Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Abstract:  Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species ( Microstegium vimineum, Alliaria petiolata, Berberis thunbergii ) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines in native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.     

A simulation model to evaluate the impacts of invasive earthworms on soil carbon dynamics

Recent studies have reported that earthworm invasions alter native communities and impact nutrient cycling in terrestrial ecosystems. We developed a simulation model to evaluate the potential impacts of earthworm invasions on carbon dynamics, taking into consideration earthworm feeding strategies and priming effects on the microorganisms through their casting activities. Responses of carbon stocks (forest litter, soil organic matter, microbial biomass and earthworm populations) and carbon fluxes (litter decomposition, earthworm consumption, and microbial respiration) were used to evaluate an earthworm invasion of a forest ecosystem. Data from a northern temperate forest (Arnot Forest, New York) were adapted for model calibration and evaluation. Simulation results suggest that the impact and outcome of earthworm invasions are affected by pre-invasion resource availability (litter and soil organic matter), invasive earthworm assemblages (particularly feeding strategy), and invasion history (associated with earthworm population dynamics). The abovementioned factors may also determine invasion progress of earthworm species. The accuracy of the model could be improved by the addition of environmental modules (e.g., soil water regimes), precise parameters accounting for individual species attributes under different environmental conditions (e.g. utilization ability of different types of food resources), as well as earthworm population dynamics (size and structure) and interactions with predators and other invasive/indigenous species during the invasion progress. Such an earthworm invasion model could provide valuable evaluation of the complicated responses of carbon dynamics to earthworm invasions in a range of forest ecosystems, particularly under global change scenarios.     

Reduced Density and Nest Survival of Ground‐Nesting Songbirds Relative to Earthworm Invasions in Northern Hardwood Forests

Abstract: European earthworms (Lumbricus spp.) are spreading into previously earthworm‐free forests in the United States and Canada and causing substantial changes, including homogenization of soil structure, removal of the litter layer, and reduction in arthropod abundance and species richness of understory plants. Whether these changes affect songbirds that nest and forage on the forest floor is unknown. In stands with and without earthworms in the Chequamegon‐Nicolet National Forest, Wisconsin (U.S.A.), we surveyed for, monitored nests of, and measured attributes of habitat of Ovenbirds (Seiurus aurocapillus) and Hermit Thrushes (Catharus guttatus), both ground‐dwelling songbirds, and we sampled earthworms at survey points and nests. Bird surveys indicated significantly lower densities of Ovenbirds and Hermit Thrushes in relation to Lumbricus invasions at survey point and stand extents (3.1 and 15–20 ha, respectively). Modeling of Ovenbird nest survival (i.e., the probability that nestlings successfully fledge) indicated that lower survival probabilities were associated with increased sedge cover and decreased litter depth, factors that are related to Lumbricus invasions, possibly due to reduced nest concealment or arthropod abundance. Our findings provide compelling evidence that earthworm invasions may be associated with local declines of forest songbird populations.     

Patterns of litter disappearance in a northern hardwood forest invaded by exotic earthworms.

A field study was conducted to evaluate the effects of exotic earthworm invasions on the rates of leaf litter disappearance in a northern hardwood forest in southcentral New York, USA. Specifically, we assessed whether differences in litter quality and the species composition of exotic earthworm communities affected leaf litter disappearance rates. Two forest sites with contrasting communities of exotic earthworms were selected, and disappearance rates of sugar maple and red oak litter were estimated in litter boxes in adjacent earthworm-free, transition, and earthworm-invaded plots within each site. After 540 days in the field, 1.7-3 times more litter remained in the reference plots than in the earthworm-invaded plots. In the earthworm-invaded plots, rates of disappearance of sugar maple litter were higher than for oak litter during the first year, but by the end of the experiment, the amount of sugar maple and oak litter remaining in the earthworm-invaded plots was identical within each site. The composition of the earthworm communities significantly affected the patterns of litter disappearance. In the site dominated by the anecic earthworm Lumbricus terrestris and the endogeic Aporrectodea tuberculata, the percentage of litter remaining after 540 days (approximately 17%) was significantly less than at the site dominated by L. rubellus and Octolasion tyrtaeum (approximately 27%). This difference may be attributed to the differences in feeding behavior of the two litter-feeding species: L. terrestris buries entire leaves in vertical burrows, whereas L. rubellus usually feeds on litter at the soil surface, leaving behind leaf petioles and veins. Our results showed that earthworms not only accelerate litter disappearance rates, but also may reduce the differences in decomposition rates that result from different litter qualities at later stages of decay. Similarly, our results indicate that earthworm effects on decomposition vary with earthworm community composition. Furthermore, because earthworm invasion can involve a predictable shift in community structure along invasion fronts or through time, the community dynamics of invasion are important in predicting the spatial and temporal effects of earthworm invasion on litter decomposition, especially at later stages of decay.     

