Benefits of Studies of Overwintering Birds for Understanding Resident Bird Ecology and Promoting Development of Conservation Capacity

Abstract:  Funding of ecological research and monitoring of Neotropical migratory birds on their overwintering grounds has benefited both migratory and permanent-resident species. Using examples from our work in Puerto Rico and the Dominican Republic, we demonstrate that ecological research of overwintering migrants often provides information about the ecology and demography of little-known tropical resident birds. Critically important long-term monitoring in Puerto Rico with a focus on winter residents has provided information on the relationships between annual rainfall and fluctuations in resident bird populations and survival rates. It also has alerted local biologists to declines in resident bird populations, including a decline apparently driven by the entry of a brood parasite. But migrant-focused research may also have had an underappreciated effect on the development of conservation capacity and conservation efforts in host countries. Investments in research on Neotropical migrants overwintering on Hispaniola have resulted in a huge increase in field training of students and wildlife professionals, promoted conservation awareness at local and national levels, played an important role in the growth and professionalization of key environmental organizations, spawned a growing ecotourism industry for bird-watching, and driven national park management planning and conservation efforts for all bird species. We encourage funding organizations and agencies to consider the broader impacts of funding migratory-bird research and monitoring efforts, and we encourage researchers in the tropics to use protocols that provide the most information about all the birds that use the study areas involved and to be aware of important opportunities that they may have to build capacity in host countries.     

Application of Genetic Bottleneck Testing to the Investigation of Amphibian Declines: a Case Study with Natterjack Toads

Abstract: Declines of amphibians are causing widespread concern and in some cases have been explained by factors such as climate change and the spread of microbial pathogens. A problem remains, however, in the unequivocal demonstration of decline in situations where populations may be undergoing natural fluctuations in abundance. We describe the application of a genetic test for bottlenecks ( Cornuet & Luikart 1996 ) that should distinguish between natural oscillations and true population declines. British natterjack toads (   Bufo calamita  ) provide examples of populations that have not declined in recent decades, populations that have declined, and populations in which trends are unclear. Microsatellite allele frequency data from these populations were tested for heterozygote excess and shifts in allele frequency distributions, and inferences from these computations about bottlenecks (i.e., persistently smaller population sizes than the recent means) were compared with demographic information. The genetic test correctly identified bottlenecks and should therefore prove useful in demonstrating whether amphibian declines have occurred where long-term demographic time series are not available.     

The Contribution of Habitat Loss to Changes in Body Size, Allometry, and Bilateral Asymmetry in Two Eleutherodactylus Frogs from Puerto Rico

Abstract:  Amphibian populations have been declining worldwide and the exact mechanisms underlying these changes are not well understood. We examined environmentally induced phenotypic changes that may reflect ongoing stresses on individuals and therefore their ability to persist in increasingly changing landscapes. Specifically, we evaluated the contribution of habitat loss on the size, allometry, and levels of fluctuating asymmetry of Eleutherodactylus antillensis and E. coqui , 2 common species that are endemic to Puerto Rico. We x-rayed frogs collected at 9 sites that differed in the amount of forest cover and measured their snout-vent, radio-ulna, femur, and tibio-fibula lengths. E. antillensis and E. coqui were smaller in the highly disturbed (≤20% forest cover) than in the intermediately (20–70% forest cover) and little-disturbed (≥70% forest cover) landscapes. In E. antillensis but not in E. coqui , the slope and intercept of the curves relating snout-vent length with the length of the 3 bones differed with degree of forest cover, suggesting an effect of habitat loss on body shape. In E. antillensis and E. coqui , differences between right and left sides corresponded to true fluctuating asymmetry; however, only the radio-ulna length of E. coqui showed a trend toward an increase in fluctuating asymmetry with habitat loss. Because body size scales with a variety of physiological, life history, and ecological traits, conservation programs aimed at monitoring morphological changes in amphibians may help in understanding the mechanisms that contribute to their persistence in changing environments.     

