Susan C. Walls

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.

Trends in Amphibian Occupancy in the United States

Though a third of amphibian species worldwide are thought to be imperiled, existing assessments simply categorize extinction risk, providing little information on the rate of population losses. We conducted the first analysis of the rate of change in the probability that amphibians occupy ponds and other comparable habitat features across the United States. We found that overall occupancy by amphibians declined 3.7% annually from 2002 to 2011. Species that are Red-listed by the International Union for Conservation of Nature (IUCN) declined an average of 11.6% annually. All subsets of data examined had a declining trend including species in the IUCN Least Concern category. This analysis suggests that amphibian declines may be more widespread and severe than previously realized.

Do geographically isolated wetlands influence landscape functions

Geographically isolated wetlands (GIWs), those surrounded by uplands, exchange materials, energy, and organisms with other elements in hydrological and habitat networks, contributing to landscape functions, such as flow generation, nutrient and sediment retention, and biodiversity support. GIWs constitute most of the wetlands in many North American landscapes, provide a disproportionately large fraction of wetland edges where many functions are enhanced, and form complexes with other water bodies to create spatial and temporal heterogeneity in the timing, flow paths, and magnitude of network connectivity. These attributes signal a critical role for GIWs in sustaining a portfolio of landscape functions, but legal protections remain weak despite preferential loss from many landscapes. GIWs lack persistent surface water connections, but this condition does not imply the absence of hydrological, biogeochemical, and biological exchanges with nearby and downstream waters. Although hydrological and biogeochemical connectivity is often episodic or slow (e.g., via groundwater), hydrologic continuity and limited evaporative solute enrichment suggest both flow generation and solute and sediment retention. Similarly, whereas biological connectivity usually requires overland dispersal, numerous organisms, including many rare or threatened species, use both GIWs and downstream waters at different times or life stages, suggesting that GIWs are critical elements of landscape habitat mosaics. Indeed, weaker hydrologic connectivity with downstream waters and constrained biological connectivity with other landscape elements are precisely what enhances some GIW functions and enables others. Based on analysis of wetland geography and synthesis of wetland functions, we argue that sustaining landscape functions requires conserving the entire continuum of wetland connectivity, including GIWs.

Morphological variation in a larval salamander: dietary induction of plasticity in head shape

We examined diet-dependent plasticity in head shape in larvae of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Larvae in some populations of this species exhibit trophic polymorphism, with some individuals possessing exaggerated trophic features characteristic of a cannibalistic morphology in larval Ambystoma; e.g. a disproportionately broad head and hypertrophied vomerine teeth. We hypothesized that 1) head shape variation results from feeding upon different types of prey and that 2) cannibal morphs are induced by consumption of conspecifics. To induce variation, we fed three groups of larvae different diets: 1) brine shrimp nauplii only; 2) nauplii plus anuran tadpoles; 3) nauplii, tadpoles and conspecific larval salamanders. Comparisons of size (mass)-adjusted means revealed that this manipulation of diet induced significant variation in six measures of head shape, but not in the area of the vomerine tooth patch. For five of the six head traits, larvae that ate tadpoles and brine shrimp nauplii developed significantly broader, longer and deeper heads than did larvae that only ate brine shrimp nauplii. The ingestion of conspecifics, in addition to nauplii and tadpoles, significantly altered two head traits (interocular-width and head depth), compared to larvae only fed nauplii and tadpoles. Canonical discriminant function analysis detected two statistically reliable canonical variables: head depth was most highly associated with the first canonical variable, whereas three measures of head width (at the jaws, gills and eyes) and interocular width were most highly associated with the second canonical variable. Despite this diet-enhanced morphological variation, there was no indication that any of the three types of diet (including conspecific prey) induced the exaggerated trophic features of the “cannibal” morph in this species. These results illustrate that ingestion of different types of prey contributes to plasticity in head shape, but that some other proximate cue(s), either alone or in combination with diet variation, is essential to induce the extremes of trophic polymorphism in this species.

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.

Geographically isolated wetlands: Rethinking a misnomer

We explore the category “geographically isolated wetlands” (GIWs; i.e., wetlands completely surrounded by uplands at the local scale) as used in the wetland sciences. As currently used, the GIW category (1) hampers scientific efforts by obscuring important hydrological and ecological differences among multiple wetland functional types, (2) aggregates wetlands in a manner not reflective of regulatory and management information needs, (3) implies wetlands so described are in some way “isolated,” an often incorrect implication, (4) is inconsistent with more broadly used and accepted concepts of “geographic isolation,” and (5) has injected unnecessary confusion into scientific investigations and discussions. Instead, we suggest other wetland classification systems offer more informative alternatives. For example, hydrogeomorphic (HGM) classes based on well-established scientific definitions account for wetland functional diversity thereby facilitating explorations into questions of connectivity without an a priori designation of “isolation.” Additionally, an HGM-type approach could be used in combination with terms reflective of current regulatory or policymaking needs. For those rare cases in which the condition of being surrounded by uplands is the relevant distinguishing characteristic, use of terminology that does not unnecessarily imply isolation (e.g., “upland embedded wetlands”) would help alleviate much confusion caused by the “geographically isolated wetlands” misnomer.

Drought, deluge and declines: the impact of precipitation extremes on amphibians in a changing climate.