Traits, not origin, explain impacts of plants on larval amphibians

Managing habitats for the benefit of native fauna is a priority for many government and private agencies. Often, these agencies view nonnative plants as a threat to wildlife habitat, and they seek to control or eradicate nonnative plant populations. However, little is known about how nonnative plant invasions impact native fauna, and it is unclear whether managing these plants actually improves habitat quality for resident animals. Here, we compared the impacts of native and nonnative wetland plants on three species of native larval amphibians; we also examined whether plant traits explain the observed impacts. Specifically, we measured plant litter quality (carbon : nitrogen : phosphorus ratios, and percentages of lignin and soluble phenolics) and biomass, along with a suite of environmental conditions known to affect larval amphibians (hydroperiod, temperature, dissolved oxygen, and pH). Hydroperiod and plant traits, notably soluble phenolics, litter C:N ratio, and litter N:P ratio, impacted the likelihood that animals metamorphosed, the number of animals that metamorphosed, and the length of larval period. As hydroperiod decreased, the likelihood that amphibians achieved metamorphosis and the percentage of tadpoles that successfully metamorphosed also decreased. Increases in soluble phenolics, litter N:P ratio, and litter C:N ratio decreased the likelihood that tadpoles achieved metamorphosis, decreased the percentage of tadpoles metamorphosing, decreased metamorph production (total metamorph biomass), and increased the length of larval period. Interestingly, we found no difference in metamorphosis rates and length of larval period between habitats dominated by native and nonnative plants. Our findings have important implications for habitat management. We suggest that to improve habitats for native fauna, managers should focus on assembling a plant community with desirable traits rather than focusing only on plant origin.     

Changes in hardwood forest understory plant communities in response to European earthworm invasions

European earthworms are colonizing earthworm-free northern hardwood forests across North America. Leading edges of earthworm invasion provide an opportunity to investigate the response of understory plant communities to earthworm invasion and whether the species composition of the earthworm community influences that response. Four sugar maple-dominated forest sites with active earthworm invasions were identified in the Chippewa National Forest in north central Minnesota, USA. In each site, we established a 30 x 150 m sample grid that spanned a visible leading edge of earthworm invasion and sampled earthworm populations and understory vegetation over four years. Across leading edges of earthworm invasion, increasing total earthworm biomass was associated with decreasing diversity and abundance of herbaceous plants in two of four study sites, and the abundance and density of tree seedlings decreased in three of four study sites. Sample points with the most diverse earthworm species assemblage, independent of biomass, had the lowest plant diversity. Changes in understory plant community composition were most affected by increasing biomass of the earthworm species Lumbricus rubellus. Where L. rubellus was absent there was a diverse community of native herbaceous plants, but where L. rubellus biomass reached its maximum, the herbaceous-plant community was dominated by Carex pensylvanica and Arisaema triphyllum and, in some cases, was completely absent. Evidence from these forest sites suggests that earthworm invasion can lead to dramatic changes in the understory community and that the nature of these changes is influenced by the species composition of the invading earthworm community.     

Effects of Forest Roads on Macroinvertebrate Soil Fauna of the Southern Appalachian Mountains

Abstract: Many forested landscapes are fragmented by roads, but our understanding of the effects of these roads on the function and diversity of the surrounding forest is in its infancy. I investigated the effect of roads in otherwise continuous forests on the macroinvertebrate fauna of the soil. I took soil samples along transects leading away from the edges of unpaved roads in the Cherokee National Forest in the Southern Appalachian mountains of the United States. Roads significantly depressed both the abundance and the richness of the macroinvertebrate soil fauna. Roads also significantly reduced the depth of the leaf-litter layer. These effects persisted up to 100 m into the forest. Wider roads and roads with more open canopies tended to produce steeper declines in abundance, richness, and leaf-litter depth, but these effects were significant only for canopy cover and litter depth. The macroinvertebrate fauna of the leaf litter plays a pivotal role in the ability of the soil to process energy and nutrients. These macroinvertebrates also provide prey for vertebrate species such as salamanders and ground-foraging birds. The effect of roads on the surrounding forest is compounded by the sprawling nature of the road system in this and many other forests. My data suggest that even relatively narrow roads through forests can produce marked edge effects that may have negative consequences for the function and diversity of the forest ecosystem.     

