Time and habitat structure shape insect acoustic activity in the Amazon.

Insects are the most diverse animal taxon on Earth and play a key role in ecosystem functioning. However, they are often neglected by ecological surveys owing to the difficulties involved in monitoring this small and hyper-diverse taxon. With technological advances in biomonitoring and analytical methods, these shortcomings may finally be addressed. Here, we performed passive acoustic monitoring at 141 sites (eight habitats) to investigate insect acoustic activity in the Viruá National Park, Brazil. We first describe the frequency range occupied by three soniferous insect groups (cicadas, crickets and katydids) to calculate the acoustic evenness index (AEI). Then, we assess how AEI varies spatially and temporally among habitat types, and finally we investigate the relationship between vegetation structure variables and AEI for each insect category. Overall, crickets occupied lower and narrower frequency bands than cicadas and katydids. AEI values varied among insect categories and across space and time. The highest acoustic activity occurred before sunrise and the lowest acoustic activity was recorded in pastures. Canopy cover was positively associated with cricket acoustic activity but not with katydids. Our findings contribute to a better understanding of the role of time, habitat and vegetation structure in shaping insect activity within diverse Amazonian ecosystems. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Invading nonnative frogs use different microhabitats and change physiology along an elevation gradient.

The coqui frog (Eleutherodactylus coqui) was introduced to the island of Hawai'i in the 1980s, and has spread across much of the island. There is concern they will invade higher elevation areas where negative impacts on native species are expected. It is not known if coqui change behavior and baseline physiology in ways that allow them to invade higher elevations. We investigated where coqui are found across the island and whether that includes recent invasion into higher elevations. We also investigated whether elevation is related to coqui's microhabitat use, including substrate use and height off the forest floor, and physiological metrics, including plasma osmolality, oxidative status, glucose, free glycerol, and triglycerides, that might be associated with invading higher elevations. We found coqui have increased the area they occupy along roads from 31% to 50% and have moved into more high-elevation locations (16% vs. 1%) compared to where they were found 14 years ago. We also found frogs at high elevation on different substrates and closer to the forest floor than frogs at lower elevations-perhaps in response to air temperatures which tended to be warmer close to the forest floor. We observed that blood glucose and triglycerides increase in frogs with elevation. An increase in glucose is likely an acclimation response to cold temperatures while triglycerides may also help frogs cope with the energetic demands of suboptimal temperatures. Finally, we found that female coqui have higher plasma osmolality, reactive oxygen metabolites (dROMs), free glycerol, and triglycerides than males. Our study suggests coqui behavior and physiology in Hawai'i may be influenced by elevation in ways that allow them to cope with lower temperatures and invade higher elevations.

Invasive frogs show persistent physiological differences to elevation and acclimate to colder temperatures.

The coqui frog (Eleutherodactylus coqui) was introduced to the island of Hawai'i in the 1980s and has spread across much of the island. Concern remains that this frog will continue to expand its range and invade higher elevation habitats where much of the island's endemic species are found. We determined whether coqui thermal tolerance and physiology change along Hawai'i's elevational gradients. We measured physiological responses using a short-term experiment to determine baseline tolerance and physiology by elevation, and a long-term experiment to determine the coqui's ability to acclimate to different temperatures. We collected frogs from low, medium, and high elevations. After both the short and long-term experiments, we measured critical thermal minimum (CTmin), blood glucose, oxidative stress, and corticosterone levels. CTmin was lower in high elevation frogs than low elevation frogs after the short acclimation experiment, signifying that they acclimate to local conditions. After the extended acclimation, CTmin was lower in frogs acclimated to cold temperatures compared to warm-acclimated frogs and no longer varied by elevation. Blood glucose levels were positively correlated with elevation even after the extended acclimation, suggesting glucose may also be related to lower temperatures. Oxidative stress was higher in females than males, and corticosterone was not significantly related to any predictor variables. The extended acclimation experiment showed that coquis can adjust their thermal tolerance to different temperatures over a 3-week period, suggesting the expansion of coqui into higher elevation habitats may still be possible, and they may not be as restricted by cold temperatures as previously thought.

A modern two-layer hypothesis helps resolve the 'savanna problem'.