Interspecific Variation in Susceptibility of Frog Tadpoles to the Pathogenic Fungus Batrachochytrium dendrobatidis

Abstract:  As part of an overall biodiversity crisis many amphibian populations are in decline throughout the world. Numerous causes have been invoked to explain these declines. These include habitat destruction, climate change, increasing levels of ultraviolet radiation, environmental contamination, and the introduction of non-native species and diseases. Several types of pathogens have been implicated in contributing to amphibian population declines: viruses, bacteria, oomycetes, and fungi. One particular fungus, the chytridiomycete Batrachochytrium dendrobatidis may have caused amphibian population declines in several regions. This pathogen causes chytridiomycosis, which is fatal to newly metamorphic and adult amphibians of certain species. We present experimental evidence that larval stages may also be susceptible to exposure to Batrachochytrium . There was, however, differential sensitivity to B. dendrobatidis in larvae we examined. In laboratory experiments, larval western toads (  Bufo boreas ) exposed to B. dendrobatidis experienced increased mortality and behaviors that suggested they were affected by exposure compared with unexposed control tadpoles. Larvae of Cascades frogs (  Rana cascadae ), bullfrogs ( R. catesbeiana ), and Pacific treefrogs ( Hyla regilla ) did not die after exposure to Batrachochytrium and appeared to behave normally. R. cascadae larvae exposed to B. dendrobatidis , however, showed an increase incidence in mouthpart abnormalities, a characteristic effect of chytridiomycosis, compared with unexposed controls. These results show that Batrachochytrium can negatively affect some species of amphibians at the larval stage and not others. The implications of interspecific variation in susceptibility to fungal infection are broad.     

Mitochondrial DNA Phylogeography and the Conservation of Endangered Lesser Antillean Icterus Orioles

Abstract: Recent natural and anthropogenic disturbances have endangered two of the three oriole species endemic to single islands in the Lesser Antilles. The ongoing eruption of the Soufriere Hills volcano may have doomed the Montserrat Oriole (   Icterus oberi ), whereas high levels of nest parasitism by a cowbird threaten the Martinique Oriole (   I. bonana ). These orioles and related Antillean and Central American forms have been considered part of the Icterus dominicensis superspecies complex, but the taxonomic status of the different Antillean island populations has been long debated. To investigate levels of evolutionary differentiation among threatened Lesser Antillean orioles, we analyzed 2507 nucleotides of protein-coding mitochondrial DNA (mtDNA) sequence from orioles on Martinique, Montserrat, St. Lucia (   I. laudabilis ), Puerto Rico (   I. dominicensis dominicensis ), Mexico (   I. d. prosthemelas ), and three Icterus outgroup species. Phylogenetic analyses of the mtDNA data supported the monophyly of Antillean members of the I. dominicensis complex and identified a star-like pattern of relationship among them. Mitochondrial distances between the Antillean populations were large (4.5–5.8% nucleotide divergence) and suggested that the Lesser Antillean orioles have been isolated evolutionarily from one another since the late Pliocene. The oriole taxa on Montserrat, Martinique, and St. Lucia meet species criteria under the phylogenetic species concept and represent evolutionarily significant units. The impending extinction of the phylogenetically unique Montserrat oriole highlights the vulnerability of island endemics to habitat degradation followed by rare and unpredictable natural catastrophes.     

Variation in Pesticide Tolerance of Tadpoles among and within Species of Ranidae and Patterns of Amphibian Decline

Abstract: There is significant variation among and within amphibian species with respect to reports of population decline; declining species are often found in environments that are physiograpically similar to environments where the same species is thriving. Because variability exists among organisms in their sensitivity to environmental stressors, it is important to determine the degree of this variation when undertaking conservation efforts. We conducted both lethal (time-to-death) and sublethal (activity change) assays to determine the degree of variation in the sensitivity of tadpoles to a pesticide, carbaryl, at three hierarchical levels: among ranid species, among several populations of a single ranid species (   Rana sphenocephala ), and within populations of R. sphenocephala . We observed significant variation in time to death among the nine ranid species and among the 10 R. sphenocephala populations we tested. Four out of eight R. sphenocephala populations exhibited significantly different times to death among families. The magnitude of the activity change in response to exposure to sublethal carbaryl levels was significantly different among species and within R. sphenocephala populations. Chemical contamination, at lethal or sublethal levels, can alter natural regulatory processes such as juvenile recruitment in amphibian populations and should be considered a contributing cause of declines in amphibian populations.     