Effects of Earthworm Invasion on Plant Species Richness in Northern Hardwood Forests

Abstract:  The invasion of non-native earthworms ( Lumbricus spp.) into a small number of intensively studied stands of northern hardwood forest has been linked to declines in plant diversity and the local extirpation of one threatened species. It is unknown, however, whether these changes have occurred across larger regions of hardwood forests, which plant species are most vulnerable, or with which earthworm species such changes are associated most closely. To address these issues we conducted a regional survey in the Chippewa and Chequamegon national forests in Minnesota and Wisconsin (U.S.A.), respectively. We sampled earthworms, soils, and vegetation, examined deer browse in 20 mature, sugar-maple-dominated forest stands in each national forest, and analyzed the relationship between invasive earthworms and vascular plant species richness and composition. Invasion by Lumbricus was a strong indicator of reduced plant richness in both national forests. The mass of Lumbricus juveniles was significantly and negatively related to plant-species richness in both forests. In addition, Lumbricus was a significant factor affecting plant richness in a full model that included multiple variables. In the Chequamegon National Forest earthworm mass was associated with higher sedge cover and lower cover of sugar maple seedlings and several forb species. The trends were similar but not as pronounced in Chippewa, perhaps due to lower deer densities and different earthworm species composition. Our results provide regional evidence that invasion by Lumbricus species may be an important mechanism in reduced plant-species richness and changes in plant communities in mature forests dominated by sugar maples.     

Regional extent of an ecosystem engineer: earthworm invasion in northern hardwood forests.

The invasion of exotic earthworms into northern temperate and boreal forests previously devoid of earthworms is an important driver of ecosystem change. Earthworm invasion can cause significant changes in soil structure and communities, nutrient cycles, and the diversity and abundance of herbaceous plants. However, the regional extent and patterns of this invasion are poorly known. We conducted a regional survey in the Chippewa and Chequamegon National Forests, in Minnesota and Wisconsin, U.S.A., respectively, to measure the extent and patterns of earthworm invasion and their relationship to potential earthworm introduction sites. We sampled earthworms, soils, and vegetation in 20 mature, sugar maple-dominated forest stands in each national forest and analyzed the relationship between the presence of five earthworm taxonomic groups, habitat variables, and distance to the nearest potential introduction site. Earthworm invasion was extensive but incomplete in the two national forests. Four of the six earthworm taxonomic groups occurred in 55-95% of transects; however 20% of all transects were invaded by only one taxonomic group that has relatively minor ecological effects. Earthworm taxonomic groups exhibited a similar sequence of invasion found in other studies: Dendrobaena > Aporrectodea = Lumbricus juveniles > L. rubellus > L. terrestris. Distance to the nearest road was the best predictor of earthworm invasion in Wisconsin while distance to the nearest cabin was the best predictor in Minnesota. These data allow us to make preliminary assessments of landscape patterns of earthworm invasion. As an example, we estimate that 82% of upland mesic hardwood stands in the Wisconsin region are likely invaded by most taxonomic groups while only 3% are unlikely to be invaded at present. Distance to roads and cabins provides a coarse-scale predictor of earthworm invasion to focus stand-level assessments that will help forest managers better understand current and potential forest conditions and identify uninvaded areas that could serve as important refugia for plant species threatened by earthworm invasion.     

Susceptibility of a Northern Hardwood Forest to Exotic Earthworm Invasion

Abstract:  Numerous exotic earthworm species are colonizing northern hardwood forests of North America, where no native earthworms exist. Upon invasion, earthworms have been shown to alter the surface soil environment and plant populations and communities. We sought to identify land-use factors in the Ottawa National Forest (ONF), Michigan (U.S.A.), that contribute to earthworm invasion in forest dominated by sugar maple ( Acer saccharum Marsh.) so that the susceptibility to additional colonization could be evaluated. We sampled earthworm communities in Sylvania Wilderness Area, a unique old-growth hardwood forest, and nonwilderness sites influenced by recreational fishing, recent timber harvesting, or roads. All the nonwilderness sites contained one to five species of exotic earthworms. In contrast, only 50% of wilderness sites contained exotic earthworms, all of a single species. Nonwilderness sites also had thinner litter and duff layers, higher soil C and N content, and higher nitrogen mineralization potentials than Sylvania sites. Two central differences between Sylvania and nonwilderness sites were that all nonwilderness sites were in close contact with roads and had a history of timber harvest, whereas these factors were not present in Sylvania Wilderness Area. Using average rates of colonization, we constructed two geographic information system models to estimate the percentage of sugar maple on the ONF falling within a theoretical 100-year invasion distance of roads and of second-growth sugar maple as relative indices of susceptibility to invasion. Both models indicated high susceptibility to invasion, with 91.7% and 98.9% of sugar maple habitat falling within a theoretical 100-year invasion distance of roads or historical harvests, respectively.     