For over a century, deep roots have been assumed to allow trees to avoid competition with grasses (i.e., the two-layer hypothesis). Yet, in part because it remains difficult to measure water uptake in the field, there has been a shift in savanna ecology away from the two-layer hypothesis and towards alternative explanations of tree-grass coexistence. Here, we combine hydrologic tracer experiments and soil water flow models to demonstrate how the distribution of active roots affects water uptake across a range of savanna conditions. Grass roots were shallower and provided pre-emptive access to enough soil water to allow nearly continuous grass cover, but slightly deeper roots provided trees with more total water under most conditions. This 'some water now or more water later' tradeoff varied with precipitation amount, soil texture, and tree and grass relative root abundance in ways that helped explain tree and grass landscape abundance.

Herbivory changes soil microbial communities and greenhouse gas fluxes in a high-latitude wetland.

Herbivory can have strong impacts on greenhouse gas fluxes in high-latitude ecosystems. For example, in the Yukon-Kuskokwim (Y-K) Delta in western Alaska, migratory goose grazing affects the magnitude of soil carbon dioxide (CO2) and methane (CH4) fluxes. However, the underlying drivers of this relationship are unclear, as few studies systematically tease apart the processes by which herbivores influences soil biogeochemistry. To examine these mechanisms in detail, we conducted a laboratory incubation experiment to quantify changes in greenhouse gas fluxes in response to three parameters altered by herbivores in situ: temperature, soil moisture content, and nutrient inputs. These treatments were applied to soils collected in grazing lawns and nearby ungrazed habitat, allowing us to assess how variation in microbial community structure influenced observed responses. We found pronounced differences in both fungal and prokaryotic community composition between grazed and ungrazed areas. In the laboratory incubation experiment, CO2 and CH4 fluxes increased with temperature, soil moisture, and goose fecal addition, suggesting that grazing-related changes in the soil abiotic environment may enhance soil C losses. Yet, these abiotic drivers were insufficient to explain variation in fluxes between soils with and without prior grazing. Differences in trace gas fluxes between grazed and ungrazed areas may result both from herbivore-induced shifts in abiotic parameters and grazing-related alterations in microbial community structure. Our findings suggest that relationships among herbivores and soil microbial communities could mediate carbon-climate feedbacks in rapidly changing high-latitude ecosystems.

Sex-related differences in aging rate are associated with sex chromosome system in amphibians.

Sex-related differences in mortality are widespread in the animal kingdom. Although studies have shown that sex determination systems might drive lifespan evolution, sex chromosome influence on aging rates have not been investigated so far, likely due to an apparent lack of demographic data from clades including both XY (with heterogametic males) and ZW (heterogametic females) systems. Taking advantage of a unique collection of capture-recapture datasets in amphibians, a vertebrate group where XY and ZW systems have repeatedly evolved over the past 200 million years, we examined whether sex heterogamy can predict sex differences in aging rates and lifespans. We showed that the strength and direction of sex differences in aging rates (and not lifespan) differ between XY and ZW systems. Sex-specific variation in aging rates was moderate within each system, but aging rates tended to be consistently higher in the heterogametic sex. This led to small but detectable effects of sex chromosome system on sex differences in aging rates in our models. Although preliminary, our results suggest that exposed recessive deleterious mutations on the X/Z chromosome (the "unguarded X/Z effect") or repeat-rich Y/W chromosome (the "toxic Y/W effect") could accelerate aging in the heterogametic sex in some vertebrate clades.

Woody plant growth increases with precipitation intensity in a cold semiarid system.