Effects of Amphibian Chytrid Fungus on Individual Survival Probability in Wild Boreal Toads

Abstract: Chytridiomycosis is linked to the worldwide decline of amphibians, yet little is known about the demographic effects of the disease. We collected capture–recapture data on three populations of boreal toads (Bufo boreas [Bufo = Anaxyrus]) in the Rocky Mountains (U.S.A.). Two of the populations were infected with chytridiomycosis and one was not. We examined the effect of the presence of amphibian chytrid fungus (Batrachochytrium dendrobatidis [Bd]; the agent of chytridiomycosis) on survival probability and population growth rate. Toads that were infected with Bd had lower average annual survival probability than uninfected individuals at sites where Bd was detected, which suggests chytridiomycosis may reduce survival by 31–42% in wild boreal toads. Toads that were negative for Bd at infected sites had survival probabilities comparable to toads at the uninfected site. Evidence that environmental covariates (particularly cold temperatures during the breeding season) influenced toad survival was weak. The number of individuals in diseased populations declined by 5–7%/year over the 6 years of the study, whereas the uninfected population had comparatively stable population growth. Our data suggest that the presence of Bd in these toad populations is not causing rapid population declines. Rather, chytridiomycosis appears to be functioning as a low‐level, chronic disease whereby some infected individuals survive but the overall population effects are still negative. Our results show that some amphibian populations may be coexisting with Bd and highlight the importance of quantitative assessments of survival in diseased animal populations.     

Multiple stressors and amphibian declines: dual impacts of pesticides and fish on yellow-legged frogs.

More than 40% of Earth's 5700+ amphibian species have undergone recent declines. Despite the likely involvement of multiple factors in driving these declines, most studies continue to focus on single stressors. In California (USA), separate studies have implicated either introduced fish or pesticides as causal agents. To date, however, no study has simultaneously evaluated the respective roles of these two potential stressors nor attempted to assess their relative importance, information critical for the development of effective conservation efforts and environmental policies. We examined the role and relative effect of fish and pesticides on the mountain yellow-legged frog (Rana muscosa) using unusually detailed data sets for a large portion of R. muscosa's historic range in California's Sierra Nevada. Habitat characteristics and presence/absence of R. muscosa and fish were quantified at each of 6831 sites during field surveys. Pesticide use upwind of each site was calculated from pesticide application records and predominant wind directions. Using generalized additive models, we found that, after accounting for habitat effects, the probability of R. muscosa presence was significantly reduced by both fish and pesticides, with the landscape-scale effect of pesticides much stronger than that of fish. The degree to which a site was sheltered from the predominant wind (and associated pesticides) was also a significant predictor of R. muscosa presence. Taken together, these results represent the strongest evidence to date that windborne pesticides are contributing to amphibian declines in pristine locations. Our results suggest that amphibian declines may have complex multi-factorial causes, and caution that single-factor studies that demonstrate the importance of one factor should not be used as evidence against the importance of other factors.     

Wood Thrush Populations Are Not All Sinks in the Agricultural Midwestern United States

Abstract: Conservation strategies for forest-breeding, Neotropical migratory birds focus on the identification and preservation of source habitats. In sources, populations maintain relatively high levels of reproductive success, whereas in sinks, reproductive success is consistently below net replacement rate. In agricultural regions of the midwestern United States, forest patches of various sizes appear to act as sinks for Neotropical migrants. In 1994–1996 I monitored Wood Thrush (   Hylocichla mustelina ) nests in four forest patches in an agricultural landscape in northern Indiana. Given the prevailing estimates of adult and juvenile survival, I report productivity data that show substantial annual variation and that are sufficient to suggest the maintenance of temporal sources. Indeed, some Wood Thrush populations may act as source habitats in regions with relatively high levels of Brown-headed Cowbird (   Molothrys ater ) parasitism and nest predation because pairs produce multiple clutches and have the ability to fledge cowbirds and their own young from parasitized nests. My results and those from other studies of Wood Thrushes in landscapes with fragmented forests illustrate the variable ways in which source-sink dynamics are affected by habitat fragmentation. In some agricultural regions, populations in small forest patches may act as sources for Wood Thrushes and may be a legitimate focus of conservation efforts aimed at some migratory songbirds.     

