Integrated analysis shows how the effects of extreme flooding events propagate through fish communities to impact amphibians.

Research Highlight: Davis, C. L., Walls, S. C., Barichivich, W. J., Brown, M. E., & Miller, D. A. (2022). Disentangling direct and indirect effects of extreme events on coastal wetland communities. Journal of Animal Ecology, https://doi.org/10.1111/1365-2656.13874. Catastrophic events such as floods, hurricanes, winter storms, droughts and wildfires increasingly touch our lives either directly or indirectly. These events draw our attention to the seriousness of changes in climate not only to human well-being but also to the integrity of ecological systems upon which we depend. Understanding the impacts of extreme events on ecological systems requires the ability to characterize the cascading effects of environmental changes on the environments in which organisms live and the altered biological interactions produced. This scientific ambition represents no small challenge for the study of animal communities, which are typically difficult to census as well as dynamic in time and space. Davis et al. (2022) in a recent study in the Journal of Animal Ecology examined the amphibian and fish communities found in depressional coastal wetlands to better understand how they respond to major rainfall and flooding events. Data from the U.S. Geological Survey's Amphibian Research and Monitoring Initiative provided an 8-year record of observations as well as environmental measurements. For this study, the authors integrated techniques for assessing the dynamics of animal populations with a Bayesian implementation of structural equation modelling. Using their integrated methodological approach permitted the authors to reveal the direct and indirect effects of extreme weather events on co-occurring amphibian and fish communities while accounting for observational uncertainty and temporal variation in population-level processes. Their findings indicate that the most prominent effects of flooding on the amphibian community were caused by changes in the fish community that led to increased predation and resource competition. In their conclusions, the authors emphasize the importance of understanding networks of abiotic and biotic effects if we are to predict and mitigate the influence of extreme weather events.

From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence.

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.

Migration and transformation of coastal wetlands in response to rising seas.

Coastal wetlands are not only among the world's most valued ecosystems but also among the most threatened by high greenhouse gas emissions that lead to accelerated sea level rise. There is intense debate regarding the extent to which landward migration of wetlands might compensate for seaward wetland losses. By integrating data from 166 estuaries across the conterminous United States, we show that landward migration of coastal wetlands will transform coastlines but not counter seaward losses. Two-thirds of potential migration is expected to occur at the expense of coastal freshwater wetlands, while the remaining one-third is expected to occur at the expense of valuable uplands, including croplands, forests, pastures, and grasslands. Our analyses underscore the need to better prepare for coastal transformations and net wetland loss due to rising seas.

Biodiversity effects on grape quality depend on variety and management intensity.

Interactions between plants can be beneficial, detrimental or neutral. In agricultural systems, competition between crop and spontaneous vegetation is a major concern. We evaluated the relative support for three non-exclusive ecological hypotheses about interactions between crop and spontaneous plants based on competition, complementarity or facilitation.The study was conducted in Swiss vineyards with different vegetation management intensities. In all, 33 vineyards planted with two different grape varieties were studied over 3 years to determine whether low-intensity vegetation management might provide benefits for grape quality parameters. Management intensity varied with the degree of control of spontaneous inter-row vegetation. Features of spontaneous vegetation measured included total cover, total species richness and abundance of nitrogen-fixing plants. Grape quality parameters of known importance to wine making (yeast assimilable nitrogen, sugars, tartaric acid and malic acid) were determined by Fourier-transform infrared spectroscopy (FTIR). Using structural equation modelling, we evaluated hypotheses about the multivariate responses of grape quality parameters as well as the direct and indirect (plant-mediated) effects of management.Observed effects of management differed between grape varieties. Management intensity and abundance of N-fixing plants significantly influenced grape quality parameters while total richness of spontaneous plants did not have detectable effects. Abundance of N-fixing plants was enhanced by low-intensity management resulting in increased N content in the red grape variety Pinot noir, potentially enhancing grape quality, while measured soil N content did not explain the increase.Synthesis and applications. Our study shows that crop quality can be enhanced by spontaneous plants, in this case by the abundance of a key functional group (N-fixers), most likely through plant-plant or plant-microbe facilitation. However, beneficial interactions may have a high specificity in terms of facilitation partners and may have contrasting effects at low taxonomic resolutions such as crop varieties. Generally, increasing plant biodiversity in agricultural systems may increase competition with crops. Thus, the identification of suitable interaction partners and a careful balance between crop variety and spontaneous plant species may be necessary to utilize beneficial interactions and to reduce the trade-off between agricultural production and biodiversity to achieve a sustainable ecological benefit in agricultural systems.