As the atmosphere warms, precipitation events become larger, but less frequent. Yet, there is fundamental disagreement about how increased precipitation intensity will affect vegetation. Walter's two-layer hypothesis and experiments testing it have demonstrated that precipitation intensity can increase woody plant growth. Observational studies have found the opposite pattern. Not only are the patterns contradictory, but inference is largely limited to grasslands and savannas. We tested the effects of increased precipitation intensity in a shrub-steppe ecosystem that receives >30% of its precipitation as snow. We used 11 (8 × 8 m) shelters to collect and redeposit rain and snow as larger, more intense events. Total annual precipitation was the same in all plots, but each plot received different precipitation event sizes ranging from 1 to 18 mm. Over three growing seasons, larger precipitation event sizes increased soil water availability, sagebrush (Artemisia tridentata) stem radius, and canopy greenness, decreased new root growth in shallow soils, and had no effect on herbaceous plant cover. Thus, we found that increased precipitation intensity can increase soil water availability and woody plant growth in a cold semiarid system. Assuming that stem growth is positively correlated with shrub reproduction, establishment and spread, results suggest that increasing precipitation intensity may have contributed to the woody plant encroachment observed around the world in the past 50 yr. Further, continuing increases in precipitation intensity caused by atmospheric warming are likely to continue to contribute to shrub encroachment in the future.

Small differences in root distributions allow resource niche partitioning.

Deep roots have long been thought to allow trees to coexist with shallow-rooted grasses. However, data demonstrating how root distributions affect water uptake and niche partitioning are uncommon.We describe tree and grass root distributions using a depth-specific tracer experiment six times over two years in a subtropical savanna, Kruger National Park, South Africa. These point-in-time measurements were then used in a soil water flow model to simulate continuous water uptake by depth and plant growth form (trees and grasses) across two growing seasons. This allowed estimates of the total amount of water a root distribution could absorb as well as the amount of water a root distribution could absorb in excess of the other rooting distribution (i.e., unique hydrological niche).Most active tree and grass roots were in shallow soils: The mean depth of water uptake was 22 cm for trees and 17 cm for grasses. Slightly deeper rooting distributions provided trees with 5% more soil water than the grasses in a drier season, but 13% less water in a wetter season. Small differences also provided each rooting distribution (tree or grass) with unique hydrological niches of 4 to 13 mm water.The effect of rooting distributions has long been inferred. By quantifying the depth and timing of water uptake, we demonstrated how even small differences in rooting distributions can provide plants with resource niches that can contribute to species coexistence. Differences in total water uptake and unique hydrological niche sizes were small in this system, but they indicated that tradeoffs in rooting strategies can be expected to contribute to tree and grass coexistence because 1) competitive advantages change over time and 2) plant growth forms always have access to a soil resource pool that is not available to the other plant growth form.

Herbivores at the highest risk of extinction among mammals, birds, and reptiles.

As a result of their extensive home ranges and slow population growth rates, predators have often been perceived to suffer higher risks of extinction than other trophic groups. Our study challenges this extinction-risk paradigm by quantitatively comparing patterns of extinction risk across different trophic groups of mammals, birds, and reptiles. We found that trophic level and body size were significant factors that influenced extinction risk in all taxa. At multiple spatial and temporal scales, herbivores, especially herbivorous reptiles and large-bodied herbivores, consistently have the highest proportions of threatened species. This observed elevated extinction risk for herbivores is ecologically consequential, given the important roles that herbivores are known to play in controlling ecosystem function.

Functional traits explain amphibian distribution in the Brazilian Atlantic Forest.

AIM: Species distributions are one of the most important ways to understand how communities interact through macroecological relationships. The functional abilities of a species, such as its plasticity in various environments, can determine its distribution, species richness and beta diversity patterns. In this study, we evaluate how functional traits influence the distribution of amphibians, and hypothesize which functional traits explain the current pattern of amphibian species composition. LOCATION: Atlantic Forest, Brazil. TAXON: Amphibia (Anura and Gymnophiona). METHODS: Using potential distributions of Brazilian amphibians from Atlantic Forest based on their functional traits, we analysed the influence of biotic and abiotic factors on species richness, endemism (with permutation multivariate analysis) and beta diversity components (i.e. total, turnover and nestedness dissimilarities). RESULTS: Environmental variables explained 59.5% of species richness, whereas functional traits explained 15.8% of species distribution (geographical species range) for Anuran and 88.8% for Gymnophiona. Body size had the strongest correlation with species distribution. Results showed that species with medium to large body size, and species that are adapted to living in open areas tended to disperse from west to east direction. Current forest changes directly affected beta diversity patterns (i.e. most species adapted to novel environments increase their ranges). Beta diversity partitioning between humid and dry forests showed decreased nestedness and increased turnover by increasing altitude in the south-eastern region of the Atlantic Forest. MAIN CONCLUSIONS: Our study shows that functional traits directly influence the ability of the species to disperse. With the alterations of the natural environment, species more apt to these alterations have dispersed or increased their distribution, which consequently changes community structure. As a result, there are nested species distribution patterns and homogenization of amphibian species composition throughout the Brazilian Atlantic Forest.

