How ecologists define drought,and why we should do better

Drought, widely studied as an important driver of ecosystem dynamics, is predicted to increase in frequency and severity globally. To study drought, ecologists must define or at least operationalize what constitutes a drought. How this is accomplished in practice is unclear, particularly given that climatologists have long struggled to agree on definitions of drought, beyond general variants of “an abnormal deficiency of water.” We conducted a literature review of ecological drought studies (564 papers) to assess how ecologists describe and study drought. We found that ecologists characterize drought in a wide variety of ways (reduced precipitation, low soil moisture, reduced streamflow, etc.), but relatively few publications (~32%) explicitly define what are, and are not, drought conditions. More troubling, a surprising number of papers (~30%) simply equated “dry conditions” with “drought” and provided little characterization of the drought conditions studied. For a subset of these, we calculated Standardized Precipitation Evapotranspiration Index values for the reported drought periods. We found that while almost 90% of the studies were conducted under conditions quantifiable as slightly to extremely drier than average, ~50% were within the range of normal climatic variability. We conclude that the current state of the ecological drought literature hinders synthesis and our ability to draw broad ecological inferences because drought is often declared but is not explicitly defined or well characterized. We suggest that future drought publications provide at least one of the following: (a) the climatic context of the drought period based on long‐term records; (b) standardized climatic index values; (c) published metrics from drought‐monitoring organizations; (d) a quantitative definition of what the authors consider to be drought conditions for their system. With more detailed and consistent quantification of drought conditions, comparisons among studies can be more rigorous, increasing our understanding of the ecological effects of drought.     

Long‐term Changes in Fruit Phenology in a West African Lowland Tropical Rain Forest are Not Explained by Rainfall

Decreases in rainfall have been proposed to have a negative impact on tropical rain forests, and West Africa is currently experiencing a decline in rainfall at the multi‐decadal scale. Here, we present analyses of a long‐term dataset on the plant fruiting status from individuals of 44 species of the tropical rain forest of Taï National Park, Côte d'Ivoire. This study includes records of 1401 individuals collected at monthly intervals for over 12 yr, 984 of which survived throughout the entire study period. The aims of this study were to: (1) quantitate inter‐annual trends in species and forest scale fruit presence; and (2) test the importance of rainfall in explaining inter‐annual fruit presence variability. Long‐term upward trends in the expected proportion of individuals with fruits were found for the majority of species, while no significant downward trends were detected, driving a significant upward trend at the community level. Peak production months of the upward trending species were not associated with the dry or wet season. Significant rainfall correlations with the total proportion of individuals showing fruit were generally negative, with only five species showing significant positive correlations. Taken together, these results suggest that the observed inter‐annual trends and variability of fruit abundance are currently not associated with rainfall. We discuss several parsimonious and complex alternative explanations.     

A Multilevel Analysis of the Impact of Land Use on Interannual Land-Cover Change in East Africa

Abstract The aim of this study was to characterize the short-term land-cover change processes that were detected in Eastern Africa, based on a set of change metrics that allow for the quantification of interannual changes in vegetation productivity, changes in vegetation phenology and a combination of both. We tested to what extent land use, fire activity and livestock grazing modified the vegetation response to short-term rainfall variability in East Africa and how this is reflected in change metrics derived from MODerate Imaging Spectrometer (MODIS) time series of remote sensing data. We used a hierarchical approach to disentangle the contribution of human activities and climate variability to the patterns of short-term vegetation change in East Africa at different levels of organization. Our results clearly show that land use significantly influences the vegetation response to rainfall variability as measured by time series of MODIS data. Areas with different types of land use react in a different way to interannual climate variability, leading to different values of the change indices depending on the land use type. The impact of land use is more reflected in interannual variability of vegetation productivity and overall change in the vegetation, whereas changes in phenology are mainly driven by climate variability and affect most vegetation types in similar ways. Our multilevel approach led to improved models and clearly demonstrated that climate influence plays at a different scale than land use, fire and herbivore grazing. It helped us to understand dynamics within and between biomes in the study area and investigate the relative importance of different factors influencing short-term variability in change indices at different scales.     