The Class Amphibia is one of the most severely impacted taxa in an on-going global biodiversity crisis. Because amphibian reproduction is tightly associated with the presence of water, climatic changes that affect water availability pose a particularly menacing threat to both aquatic and terrestrial-breeding amphibians. We explore the impacts that one facet of climate change—that of extreme variation in precipitation—may have on amphibians. This variation is manifested principally as increases in the incidence and severity of both drought and major storm events. We stress the need to consider not only total precipitation amounts but also the pattern and timing of rainfall events. Such rainfall “pulses” are likely to become increasingly more influential on amphibians, especially in relation to seasonal reproduction. Changes in reproductive phenology can strongly influence the outcome of competitive and predatory interactions, thus potentially altering community dynamics in assemblages of co-existing species. We present a conceptual model to illustrate possible landscape and metapopulation consequences of alternative climate change scenarios for pond-breeding amphibians, using the Mole Salamander, Ambystoma talpoideum, as an example. Although amphibians have evolved a variety of life history strategies that enable them to cope with environmental uncertainty, it is unclear whether adaptations can keep pace with the escalating rate of climate change. Climate change, especially in combination with other stressors, is a daunting challenge for the persistence of amphibians and, thus, the conservation of global biodiversity.

Density dependence in a larval salamander: The effects of interference and food limitation

I examined the roles of food limitation and aggressive interference in densitydependent growth and survival in larvae of the spotted salamander, Ambystoma maculatum. In a laboratory experiment, I raised larvae at a low density, low food level; high density, low food level; and high density, high food level. An increased density of larvae fed low food levels significantly decreased mean body size yet increased the size ratio of the largest:smallest larva (a factor that may promote cannibalism). High density also significantly decreased production of metamorphs, although time to metamorphosis did not differ among the three treatments. Food supplementation did not ameliorate the effects of increased density on these variables. Survival was significantly reduced due to an increase in the proportion of larvae cannibalized at the high density. Food supplementation reduced the incidence of cannibalism and improved survival, although it was inadequate to enhance growth significantly. Behavioral trials revealed no difference in aggression across the three treatments, suggesting that it was not a major factor in the observed responses. Rather, densitydependent survival was more a consequence of cannibalism resulting from food limitation. These results provide experimental evidence that, for this species, food limitation may be of greater importance than interference as a mechanism of density dependence.

Effects of conservation practices on wetland ecosystem services in the Mississippi Alluvial Valley

Restoration of wetland ecosystems is an important priority for many state and federal agencies, as well as nongovernmental conservation organizations. The historic conversion of wetlands in the Mississippi Alluvial Valley (MAV) has resulted in large-scale implementation of a variety of conservation practices designed to restore and enhance wetland ecosystem services. As a consequence, the effectiveness of multiple approaches in achieving desired conservation goals varies depending on site conditions, practices employed, and specific ecosystem services. We reviewed government agency programs and the scientific literature to evaluate the effects of conservation practices on wetlands in the MAV. There were 68 different conservation practices applied to a combined total of 1.27 million ha in the MAV between 2000 and 2006. These practices fell into two categories: Wetland Conservation Practices and Upland Conservation Practices. Sixteen different practices accounted for nearly 92% of the total area, and only three of these are directly related to wetlands: Wetland Wildlife Habitat Management, Wetland Restoration, and Riparian Forest Buffer. All three of these practices involve reforestation, primarily planting hard-mast species such as Quercus sp. and Carya sp. These plantings are likely to develop into even-aged stands of low tree diversity with little structural heterogeneity, which will impact future wildlife habitat. Since hydrology is a critical driver of wetland processes, the ability of a given conservation practice to restore wetland hydrology is a key determinant of how well it can restore ecosystem services. However, there is little to no follow-up monitoring of projects, so it is difficult to know how much variability exists for any given practice or the efficacy of specific practices. Conservation practices that only plant trees without reconnecting the wetland to the hydrologic and nutrient fluxes in the watershed may restore some wildlife habitat but will do little for regulating services such as nitrogen retention. While conservation practices have overall beneficial effects on many ecosystem services in the MAV, the most effective are those with a direct link between the actions associated with a given practice and controls over ecosystem processes and services.

Occurrence of Oral Deformities in Larval Anurans

Abstract We quantified deformities in the marginal papillae, tooth rows, and jaw sheaths of tadpoles from 13 population samples representing three families and 11 sites in the southeastern United States. Oral deformities were observed in all samples and in 13.5–98% of the specimens per sample. Batrachochytrium dendrobatidis (chytrid) infections were detected in three samples. There was high variability among samples in the pattern and number of discovered deformities. Pairwise associations between oral structures containing deformities were nonrandom for several populations, especially those with B. dendrobatidis infections or high total numbers of deformities. Comparisons of deformities among samples using multivariate analyses revealed that tadpole samples grouped together by family. Analyses of ordination indicated that three variables, the number of deformities, the number of significant associations among deformity types within populations, and whether populations were infected with B. dendrobatidis, were significantly correlated with the pattern of deformities. Our data indicate that the incidence of oral deformities can be high in natural populations and that phylogeny and B. dendrobatidis infection exert a strong influence on the occurrence and type of oral deformities in tadpoles.