The Missing Angle: Ecosystem Consequences of Phenological Mismatch.

Climate change leads to unequal shifts in the phenology of interacting species, such as consumers and their resources, leading to potential phenological mismatches. While studies have investigated how phenological mismatch affects wild populations, we still lack studies and a framework for investigating how phenological mismatch affects ecosystems, particularly nutrient cycling.

Amphibians of Santa Teresa, Brazil: the hotspot further evaluated.

A checklist of the amphibians of Santa Teresa municipality, in southeastern Brazil is presented based on fieldwork, examination of specimens in collections, and a literature review. This new amphibian list of Santa Teresa includes 108 species, of which 106 (~98%) belong to Anura and two (~2%) to Gymnophiona. Hylidae was the most represented family with 47 species (43%). Compared to the previous amphibian lists for Santa Teresa, 14 species were added, 17 previously reported species were removed, and 13 species were re-identified based on recent taxonomic rearrangements. Of the 14 species added, 11 (79%) were first recorded during our fieldwork and specimen examination. It is also the first list of caecilians for Santa Teresa. This list suggests that Santa Teresa has 0.16 species per km2 (i.e., 108 species/683 km2), one of the highest densities of amphibian species in the world at a regional scale. This richness represents 78% of the 136 anurans from Espírito Santo state and 10% of the 1,080 amphibians from Brazil. We highlight the need for long-term monitoring to understand population trends and develop effective conservation plans to safeguard this remarkable amphibian richness.

Migratory goose arrival time plays a larger role in influencing forage quality than advancing springs in an Arctic coastal wetland.

With warmer springs, herbivores migrating to Arctic breeding grounds may experience phenological mismatches between their energy demands and the availability of high quality forage. Yet, how the timing of the start of the season and herbivore arrival influences forage quality is often unknown. In coastal western Alaska, approximately one million migratory geese arrive each spring to breed, where foliar %N and C:N ratios are linked to gosling survival and population growth. We conducted a three-year experiment where we manipulated the start of the growing season using warming chambers and grazing times using captive Pacific black brant (Branta bernicla nigricans) to examine how the timing of these events influences the quality of an important forage species. Our results suggest that grazing timing plays a much greater role than an advanced growing season in determining forage quality. All top models included grazing timing, and suggested that compared to typical grazing timing, early grazing significantly reduced foliar %C by 6% and C:N ratios by 16%, while late goose grazing significantly reduced foliar %N by 15% and increased foliar C:N ratios by 21%. While second-ranking top models included the effect of season, the advanced growing season effect was not significant and only reduced %N by 4%, increased %C by <1%, and increased C:N ratios by 5% compared to an ambient growing season. In summary, in years where geese arrive early, they will consume higher quality forage when they arrive and throughout the season, while in years that geese arrive late they will consume lower quality forage when they arrive and for the remainder of the season. When the growing season starts has only a minor influence on this pattern. Our findings suggest that cues determining migration and arrival times to breeding areas are important factors influencing forage quality for geese in western Alaska.

Chronosequence and direct observation approaches reveal complementary community dynamics in a novel ecosystem.

Non-native, early-successional plants have been observed to maintain dominance for decades, particularly in semi-arid systems. Here, two approaches were used to detect potentially slow successional patterns in an invaded semi-arid system: chronosequence and direct observation. Plant communities in 25 shrub-steppe sites that represented a 50-year chronosequence of agricultural abandonment were monitored for 13 years. Each site contained a field abandoned from agriculture (ex-arable) and an adjacent never-tilled field. Ex-arable fields were dominated by short-lived, non-native plants. These 'weedy' communities had lower species richness, diversity and ground cover, and greater annual and forb cover than communities in never-tilled fields. Never-tilled fields were dominated by long-lived native plants. Across the chronosequence, plant community composition remained unchanged in both ex-arable and never-tilled fields. In contrast, 13 years of direct observation detected directional changes in plant community composition within each field type. Despite within-community changes in both field types during direct observation, there was little evidence that native plants were invading ex-arable fields or that non-native plants were invading never-tilled fields. The more-controlled, direct observation approach was more sensitive to changes in community composition, but the chronosequence approach suggested that these changes are unlikely to manifest over longer time periods, at least in part because of disturbances in the system. Results highlight the long-term consequences of soil disturbance and the difficulty of restoring native perennials in disturbed semi-arid systems.