Contributions of MIR to soundscape ecology. Part I: Potential methodological synergies

We present here the potential synergies between a sub-discipline of music called computational musicology and its associated toolkit Music Information Retrieval (MIR) systems and the current needs of soundscape ecologists who use passive acoustic recorders to collect massive amounts of audio data. These cross-sub-disciplinary synergies fall into three broad methodological categories: (1) automated feature extraction, (2) sound classification and labeling using machine learning, and (3) data visualization. We argue that the use of the MIR toolkit by soundscape ecologists could represent the development of a second generation of soundscape metrics, which are more translatable to policy and natural resource management problems. Two companion articles in this special issue by the authors provide a more detailed examination of the potential for computational musicology's MIR toolkit as applied to questions common to the emerging field of soundscape ecology.     

Robustness to thermal variability differs along a latitudinal gradient in zooplankton communities

Determining how thermal variability will affect the structure, stability, and function of ecological communities is becoming increasingly important as global warming is predicted to affect not only average temperatures but also increase the frequency of long runs of high temperatures. Latitudinal differences in the responses of ecological communities to changes in their thermal regimes have also been predicted based on adaptations over evolutionary time to different thermal environments. We conducted an experiment to determine whether variability in temperature leads to consistent changes in community structure, temporal dynamics, and ecosystem functioning in laboratory analogues of natural freshwater supralittoral rock pool communities inhabited by meiofauna and zooplankton collected from sub‐Arctic, temperate, and tropical regions. Thermal variability of +4 °C around mean temperature led to increased extinction frequency, decreases in consumer abundance, increases in temporal variability of consumer abundance, and shifts from predominately negative interactions observed under constant temperature to positive interactions in the temperate and tropical communities but not in the sub‐Arctic communities. That sub‐Arctic zooplankton communities may be more robust to thermal variability than temperate or tropical communities’ supports recent studies on macrophysiological adaptations of species along latitudinal gradients and suggests that increasing thermal variability may have the greatest effects on community structure and function in tropical and temperate regions.     

Influence of climatic variables, forest type, and condition on activity patterns of Geoffroyi's spider monkeys throughout Mesoamerica

Understanding how species cope with variations in climatic conditions, forest types and habitat amount is a fundamental challenge for ecologists and conservation biologists. We used data from 18 communities of Mesoamerican spider monkeys (Ateles geoffroyi) throughout their range to determine whether their activity patterns are affected by climatic variables (temperature and rainfall), forest types (seasonal and nonseasonal forests), and forest condition (continuous and fragmented). Data were derived from 15 published and unpublished studies carried out in four countries (Mexico, El Salvador, Costa Rica, and Panama), cumulatively representing more than 18 years (221 months, >3,645 hr) of behavioral observations. Overall, A. geoffroyi spent most of their time feeding (38.4 ± 14.0%, mean ± SD) and resting (36.6 ± 12.8%) and less time traveling (19.8 ± 11.3%). Resting and feeding were mainly affected by rainfall: resting time increased with decreasing rainfall, whereas feeding time increased with rainfall. Traveling time was negatively related to both rainfall and maximum temperature. In addition, both resting and traveling time were higher in seasonal forests (tropical dry forest and tropical moist forest) than in nonseasonal forests (tropical wet forest), but feeding time followed the opposite pattern. Furthermore, spider monkeys spent more time feeding and less time resting (i.e., higher feeding effort) in forest fragments than in continuous forest. These findings suggest that global climate changes and habitat deforestation and fragmentation in Mesoamerica will threaten the survival of spider monkeys and reduce the distributional range of the species in the coming decades. Am. J. Primatol. 73:1189–1198, 2011. © 2011 Wiley Periodicals, Inc.     

