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.

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.

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.

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.

Live long and prosper: plant-soil feedback, lifespan, and landscape abundance covary.

Plant soil feedbacks (PSFs) are thought to be important to plant growth and species coexistence, but most support for these hypotheses is derived from short-term greenhouse experiments. Here we use a seven-year, common garden experiment to measure PSFs for seven native and six nonnative species common to the western United States. We use these long-term, field-based estimates to test correlations between PSF and plant landscape abundance, species origin, functional type, and lifespan. To assess potential PSF mechanisms, we also measured soil microbial community composition, root biomass, nitrogen cycling, bulk density, penetration resistance, and shear strength. Plant abundance on the landscape and plant lifespan were positively correlated with PSFs, though this effect was due to the relationships for native plants. PSFs were correlated with indices of soil microbial community composition. Soil nutrient and physical traits and root biomass differed among species but were not correlated with PSF. While results must be taken with caution because only 13 species were examined, these species represent most of the dominant plant species in the system. Results suggest that native plant abundance is associated with the ability of long-lived plants to create positive plant-soil microbe interactions, while short-lived nonnative plants maintain dominance by avoiding soil-borne antagonists, increasing nitrogen cycling and dedicating resources to aboveground growth and reproduction rather than to belowground growth. Broadly, results suggest that PSFs are correlated with a suite of traits that determine plant abundance.

Using plant-soil feedbacks to predict plant biomass in diverse communities.

It has become clear that plants can create soils that affect subsequent plant growth. However, because plant-soil feedbacks (PSFs) are typically measured in monoculture experiments, it remains unclear to what extent PSFs affect plant growth in communities. Here we used data from a factorial PSF experiment to predict the biomass of 12 species grown in 162 plant community combinations. Five different plant growth models were parameterized with either monoculture biomass data (Null) or with PSF data (PSF) and model predictions were compared to plant growth observed in communities. For each of the five models, PSF model predictions were closer to observed species biomass in communities than Null model predictions. PSFs, which were associated with a 28% difference in plant biomass across soil types, explained 10% more variance than Null models. Results provided strong support for a small role for PSFs in predicting plant growth in communities and suggest several reasons that PSFs, as traditionally measured in monoculture experiments, may overestimate PSF effects in communities. First, monoculture data used in Null models inherently includes "self " PSF effects. Second, PSFs must be large relative to differences in intrinsic growth rates among species to change competitive outcomes. Third, PSFs must vary among species to change species relative abundances.

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'.

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.

Root niche partitioning among grasses, saplings, and trees measured using a tracer technique.

Niche partitioning of resources by plants is believed to be a fundamental aspect of plant coexistence and biogeochemical cycles; however, measurements of the timing and location of resource use are often lacking because of the difficulties of belowground research. To measure niche partitioning of soil water by grasses, planted saplings, and trees in a mesic savanna (Kruger National Park, South Africa), we injected deuterium oxide into 102,000 points in 15, 154-m(2) plots randomly assigned to one of five depths (0-120 cm) and one of three time periods during the 2008/2009 growing season. Grasses, saplings and trees all demonstrated an exponential decline in water uptake early in the season when resources were abundant. Later in the season, when resources were scarce, grasses continued to extract the most water from the shallowest soil depths (5 cm), but saplings and trees shifted water uptake to deeper depths (30-60 cm). Saplings, in particular, rapidly established roots to at least 1 m and used these deep roots to a greater extent than grasses or trees. Helping to resolve contradictory observations of the relative importance of deep and shallow roots, our results showed that grasses, saplings and trees all extract the most water from shallow soils when it is available but that woody plants can rapidly shift water uptake to deeper soils when resources are scarce. Results highlight the importance of temporal changes in water uptake and the problems with inferring spatial and temporal partitioning of soil water uptake from root biomass measurements alone.

Plant-soil feedbacks provide an additional explanation for diversity-productivity relationships.

Plant-soil feedbacks (PSFs) have gained attention for their role in plant community dynamics, but their role in productivity has been overlooked. We developed and tested a biomass-specific, multi-species model to examine the role of PSFs in diversity-productivity relationships. The model predicts a negative relationship between PSFs and overyielding: plants with negative PSFs grow more in communities than in monoculture (i.e. overyield), and plants with positive PSFs grow less in communities than in monoculture (i.e. underyield). This effect is predicted to increase with diversity and saturate at low species richness because the proportion of 'self-cultivated' soils rapidly decreases as species are added to a community. Results in a set of glasshouse experiments supported model predictions. We found that PSFs measured in one experiment were negatively correlated with overyielding in three-species plant communities measured in a separate experiment. Furthermore, when parametrized with our experimental PSF data, our model successfully predicted species-level overyielding and underyielding. The model was less effective at predicting community-level overyielding and underyielding, although this appeared to reflect large differences between communities with or without nitrogen-fixing plants. Results provide conceptual and experimental support for the role of PSFs in diversity-productivity relationships.