Effects of Invasive Plants on Arthropods

Non‐native plants have invaded nearly all ecosystems and represent a major component of global ecological change. Plant invasions frequently change the composition and structure of vegetation communities, which can alter animal communities and ecosystem processes. We reviewed 87 articles published in the peer‐reviewed literature to evaluate responses of arthropod communities and functional groups to non‐native invasive plants. Total abundance of arthropods decreased in 62% of studies and increased in 15%. Taxonomic richness decreased in 48% of studies and increased in 13%. Herbivorous arthropods decreased in response to plant invasions in 48% of studies and increased in 17%, likely due to direct effects of decreased plant diversity. Predaceous arthropods decreased in response to invasive plants in 44% of studies, which may reflect indirect effects due to reductions in prey. Twenty‐two percent of studies documented increases in predators, which may reflect changes in vegetation structure that improved mobility, survival, or web‐building for these species. Detritivores increased in 67% of studies, likely in response to increased litter and decaying vegetation; no studies documented decreased abundance in this functional group. Although many researchers have examined effects of plant invasions on arthropods, sizeable information gaps remain, specifically regarding how invasive plants influence habitat and dietary requirements. Beyond this, the ability to predict changes in arthropod populations and communities associated with plant invasions could be improved by adopting a more functional and mechanistic approach. Understanding responses of arthropods to invasive plants will critically inform conservation of virtually all biodiversity and ecological processes because so many organisms depend on arthropods as prey or for their functional roles, including pollination, seed dispersal, and decomposition. Given their short generation times and ability to respond rapidly to ecological change, arthropods may be ideal targets for restoration and conservation activities. Efectos de las Plantas Invasoras sobre los Artrópodos     

Habitat Fragmentation and Species Loss across Three Interacting Trophic Levels: Effects of Life-History and Food-Web Traits

Abstract:  Not all species are likely to be equally affected by habitat fragmentation; thus, we evaluated the effects of size of forest remnants on trophically linked communities of plants, leaf-mining insects, and their parasitoids. We explored the possibility of differential vulnerability to habitat area reduction in relation to species-specific and food-web traits by comparing species–area regression slopes. Moreover, we searched for a synergistic effect of these traits and of trophic level . We collected mined leaves and recorded plant, leaf miner, and parasitoid species interactions in five 100-m² transects in 19 Chaco Serrano woodland remnants in central Argentina. Species were classified into extreme categories according to body size, natural abundance, trophic breadth, and trophic level . Species–area slopes differed between groups with extreme values of natural abundance or trophic specialization. Nevertheless, synergistic effects of life-history and food-web traits were only found for trophic level and trophic breadth: area-related species loss was highest for specialist parasitoids. It has been suggested that species position within interaction webs could determine their vulnerability to extinction. Our results provide evidence that food-web parameters, such as trophic level and trophic breadth, affect species sensitivity to habitat fragmentation .     

Remarkable Amphibian Biomass and Abundance in an Isolated Wetland: Implications for Wetland Conservation

Abstract:  Despite the continuing loss of wetland habitats and associated declines in amphibian populations, attempts to translate wetland losses into measurable losses to ecosystems have been lacking. We estimated the potential productivity from the amphibian community that would be compromised by the loss of a single isolated wetland that has been protected from most industrial, agricultural, and urban impacts for the past 54 years. We used a continuous drift fence at Ellenton Bay, a 10-ha freshwater wetland on the Savannah River Site, near Aiken, South Carolina (U.S.A.), to sample all amphibians for 1 year following a prolonged drought. Despite intensive agricultural use of the land surrounding Ellenton Bay prior to 1951, we documented 24 species and remarkably high numbers and biomass of juvenile amphibians (>360,000 individuals; >1,400 kg) produced during one breeding season. Anurans (17 species) were more abundant than salamanders (7 species), comprising 96.4% of individual captures. Most (95.9%) of the amphibian biomass came from 232095 individuals of a single species of anuran (southern leopard frog [Rana sphenocephala ]). Our results revealed the resilience of an amphibian community to natural stressors and historical habitat alteration and the potential magnitude of biomass and energy transfer from isolated wetlands to surrounding terrestrial habitat. We attributed the postdrought success of amphibians to a combination of adult longevity (often >5 years), a reduction in predator abundance, and an abundance of larval food resources. Likewise, the increase of forest cover around Ellenton Bay from <20% in 1951 to >60% in 2001 probably contributed to the long-term persistence of amphibians at this site. Our findings provide an optimistic counterpoint to the issue of the global decline of biological diversity by demonstrating that conservation efforts can mitigate historical habitat degradation.     