Drier tropical forests are susceptible to functional changes in response to a long‐term drought

Climatic changes have profound effects on the distribution of biodiversity, but untangling the links between climatic change and ecosystem functioning is challenging, particularly in high diversity systems such as tropical forests. Tropical forests may also show different responses to a changing climate, with baseline climatic conditions potentially inducing differences in the strength and timing of responses to droughts. Trait‐based approaches provide an opportunity to link functional composition, ecosystem function and environmental changes. We demonstrate the power of such approaches by presenting a novel analysis of long‐term responses of different tropical forest to climatic changes along a rainfall gradient. We explore how key ecosystem's biogeochemical properties have shifted over time as a consequence of multi‐decadal drying. Notably, we find that drier tropical forests have increased their deciduous species abundance and generally changed more functionally than forests growing in wetter conditions, suggesting an enhanced ability to adapt ecologically to a drying environment.     

Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.     

A conceptual guide to measuring species diversity

Three metrics of species diversity – species richness, the Shannon index and the Simpson index – are still widely used in ecology, despite decades of valid critiques leveled against them. Developing a robust diversity metric has been challenging because, unlike many variables ecologists measure, the diversity of a community often cannot be estimated in an unbiased way based on a random sample from that community. Over the past decade, ecologists have begun to incorporate two important tools for estimating diversity: coverage and Hill diversity. Coverage is a method for equalizing samples that is, on theoretical grounds, preferable to other commonly used methods such as equal-effort sampling, or rarefying datasets to equal sample size. Hill diversity comprises a spectrum of diversity metrics and is based on three key insights. First, species richness and variants of the Shannon and Simpson indices are all special cases of one general equation. Second, richness, Shannon and Simpson can be expressed on the same scale and in units of species. Third, there is no way to eliminate the effect of relative abundance from estimates of any of these diversity metrics, including species richness. Rather, a researcher must choose the relative sensitivity of the metric towards rare and common species, a concept which we describe as ‘leverage.' In this paper we explain coverage and Hill diversity, provide guidelines for how to use them together to measure species diversity, and demonstrate their use with examples from our own data. We show why researchers will obtain more robust results when they estimate the Hill diversity of equal-coverage samples, rather than using other methods such as equal-effort sampling or traditional sample rarefaction.     

Rainfall induces time-lagged changes in the proportion of tropical aquatic hosts infected with metazoan parasites

Rainfall serves as a powerful driving force, shifting temporal abundance and prevalence patterns in parasites and free-living aquatic organisms in tropical environments. However, there is a lack of sound evidence showing the temporal scales at which rainfall influences infection parameters of parasites in the tropics either directly by affecting the parasite life cycle or indirectly by modifying host population abundance. In the present study, we demonstrate that changes in rainfall patterns lead to changes in the proportion of infected hosts with several parasite species, causing immediate or lagged favourable conditions for an increase in levels of infection. However, the temporal scale of the influence of rainfall varied depending on the ecological characteristics of aquatic ecosystems. Despite the environmental heterogeneity and stochastic events (storms and hurricanes) which affect the study sites, the proportion of infected hosts shows frequency cycles on a yearly scale, suggesting that environmental changes are within the range of variability that naturally occur at the study sites. We propose that the incorporation of stochastic events into long-term predictive models is crucial for understanding the potential effects of global climate change on infection parameters of tropical parasites.     