Epidemic Disease and the Catastrophic Decline of Australian Rain Forest Frogs

In the montane rain forests of eastern Australia at least 14 species of endemic, stream-dwelling frogs have disappeared or declined sharply (by more than 90%) during the past 15 years. We review available information on these declines and present eight lines of evidence that collectively suggest that a rapidly spreading, epidemic disease is the most likely responsible agent. The extreme virulence of the putative frog patbogen suggests it is likely exotic to Australian rain forests. We propose that exotic pathogens may be responsible for some recent declines of amphibian populations on other continents and that the intercontinental spread of such pathogens is greatly facilitated by human activities such as the thriving international trade in aquarium fish. Our hypothesis may help explain why some amphibian populations in seemingly pristine environments have mysteriously declined.     

Hypothesis Concerning the Causes of the Disappearance of Boreal Toads from the Mountains of Colorado

Recent reports of world-wide declines and extinctions of amphibian populations have raised questions about the relation of environmental change to the demise of certain amphibian species. Between 1974 and 1982, 11 populations of boreal toads ( Bufo boreas boreas ) in the West Elk Mountains of Colorado totally disappeared. The apparent cause of extinction of these toads was infection with the bacteria Aeromonas hydrophila . In this paper, the presence of disease in declining populations of these toads is used in conjunction with a variety of data from the literature to formulate a working hypothesis for explaining the cause of the decline of this species, and perhaps others:

     

The consequences of poaching and anthropogenic change for forest elephants

Poaching has devastated forest elephant populations (Loxodonta cyclotis), and their habitat is dramatically changing. The long‐term effects of poaching and other anthropogenic threats have been well studied in savannah elephants (Loxodonta africana), but the impacts of these changes for Central Africa's forest elephants have not been discussed. We examined potential repercussions of these threats and the related consequences for forest elephants in Central Africa by summarizing the lessons learned from savannah elephants and small forest elephant populations in West Africa. Forest elephant social organization is less known than the social organization of savannah elephants, but the close evolutionary history of these species suggests that they will respond to anthropogenic threats in broadly similar ways. The loss of older, experienced individuals in an elephant population disrupts ecological, social, and population parameters. Severe reduction of elephant abundance within Central Africa's forests can alter plant communities and ecosystem functions. Poaching, habitat alterations, and human population increase are probably compressing forest elephants into protected areas and increasing human–elephant conflict, which negatively affects their conservation. We encourage conservationists to look beyond documenting forest elephant population decline and address the causes of these declines when developing conversation strategies. We suggest assessing the effectiveness of the existing protected‐area networks for landscape connectivity in light of current industrial and infrastructure development. Longitudinal assessments of the effects of landscape changes on forest elephant sociality and behavior are also needed. Finally, lessons learned from West African elephant population loss and habitat fragmentation should be used to inform strategies for land‐use planning and managing human–elephant interactions.     

Effects of Landscape Composition and Wetland Fragmentation on Frog and Toad Abundance and Species Richness in Iowa and Wisconsin, U.S.A.

Abstract: Management of amphibian populations to reverse recent declines will require defining high-quality habitat for individual species or groups of species, followed by efforts to retain or restore these habitats on the landscape. We examined landscape-level habitat relationships for frogs and toads by measuring associations between relative abundance and species richness based on survey data derived from anuran calls and features of land-cover maps for Iowa and Wisconsin. The most consistent result across all anuran guilds was a negative association with the presence of urban land. Upland and wetland forests and emergent wetlands tended to be positively associated with anurans. Landscape metrics that represent edges and patch diversity also had generally positive associations, indicating that anurans benefit from a complex of habitats that include wetlands. In Iowa the most significant associations with relative abundance were the length of the edge between wetland and forest ( positive) and the presence of urban land (negative). In Wisconsin the two most significant associations with relative abundance were forest area and agricultural area ( both positive). Anurans had positive associations with agriculture in Wisconsin but not in Iowa. Remnant forest patches in agricultural landscapes may be providing refuges for some anuran species. Differences in anuran associations with deep water and permanent wetlands between the two states suggest opportunities for management action. Large-scale maps can contribute to predictive models of amphibian habitat use, but water quality and vegetation information collected from individual wetlands will likely be needed to strengthen those predictions. Landscape habitat analyses provide a framework for future experimental and intensive research on specific factors affecting the health of anurans.     