Delayed herbivory by migratory geese increases summer-long CO2 uptake in coastal western Alaska.

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to "phenological mismatches" as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three-year experiment to examine the consequences for CO2 exchange of advanced spring green-up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green-up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO2 uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green-up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m-2  s-1 toward a greater CO2 sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO2 uptake. Importantly, while the direct effect of climate warming on phenology of green-up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.

Predictors of Participation in Invasive Species Control Activities Depend on Prior Experience with the Species.

The increasing worldwide spread of non-native species is both a component and a consequence of environmental change, and islands are especially vulnerable to negative effects. Efforts to control non-native species often include public education intended to promote behaviors designed to reduce or reverse their spread. To inform the use of information strategies to control the invasive, non-native frog Eleutherodactylus coqui in Hawaii, USA, we surveyed over 700 property owners about their attitudes and behaviors regarding the species. Included were residents of the island of Hawaii, where the species is common and management emphasizes prevention of further spread, and three other islands where the species is largely absent and management emphasizes detection and eradication. Where frogs are present, 61% of respondents reported taking actions to reduce their population, typically clearing vegetation or hand-capturing individual frogs. For these individuals, intentions to engage in future control activities were not significantly related to reports of past behavior. Intentions to participate in future control efforts on the island of Hawaii were best predicted by attitudes toward practices. On the other islands, behavioral intentions were best predicted by subjective norms (i.e., beliefs about others' expectations that they should manage frogs). Thus, intentions to engage in non-native species management behaviors appear to be influenced by prior exposure to, and experience with, that species. Understanding the predictors of behavioral intentions at different stages of invasion have implications for the design of information strategies that can promote participation in control activities.

Change in dominance determines herbivore effects on plant biodiversity.

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition.

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra- and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five-fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra- and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.

Direct effects of warming increase woody plant abundance in a subarctic wetland.

Both the direct effects of warming on a species' vital rates and indirect effects of warming caused by interactions with neighboring species can influence plant populations. Furthermore, herbivory mediates the effects of warming on plant community composition in many systems. Thus, determining the importance of direct and indirect effects of warming, while considering the role of herbivory, can help predict long-term plant community dynamics. We conducted a field experiment in the coastal wetlands of western Alaska to investigate how warming and herbivory influence the interactions and abundances of two common plant species, a sedge, Carex ramenskii, and a dwarf shrub, Salix ovalifolia. We used results from the experiment to model the equilibrium abundances of the species under different warming and grazing scenarios and to determine the contribution of direct and indirect effects to predict population changes. Consistent with the current composition of the landscape, model predictions suggest that Carex is more abundant than Salix under ambient temperatures with grazing (53% and 27% cover, respectively). However, with warming and grazing, Salix becomes more abundant than Carex (57% and 41% cover, respectively), reflecting both a negative response of Carex and a positive response of Salix to warming. While grazing reduced the cover of both species, herbivory did not prevent a shift in dominance from sedges to the dwarf shrub. Direct effects of climate change explained about 97% of the total predicted change in species cover, whereas indirect effects explained only 3% of the predicted change. Thus, indirect effects, mediated by interactions between Carex and Salix, were negligible, likely due to use of different niches and weak interspecific interactions. Results suggest that a 2°C increase could cause a shift in dominance from sedges to woody plants on the coast of western Alaska over decadal timescales, and this shift was largely a result of the direct effects of warming. Models predict this shift with or without goose herbivory. Our results are consistent with other studies showing an increase in woody plant abundance in the Arctic and suggest that shifts in plant-plant interactions are not driving this change.