Ecological niche modeling re‐examined: A case study with the Darwin's fox

Many previous studies have attempted to assess ecological niche modeling performance using receiver operating characteristic (ROC) approaches, even though diverse problems with this metric have been pointed out in the literature. We explored different evaluation metrics based on independent testing data using the Darwin's Fox (Lycalopex fulvipes) as a detailed case in point. Six ecological niche models (ENMs; generalized linear models, boosted regression trees, Maxent, GARP, multivariable kernel density estimation, and NicheA) were explored and tested using six evaluation metrics (partial ROC, Akaike information criterion, omission rate, cumulative binomial probability), including two novel metrics to quantify model extrapolation versus interpolation (E‐space index I) and extent of extrapolation versus Jaccard similarity (E‐space index II). Different ENMs showed diverse and mixed performance, depending on the evaluation metric used. Because ENMs performed differently according to the evaluation metric employed, model selection should be based on the data available, assumptions necessary, and the particular research question. The typical ROC AUC evaluation approach should be discontinued when only presence data are available, and evaluations in environmental dimensions should be adopted as part of the toolkit of ENM researchers. Our results suggest that selecting Maxent ENM based solely on previous reports of its performance is a questionable practice. Instead, model comparisons, including diverse algorithms and parameterizations, should be the sine qua non for every study using ecological niche modeling. ENM evaluations should be developed using metrics that assess desired model characteristics instead of single measurement of fit between model and data. The metrics proposed herein that assess model performance in environmental space (i.e., E‐space indices I and II) may complement current methods for ENM evaluation.     

Quantifying the multiple facets of isotopic diversity: New metrics for stable isotope ecology

Stable isotope analyses have emerged as an insightful tool for ecologists, with quantitative methods being developed to analyse data at the population, community and food web levels. In parallel, functional ecologists have developed metrics to quantify the multiple facets of functional diversity in a n-dimensional space based on functional traits. Here, we transferred and adapted metrics developed by functional ecologists into a set of four isotopic diversity metrics (isotopic divergence, dispersion, evenness and uniqueness) complementary to the existing metrics. Specifically, these new metrics are mathematically independent of the number of organisms analysed and account for the abundance of organisms. They can also be calculated with more than two stable isotopes. In addition, we also provide a procedure for calculating the levels of isotopic overlap (similarity and turnover) between two groups of organisms. These metrics have been implemented into new functions in R made freely available to users and we illustrated their application using stable isotope values from a freshwater fish community. Transferring the framework developed initially for measuring functional diversity to stable isotope ecology will allow more efficient assessments of changes in the multiple facets of isotopic diversity following anthropogenic disturbances.     

Effects of climate variability on the demography of wild geladas

Nonhuman primates are an essential part of tropical biodiversity and play key roles in many ecosystem functions, processes, and services. However, the impact of climate variability on nonhuman primates, whether anthropogenic or otherwise, remains poorly understood. In this study, we utilized age‐structured matrix population models to assess the population viability and demographic variability of a population of geladas (Theropithecus gelada) in the Simien Mountains, Ethiopia with the aim of revealing any underlying climatic influences. Using data from 2008 to 2019 we calculated annual, time‐averaged, and stochastic population growth rates (λ) and investigated relationships between vital rate variability and monthly cumulative rainfall and mean temperature. Our results showed that under the prevailing environmental conditions, the population will increase (λ _s = 1.021). Significant effects from rainfall and/or temperature variability were widely detected across vital rates; only the first year of infant survival and the individual years of juvenile survival were definitively unaffected. Generally, the higher temperature in the hot‐dry season led to lower survival and higher fecundity, while higher rainfall in the hot‐dry season led to increased survival and fecundity. Overall, these results provide evidence of greater effects of climate variability across a wider range of vital rates than those found in previous primate demography studies. This highlights that although primates have often shown substantial resilience to the direct effects of climate change, their vulnerability may vary with habitat type and across populations.     