Inconclusiveness of Chytridiomycosis as the Agent in Widespread Frog Declines

Abstract:  Although there is considerable evidence to support the hypothesis that the chytrid fungus Batrachochytrium dendrobatidis is the primary agent responsible for widespread declines in amphibian populations, particularly rainforest frog populations in Australia and Central America, I argue the case has not yet been made conclusively. Few specimens were collected at the time of population declines, so it may never be possible to conclusively determine their cause. It remains unclear whether the pathogen is novel where declines have occurred. Although it is not necessary that the infection be novel for it to be implicated in declines, if a preexisting pathogen has only recently caused extinctions, cofactors must be important. Whether the pattern of outbreaks represents a "wave" of extinctions is unclear, but if it does, the rate of spread in Australia is implausibly high for a waterborne pathogen, given the most likely estimates of epidemiological parameters. Although B. dendrobatidis is an amphibian pathogen according to Koch's postulates, the postulates are neither necessary nor sufficient criteria to identify a pathogen. The following key pieces of information are necessary to better understand the impact of this fungus on frog communities: better knowledge of the means and rate of transmission under field conditions, prevalence of infection among frog populations, as distinct from morbid individuals, and the effect of the fungus on frogs in the wild. It is crucial to determine whether there are strains of the fungus with differing pathogenicity to particular frog species and whether host-pathogen coevolution has occurred or is occurring. Recently developed diagnostic tools bring into reach the possibility of addressing these questions and thus developing appropriate strategies to manage frog communities that may be affected by this fungus.     

How Broad-Scale Studies of Patterns and Processes Can Serve to Guide Conservation Planning in Africa

Abstract:  Analysis of large-scale biodiversity patterns can uncover general relationships and problems that need to be taken into account when conservation strategies are developed. Nevertheless, these large-scale patterns need to be supplemented with information from local studies that can identify specific problems and determine how the land can be divided between conservation and development interests. I analyzed biodiversity patterns at three different scales to show how various scales of research contributed to conservation planning. A gap analysis for all of sub-Saharan Africa revealed that the network of wildlife reserves provides insufficient protection of narrowly endemic and threatened species, mainly because such species are aggregated in certain areas with dense human populations. A more fine-grained analysis of the distribution of forest birds of eastern Africa generally confirmed the results obtained with coarse-scale data and added precision by identifying forest tracts where conservation actions should be concentrated. Detailed local distribution data for one of the prioritized areas, the Uluguru Mountains in Tanzania, suggest that the actions to halt the loss of biodiversity should be concentrated in the submontane zone, immediately adjacent to densely populated areas. To achieve conservation on the ground, these general planning tools must be supplemented with other kinds of research concerning land-use and local knowledge and with approaches that promote more sustainable development. Different types of institutions will be needed for these different tasks, but it is essential that researchers maintain a dialogue with planners in this area .     

Clinal variation in anuran advertisement calls: basis for acoustic isolation?

Summary The calls of male treefrogs (Eleutherodactylus coqui) were studied along an altitudinal gradient in the Luquillo Mountains in eastern Puerto Rico. The fundamental frequency of each note in the advertisement call was highly negatively correlated with altitude. Although males responded vigorously to playbacks of calls of other males recorded at their own altitude, males at high altitudes responded less frequently to recorded calls of males from low altitudes, and males at low altitudes responded less frequently to recorded calls of males from high altitudes. These results are discussed in relation to potential isolation in contiguous populations of E. coqui.     

Epidemiologic Analysis of Factors Associated with Local Disappearances of Native Ranid Frogs in Arizona

