and the absence of a correlation between predator abundance and deformity frequencies in natural amphibian populations undermine such a conclusion. Drawing upon our experiences with frog malformations, we suggest that the study of amphibian deformities has been hindered by two, interrelated problems. First, empirical studies often fail to critically define the expected baseline level of abnormalities and differentiate between "epidemic" and "endemic" frequencies of malformations. Second, recognizing the likelihood of multiple causes in driving amphibian malformations, continued research needs to embrace a "multiple lines of evidence" approach that allows for complex etiologies by integrating field surveys, diagnostic pathology, comparative modeling, and experiments across a range of ecological scales. We conclude by highlighting the results of a recent study that uses this approach to identify the role of aquatic predators (i.e., fishes and dragonflies) in causing high frequencies of deformed frogs in Oregon. By combining long-term data, comparative data and mechanistic experiments, this study provides compelling evidence that certain predators do cause deformities under ecologically relevant conditions. In light of continuing concerns about amphibian deformities and population declines, we emphasize the need to integrate ecological, epidemiological, and developmental tools in addressing such environmental enigmas.

Amphibian chytridiomycosis, caused by the fungus Batrachochytrium dendrobatidis (Bd), has been implicated in the decline and extinction of amphibian species worldwide, in addition to catastrophic losses of animals in captivity. Conservation of threatened amphibians, including captive breeding and maintenance of animals in zoos, research facilities, and private collections, requires effective control of pathogens. Several chemical compounds, including Formalite III®, itraconazole, and chloramphenicol, have been used to treat amphibians infected with Bd, with varying levels of success. Here, we report successful clearance of Bd in five species of post-metamorphic anurans and one caudate species using terbinafine hydrochloride (HCl) in alcohol, which is available over the counter as Lamisil AT™ (Novartis Pharmaceuticals Inc., New York, NY). Treatments consisting of 5 min soak in fresh 0.01% or 0.005% terbinafine HCl in alcohol for either five consecutive days or for six treatments spread across 10 days successfully cleared Bd from 100% of 81 test subjects in eight trials. Our results indicate that terbinafine HCl in alcohol has a high therapeutic index as a treatment for Bd infection in living post-metamorphic amphibians.

While many predators completely consume their prey, others feed only on blood or tissue without killing the prey, sometimes causing ecologically significant levels of injury. We investigated the importance of sublethal predator attacks in driving an emerging issue of conservation importance: missing-limb deformities in amphibians. We combined long-term field data and manipulative experiments to evaluate the role of sublethal predation in causing abnormalities in two regions of central Oregon, U.S.A. Since 1988, western toads (Bufo boreas) in Lake Aspen have exhibited abnormalities dominated by partially missing limbs and digits at annual frequencies from <1% to 35%. On Broken Top volcano, we found comparable types and frequencies of abnormalities in Cascades frogs (Rana cascadae). Field sampling and observational data implicated two aquatic predators in these abnormality phenomena: introduced sticklebacks (Gasterosteus aculeatus) at Lake Aspen and corduliid dragonfly larvae (Somatochlora albicincta) at Broken Top. In experiments, these predators produced limb abnormalities identical to those observed in the respective regions. At Lake Aspen, in situ predator exclosures effectively eliminated abnormalities in toads, while comparisons among years with low and high stickleback abundance and between wetlands with and without sticklebacks reinforced the link between fish and amphibian abnormalities. Neither trematode parasite infection nor pesticide contamination could explain observed abnormalities. Our results suggest that predators are an important explanation for missing-limb abnormalities and highlight the ecological significance of sublethal predation in nature.

The pathogen Batrachochytrium dendrobatidis (Bd) has been associated with amphibian declines in multiple continents, including western North America. We investigated Bd prevalence in Oregon spotted frog (Rana pretiosa), a species that has declined across its range in the Pacific Northwest. Polymerase chain reaction analysis of skin swabs indicated that Bd was prevalent within populations (420 of 617 juvenile and adults) and widespread among populations (36 of 36 sites) where we sampled R. pretiosa in Oregon and Washington. We rarely detected Bd in R. pretiosa larvae (2 of 72). Prevalence of Bd in postmetamorphic R. pretiosa was inversely related to frog size. We found support for an interactive effect of elevation and sampling date on Bd: prevalence of Bd generally increased with date, but this effect was more pronounced at lower elevations. We also found evidence that the body condition of juvenile R. pretiosa with Bd decreased after their first winter. Our data indicate that some Oregon spotted frog populations are currently persisting with relatively high Bd prevalence, but the risk posed by Bd is unknown.