Spine and dine: A key defensive trait promotes ecological success in spiny ants

A key focus of ecologists is explaining the origin and maintenance of morphological diversity and its association with ecological success. We investigate potential benefits and costs of a common and varied morphological trait, cuticular spines, for foraging behavior, interspecific competition, and predator–prey interactions in naturally co‐occurring spiny ants (Hymenoptera: Formicidae: Polyrhachis) in an experimental setting. We expect that a defensive trait like spines might be associated with more conspicuous foraging, a greater number of workers sent out to forage, and potentially increased competitive ability. Alternatively, consistent with the ecological trade‐off hypothesis, we expect that investment in spines for antipredator defense might be negatively correlated with these other ecological traits. We find little evidence for any costs to ecological traits, instead finding that species with longer spines either outperform or do not differ from species with shorter spines for all tested metrics, including resource discovery rate and foraging effort as well as competitive ability and antipredator defense. Spines appear to confer broad antipredator benefits and serve as a form of defense with undetectable costs to key ecological abilities like resource foraging and competitive ability, providing an explanation for both the ecological success of the study genus and the large number of evolutionary origins of this trait across all ants. This study also provides a rare quantitative empirical test of ecological effects related to a morphological trait in ants.     

Policy Language in Restoration Ecology

Relating restoration ecology to policy is one of the aims of the Society for Ecological Restoration and its journal Restoration Ecology. As an interdisciplinary team of researchers in both ecological science and political science, we have struggled with how policy‐relevant language is and could be deployed in restoration ecology. Using language in scientific publications that resonates with overarching policy questions may facilitate linkages between researcher investigations and decision‐makers' concerns on all levels. Climate change is the most important environmental problem of our time and to provide policymakers with new relevant knowledge on this problem is of outmost importance. To determine whether or not policy‐specific language was being included in restoration ecology science, we surveyed the field of restoration ecology from 2008 to 2010, identifying 1,029 articles, which we further examined for the inclusion of climate change as a key element of the research. We found that of the 58 articles with “climate change” or “global warming” in the abstract, only 3 identified specific policies relevant to the research results. We believe that restoration ecologists are failing to include themselves in policy formation and implementation of issues such as climate change within journals focused on restoration ecology. We suggest that more explicit reference to policies and terminology recognizable to policymakers might enhance the impact of restoration ecology on decision‐making processes.     

Designing ecological climate change impact assessments to reflect key climatic drivers

Identifying the climatic drivers of an ecological system is a key step in assessing its vulnerability to climate change. The climatic dimensions to which a species or system is most sensitive – such as means or extremes – can guide methodological decisions for projections of ecological impacts and vulnerabilities. However, scientific workflows for combining climate projections with ecological models have received little explicit attention. We review Global Climate Model (GCM) performance along different dimensions of change and compare frameworks for integrating GCM output into ecological models. In systems sensitive to climatological means, it is straightforward to base ecological impact assessments on mean projected changes from several GCMs. Ecological systems sensitive to climatic extremes may benefit from what we term the ‘model space’ approach: a comparison of ecological projections based on simulated climate from historical and future time periods. This approach leverages the experimental framework used in climate modeling, in which historical climate simulations serve as controls for future projections. Moreover, it can capture projected changes in the intensity and frequency of climatic extremes, rather than assuming that future means will determine future extremes. Given the recent emphasis on the ecological impacts of climatic extremes, the strategies we describe will be applicable across species and systems. We also highlight practical considerations for the selection of climate models and data products, emphasizing that the spatial resolution of the climate change signal is generally coarser than the grid cell size of downscaled climate model output. Our review illustrates how an understanding of how climate model outputs are derived and downscaled can improve the selection and application of climatic data used in ecological modeling.     

Deciduousness in a seasonal tropical forest in western Thailand: interannual and intraspecific variation in timing, duration and environmental cues