Abstract:  We examined factors that may independently or synergistically contribute to amphibian population declines. We used epidemiologic case–control methodology to sample and analyze a large database developed and maintained by the Arizona Game and Fish Department that describes historical and currently known ranid frog localities in Arizona, U.S.A. Sites with historical documentation of target ranid species ( n = 324) were evaluated to identify locations where frogs had disappeared during the study period (case sites) and locations where frog populations persisted (control sites). Between 1986 and 2003, 117 (36%) of the 324 sites became case sites, of which 105 were used in the analyses. An equal number of control sites were sampled to control for the effects of time. Risk factors, or predictor variables, were defined from environmental data summarized during site surveys and geographic information system data layers. We evaluated risk factors with univariate and multifactorial logistic-regression analyses to derive odds ratios (OR). Odds for local population disappearance were significantly related to 4 factors in the multifactorial model. Disappearance of frog populations increased with increasing elevation (OR = 2.7 for every 500 m, p < 0.01). Sites where disappearances occurred were 4.3 times more likely to have other nearby sites that also experienced disappearances (OR = 4.3, p < 0.01), whereas the odds of disappearance were 6.7 times less (OR = 0.15, p < 0.01) when there was a source population nearby. Sites with disappearances were 2.6 times more likely to have introduced crayfish than were control sites (OR = 2.6, p = 0.04). The identification of factors associated with frog disappearances increases understanding of declines occurring in natural populations and aids in conservation efforts to reestablish and protect native ranids by identifying and prioritizing implicated threats.     

Estimates of species- and ecosystem-level respiration of woody stems along an elevational gradient in the Luquillo Mountains, Puerto Rico

We measured CO₂ efflux from stems of seven subtropical tree species situated along an elevational gradient in the Luquillo Mountains, Puerto Rico and scaled these measurements up to the landscape level based on modeled and empirical relations. The most important determinants of ecosystem stem respiration were species composition and stem temperature. At a species scale, measured CO₂ efflux per unit bole surface area at a given temperature was highest in the early successional species Cecropia schreberiana and lowest in species that inhabit high elevations such as Micropholis garciniifolia and Tabebuia rigida. Carbon dioxide efflux rates followed a diel pattern that lagged approximately 6 h behind changes in sapwood temperatures. At an ecosystem scale, our simulation model indicates a decreasing trend of stem respiration rates with increasing elevation due to shifts in species composition, lower temperatures and reductions in branch surface area. The highest estimated stem respiration rates were present in the lowland tabonuco forest type and the lowest rates were present in the elfin forest type (mean 7.4 and 2.1 Mg C ha⁻¹ yr⁻¹, respectively). There was slight temperature-induced seasonal variation in simulated stem respiration rates at low elevations, with a maximum difference of 19% between the months of February and July. Our results coincide well with those of Odum and Jordan [Odum, H.T., Jordan, C.F., 1970. Metabolism and evapotranspiration of the lower forest in a giant plastic cylinder. In: Odum, H.T., Pigeon, R.F. (Eds.), A Tropical Rain Forest: A Study of Irradiation and Ecology at El Verde, Puerto Rico. U.S. Atomic Energy Commission, Oak Ridge, TN, pp. I165–I189] for the tabonuco forest type and extend their work by presenting estimates and spatial patterns of woody tissue respiration for the entire mountain rather than for a single forested plot.     

Projecting population trends of endangered amphibian species in the face of uncertainty: A pattern-oriented approach

Amphibian populations have been declining worldwide for the last three decades. Determining the risk of extinction is one of the major goals of amphibian conservation, yet few quantitative models have been developed for amphibian populations. Like most rare or threatened populations, there is a paucity of life history data available for most amphibian populations. Data on the critical juvenile life stage are particularly lacking. Pattern oriented modeling (POM) has been used successfully to estimate life history parameters indirectly when critical data lacking, but has not been applied to amphibian populations. We describe a spatially explicit, individual-based, stochastic simulation model developed to project population dynamics of pond-breeding amphibian populations. We parameterized the model with life history and habitat data collected for the endangered Houston toad (Bufohoustonensis), a species for which there is a high degree of uncertainty for juvenile and adult male survival. During model evaluation, we focused on explicitly reducing this uncertainty, evaluating 16 different versions of the model that represented the range of parametric uncertainty for juvenile and adult male survival. Following POM protocol, we compared simulation results to four population-level patterns observed in the field: population size, adult sex ratio, proportion of toads returning to their natal pond, and mean maximum distance moved. Based on these comparisons, we rejected 11 of the 16 model versions. Results of the remaining versions confirmed that population persistence depends heavily on juvenile survival, and further suggested that probability of juvenile survival is likely between 0.0075 and 0.015 (previous estimates ranged from 0.003 to 0.02), and that annual male survival is near 0.15 (previous estimates ranged up to 0.43).