Cast adrift on an island: introduced populations experience an altered balance between selection and drift.

A long-standing question in evolutionary biology is what becomes of adaptive traits when a species expands its range into novel environments. Here, we report the results of a study on an adaptive colour pattern polymorphism (stripes) of the coqui frog, Eleutherodactylus coqui, following its introduction to Hawaii from Puerto Rico. We compared population differentiation (Φ ST and F ST ) for the stripes locus--which underlies this colour pattern polymorphism--with neutral microsatellite loci to test for a signature of selection among native and introduced populations. Among native populations, Φ ST and F ST for stripes were lower than expected under the neutral model, suggesting uniform balancing selection. Alternatively, among introduced populations, Φ ST and F ST for stripes did not differ from the neutral model. These results suggest that the evolutionary dynamics of this previously adaptive trait have become dominated by random genetic drift following the range expansion.

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.

A depth-controlled tracer technique measures vertical, horizontal and temporal patterns of water use by trees and grasses in a subtropical savanna.

• As described in the two-layer hypothesis, woody plants are often assumed to use deep soils to avoid competition with grasses. Yet the direct measurements of root activity needed to test this hypothesis are rare. • Here, we injected deuterated water into four soil depths, at four times of year, to measure the vertical and horizontal location of water uptake by trees and grasses in a mesic savanna in Kruger National Park, South Africa. • Trees absorbed 24, 59, 14 and 4% of tracer from the 5, 20, 50, and 120 cm depths, respectively, while grasses absorbed 61, 29, 9 and 0.3% of tracer from the same depths. Only 44% of root mass was in the top 20 cm. Trees absorbed tracer under and beyond their crowns, while 98% of tracer absorbed by grasses came from directly under the stem. • Trees and grasses partitioned soil resources (20 vs 5 cm), but this partitioning did not reflect, as suggested by the two-layer hypothesis, the ability of trees to access deep soil water that was unavailable to grasses. Because root mass was a poor indicator of root activity, our results highlight the importance of precise root activity measurements.

A meta-analytic review of corridor effectiveness.

Using corridors for conservation is increasing despite a lack of consensus on their efficacy. Specifically, whether corridors increase movement of plants and animals between habitat fragments has been addressed on a case-by-case basis with mixed results. Because of the growing number of well-designed experiments that have addressed this question, we conducted a meta-analysis to determine whether corridors increase movement; whether corridor effectiveness differs among taxa; how recent changes in experimental design have influenced findings; and whether corridor effectiveness differs between manipulative and natural experiments. To conduct our meta-analysis, we analyzed 78 experiments from 35 studies using a conservative hierarchical Bayesian model that accounts for hierarchical and sampling dependence. We found a highly significant result that corridors increase movement between habitat patches by approximately 50% compared to patches that are not connected with corridors. We found that corridors were more important for the movement of invertebrates, nonavian vertebrates, and plants than they were for birds. Recent methodological advances in corridor experiments, such as controlling for the area added by corridors, did not influence whether corridors increased movement, whereas controlling for the distance between source and connected or unconnected recipient patches decreased movement through corridors. After controlling for taxa differences and whether studies controlled for distance in experimental design, we found that natural corridors (those existing in landscapes prior to the study) showed more movement than manipulated corridors (those created and maintained for the study). Our results suggest that existing corridors increase species movement in fragmented landscapes and that efforts spent on maintaining and creating corridors are worthwhile.

Genetic basis of a color pattern polymorphism in the Coqui Frog Eleutherodactylus coqui.

Many species of frog exhibit striking color and pattern polymorphisms, but the genetic bases of these traits are not known for most species. The coqui frog, Eleutherodactylus coqui, a species endemic to the island of Puerto Rico, exhibits a wide variety of color and pattern polymorphisms including 4 discrete stripe patterns on its dorsal surface and an unstriped morph. We conducted breeding experiments to determine the mode of inheritance for these 5 dorsal color patterns in E. coqui. We analyzed results from 14 different cross types, which included 1519 offspring from 71 clutches. We found that color patterns segregate at ratios consistent with a single autosomal locus, 5-allele model, in which all alleles coding for stripes are codominant and the allele coding for the unstriped morph is recessive. We propose that this locus be named "stripes" with alleles B (interocular bar), L (dorsolateral stripes), N (narrow middorsal stripe), W (wide middorsal stripe), and u (unstriped). The results of this experiment suggest the genetic basis of stripe patterns in this well-studied species and provide a model for studying the evolution and maintenance of this phenotypic polymorphism.

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.