Lethal Effects of Water Quality on Threatened California Salamanders but Not on Co‐Occurring Hybrid Salamanders

Biological invasions and habitat alteration are often detrimental to native species, but their interactions are difficult to predict. Interbreeding between native and introduced species generates novel genotypes and phenotypes, and human land use alters habitat structure and chemistry. Both invasions and habitat alteration create new biological challenges and opportunities. In the intensively farmed Salinas Valley, California (U.S.A.), threatened California tiger salamanders (Ambystoma californiense) have been replaced by hybrids between California tiger salamander and introduced barred tiger salamanders (Ambystoma tigrinum mavortium). We conducted an enclosure experiment to examine the effects habitat modification and relative frequency of hybrid and native California tiger salamanders have on recruitment of salamanders and their prey, Pacific chorus frogs (Pseudacris regilla). We tested whether recruitment differed among genetic classes of tiger salamanders (hybrid or native) and pond hydroperiod (seasonal or perennial). Roughly 6 weeks into the experiment, 70% (of 378 total) of salamander larvae died in 4 out of 6 ponds. Native salamanders survived (n = 12) in these ponds only if they had metamorphosed prior to the die‐offs. During die‐offs, all larvae of native salamanders died, whereas 56% of hybrid larvae died. We necropsied native and hybrid salamanders, tested water quality, and queried the California Department of Pesticide Regulation database to investigate possible causes of the die‐offs. Salamander die‐offs, changes in the abundance of other community members (invertebrates, algae, and cyanobacteria), shifts in salamander sex ratio, and patterns of pesticide application in adjacent fields suggest that pesticide use may have contributed to die‐offs. That all survivors were hybrids suggests that environmental stress may promote rapid displacement of native genotypes. Efectos Letales de la Calidad del Agua sobre Salamandras de California Amenazadas pero no sobre Salamandras Híbridas Concurrentes     

Batrachochytrium dendrobatidis and the Collapse of Anuran Species Richness and Abundance in the Upper Manu National Park,Southeastern Peru

Abstract: Amphibians are declining worldwide, but these declines have been particularly dramatic in tropical mountains, where high endemism and vulnerability to an introduced fungal pathogen, Batrachochytrium dendrobatidis (Bd), is associated with amphibian extinctions. We surveyed frogs in the Peruvian Andes in montane forests along a steep elevational gradient (1200–3700 m). We used visual encounter surveys to sample stream‐dwelling and arboreal species and leaf‐litter plots to sample terrestrial‐breeding species. We compared species richness and abundance among the wet seasons of 1999, 2008, and 2009. Despite similar sampling effort among years, the number of species (46 in 1999) declined by 47% between 1999 and 2008 and by 38% between 1999 and 2009. When we combined the number of species we found in 2008 and 2009, the decline from 1999 was 36%. Declines of stream‐dwelling and arboreal species (a reduction in species richness of 55%) were much greater than declines of terrestrial‐breeding species (reduction of 20% in 2008 and 24% in 2009). Similarly, abundances of stream‐dwelling and arboreal frogs were lower in the combined 2008–2009 period than in 1999, whereas densities of frogs in leaf‐litter plots did not differ among survey years. These declines may be associated with the infection of frogs with Bd. B. dendrobatidis prevalence correlated significantly with the proportion of species that were absent from the 2008 and 2009 surveys along the elevational gradient. Our results suggest Bd may have arrived at the site between 1999 and 2007, which is consistent with the hypothesis that this pathogen is spreading in epidemic waves along the Andean cordilleras. Our results also indicate a rapid decline of frog species richness and abundance in our study area, a national park that contains many endemic amphibian species and is high in amphibian species richness.     