Seasonal tropical forests exhibit a great diversity of leaf exchange patterns. Within these forests variation in the timing and intensity of leaf exchange may occur within and among individual trees and species, as well as from year to year. Understanding what generates this diversity of phenological behaviour requires a mechanistic model that incorporates rate-limiting physiological conditions, environmental cues, and their interactions. In this study we examined long-term patterns of leaf flushing for a large proportion of the hundreds of tree species that co-occur in a seasonal tropical forest community in western Thailand. We used the data to examine community-wide variation in deciduousness and tested competing hypotheses regarding the timing and triggers of leaf flushing in seasonal tropical forests. We developed metrics to quantify the nature of deciduousness (its magnitude, timing and duration) and its variability among survey years and across a range of taxonomic levels. Tree species varied widely in the magnitude, duration, and variability of leaf loss within species and across years. The magnitude of deciduousness ranged from complete crown loss to no crown loss. Among species that lost most of their crown, the duration of deciduousness ranged from 2 to 21 weeks. The duration of deciduousness in the majority of species was considerably shorter than in neotropical forests with similar rainfall periodicity. While the timing of leaf flushing varied among species, most (∼70%) flushed during the dry season. Leaf flushing was associated with changes in photoperiod in some species and the timing of rainfall in other species. However, more than a third of species showed no clear association with either photoperiod or rainfall, despite the considerable length and depth of the dataset. Further progress in resolving the underlying internal and external mechanisms controlling leaf exchange will require targeting these species for detailed physiological and microclimatic studies.     

Annual Rainfall and Seasonality Predict Pan-tropical Patterns of Liana Density and Basal Area

We test the hypotheses proposed by Gentry and Schnitzer that liana density and basal area in tropical forests vary negatively with mean annual precipitation (MAP) and positively with seasonality. Previous studies correlating liana abundance with these climatic variables have produced conflicting results, warranting a new analysis of drivers of liana abundance based on a different dataset. We compiled a pan-tropical dataset containing 28,953 lianas (≥2.5 cm diam.) from studies conducted at 13 Neotropical and 11 Paleotropical dry to wet lowland tropical forests. The ranges in MAP and dry season length (DSL) (number of months with mean rainfall <100 mm) represented by these datasets were 860–7250 mm/yr and 0–7 mo, respectively. Pan-tropically, liana density and basal area decreased significantly with increasing annual rainfall and increased with increasing DSL, supporting the hypotheses of Gentry and Schnitzer. Our results suggest that much of the variation in liana density and basal area in the tropics can be accounted for by the relatively simple metrics of MAP and DSL.     

Research Priorities for Neotropical Dry Forests1

Our understanding of the human and biophysical dimensions of tropical dry forest change and its cumulative effects is still in the early stages of academic discovery. The papers in this special section on Neotropical dry forests cover a wide range of sites and problems ranging from the use of multispectral and hyperspectral remote sensing platforms to the impact of hurricanes on tropical dry forest regeneration. Here, we present to the scientific community the results of a workshop on which research priorities for tropical dry forests were discussed. This discussion focuses on the need to develop linkages between remote sensing, ecological, and social science research. The incorporation of social sciences into ecological research could contribute dramatically to our understandings of tropical dry forests by providing important contextual information to ecologists, and by helping to develop an important science–policy–public nexus on which environmental management can succeed.     

Temperature and rainfall interact to control carbon cycling in tropical forests

Tropical forests dominate global terrestrial carbon (C) exchange, and recent droughts in the Amazon Basin have contributed to short‐term declines in terrestrial carbon dioxide uptake and storage. However, the effects of longer‐term climate variability on tropical forest carbon dynamics are still not well understood. We synthesised field data from more than 150 tropical forest sites to explore how climate regulates tropical forest aboveground net primary productivity (ANPP) and organic matter decomposition, and combined those data with two existing databases to explore climate – C relationships globally. While previous analyses have focused on the effects of either temperature or rainfall on ANPP, our results highlight the importance of interactions between temperature and rainfall on the C cycle. In cool forests (< 20 °C), high rainfall slowed rates of C cycling, but in warm tropical forests (> 20 °C) it consistently enhanced both ANPP and decomposition. At the global scale, our analysis showed an increase in ANPP with rainfall in relatively warm sites, inconsistent with declines in ANPP with rainfall reported previously. Overall, our results alter our understanding of climate – C cycle relationships, with high precipitation accelerating rates of C exchange with the atmosphere in the most productive biome on earth.