Disappearance and Return of Plethodontid Salamanders to Clearcut Plots in the Southern Blue Ridge Mountains

The disappearance and return of plethodontid salamanders on clearcuts has been monitored since 1979 in the southern Blue Ridge Mountains at three sites near Highlands, North Carolina. Salamander abundance on 225 m² plots located on clearcuts and in nearby forest was determined by nightly non-destructive searches. Abundances on clearcut and forest plots at a given site were compared for each year in which sampling occurred. Numbers of salamanders on clearcut plots decreased to 30–50% of forested plots in the first year after logging and were almost zero by the second year. Decreases in standing crop and moisture content of leaf litter seem responsible for salamander disappearance. Salamanders returned to clearcuts 4–6 years after cutting, and their numbers increased rapidly. Linear regressions estimate that salamander numbers on clearcut plots will equal or exceed numbers on forested plots by 20–24 years after cutting. The pattern of salamander return to clearcuts appears closely correlated with the timing of litter layer reformation. All sex and age classes of the most common species, Plethodon jordani, disappear from clearcuts at equal rates, whereas the earliest colonizers are predominantly large adults. Plethodon oconaluftee, a desiccation-resistant species, exists on regenerating clearcuts in disproportionately large numbers. Large adults of all species, including Plethodon oconaluftee, may be better able to withstand the drier, sparse litter cover of young, regenerating stands. Adults might move to clearcuts to avoid competition from smaller and immature salamanders restricted to mature forests with abundant, moist litter.     

Modeling the Effects of Anthropogenic Habitat Change on Savanna Snake Invasions into African Rainforest

Abstract:  We used a species-distribution modeling approach, ground-based climate data sets, and newly available remote-sensing data on vegetation from the MODIS and Quick Scatterometer sensors to investigate the combined effects of human-caused habitat alterations and climate on potential invasions of rainforest by 3 savanna snake species in Cameroon, Central Africa: the night adder (Causus maculatus) , olympic lined snake (Dromophis lineatus) , and African house snake (Lamprophis fuliginosus) . Models with contemporary climate variables and localities from native savanna habitats showed that the current climate in undisturbed rainforest was unsuitable for any of the snake species due to high precipitation. Limited availability of thermally suitable nest sites and mismatches between important life-history events and prey availability are a likely explanation for the predicted exclusion from undisturbed rainforest. Models with only MODIS-derived vegetation variables and savanna localities predicted invasion in disturbed areas within the rainforest zone, which suggests that human removal of forest cover creates suitable microhabitats that facilitate invasions into rainforest. Models with a combination of contemporary climate, MODIS- and Quick Scatterometer-derived vegetation variables, and forest and savanna localities predicted extensive invasion into rainforest caused by rainforest loss. In contrast, a projection of the present-day species-climate envelope on future climate suggested a reduction in invasion potential within the rainforest zone as a consequence of predicted increases in precipitation. These results emphasize that the combined responses of deforestation and climate change will likely be complex in tropical rainforest systems.     

Reservoir‐host amplification of disease impact in an endangered amphibian

Emerging wildlife pathogens are an increasing threat to biodiversity. One of the most serious wildlife diseases is chytridiomycosis, caused by the fungal pathogen, Batrachochytrium dendrobatidis (Bd), which has been documented in over 500 amphibian species. Amphibians vary greatly in their susceptibility to Bd; some species tolerate infection, whereas others experience rapid mortality. Reservoir hosts—species that carry infection while maintaining high abundance but are rarely killed by disease—can increase extinction risk in highly susceptible, sympatric species. However, whether reservoir hosts amplify Bd in declining amphibian species has not been examined. We investigated the role of reservoir hosts in the decline of the threatened northern corroboree frog (Pseudophryne pengilleyi) in an amphibian community in southeastern Australia. In the laboratory, we characterized the response of a potential reservoir host, the (nondeclining) common eastern froglet (Crinia signifera), to Bd infection. In the field, we conducted frog abundance surveys and Bd sampling for both P. pengilleyi and C. signifera. We built multinomial logistic regression models to test whether Crinia signifera and environmental factors were associated with P. pengilleyi decline. C. signifera was a reservoir host for Bd. In the laboratory, many individuals maintained intense infections (>1000 zoospore equivalents) over 12 weeks without mortality, and 79% of individuals sampled in the wild also carried infections. The presence of C. signifera at a site was strongly associated with increased Bd prevalence in sympatric P. pengilleyi. Consistent with disease amplification by a reservoir host, P. pengilleyi declined at sites with high C. signifera abundance. Our results suggest that when reservoir hosts are present, population declines of susceptible species may continue long after the initial emergence of Bd, highlighting an urgent need to assess extinction risk in remnant populations of other declined amphibian species.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

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.