The importance of natural versus human factors for ecological conditions of streams and rivers.

Streams are influenced by watershed-scale factors, such as climate, geology, topography, hydrology, and soils, which mostly vary naturally among sites, as well as human factors, agriculture and urban development. Thus, natural factors could complicate assessment of human disturbance. In the present study, we use structural equation modeling and data from the 2008-2009 United States National Rivers and Streams Assessment to quantify the relative importance of watershed-scale natural and human factors for in-stream conditions. We hypothesized that biological condition, represented using a diatom multimetric index (MMI), is directly affected by in-stream physicochemical environment, which in turn is regulated by natural and human factors. We evaluated this hypothesis at both national and ecoregion scales to understand how influences vary among regions. We found that direct influences of in-stream environment on diatom MMIs were greater than natural and human factors at the national scale and in all but one ecoregion. Meanwhile, in-stream environments were jointly explained by natural variations in precipitation, base flow index, hydrological stability, % volcanic rock, soil water table depth, and soil depth and by human factors measured as % crops, % other agriculture, and % urban land use. The explained variance of in-stream environment by natural and human factors ranged from 0.30 to 0.75, for which natural factors independently accounted for the largest proportion of explained variance at the national scale and in seven ecoregions. Covariation between natural and human factors accounted for a higher proportion of explained variance of in-stream environment than unique effects of human factors in most ecoregions. Ecoregions with relatively weak effects by human factors had relatively high levels of covariance, high levels of human disturbance, or small ranges in human disturbance. We conclude that accounting for effects of natural factors and their covariation with human factors will be important for accurate ecological assessments.

Scientist's guide to developing explanatory statistical models using causal analysis principles.

Recent discussions of model selection and multimodel inference highlight a general challenge for researchers: how to convey the explanatory content of a hypothesized model or set of competing models clearly. The advice from statisticians for scientists employing multimodel inference is to develop a well-thought-out set of candidate models for comparison, though precise instructions for how to do that are typically not given. A coherent body of knowledge, which falls under the general term causal analysis, now exists for examining the explanatory scientific content of candidate models. Much of the literature on causal analysis has been recently developed, and we suspect may not be familiar to many ecologists. This body of knowledge comprises a set of graphical tools and axiomatic principles to support scientists in their endeavors to create "well-formed hypotheses," as statisticians are asking them to do. Causal analysis is complementary to methods such as structural equation modeling, which provides the means for evaluation of proposed hypotheses against data. In this paper, we summarize and illustrate a set of principles that can guide scientists in their quest to develop explanatory hypotheses for evaluation. The principles presented in this paper have the capacity to close the communication gap between statisticians, who urge scientists to develop well-thought-out coherent models, and scientists, who would like some practical advice for exactly how to do that.

Climate and plant controls on soil organic matter in coastal wetlands.

Coastal wetlands are among the most productive and carbon-rich ecosystems on Earth. Long-term carbon storage in coastal wetlands occurs primarily belowground as soil organic matter (SOM). In addition to serving as a carbon sink, SOM influences wetland ecosystem structure, function, and stability. To anticipate and mitigate the effects of climate change, there is a need to advance understanding of environmental controls on wetland SOM. Here, we investigated the influence of four soil formation factors: climate, biota, parent materials, and topography. Along the northern Gulf of Mexico, we collected wetland plant and soil data across elevation and zonation gradients within 10 estuaries that span broad temperature and precipitation gradients. Our results highlight the importance of climate-plant controls and indicate that the influence of elevation is scale and location dependent. Coastal wetland plants are sensitive to climate change; small changes in temperature or precipitation can transform coastal wetland plant communities. Across the region, SOM was greatest in mangrove forests and in salt marshes dominated by graminoid plants. SOM was lower in salt flats that lacked vascular plants and in salt marshes dominated by succulent plants. We quantified strong relationships between precipitation, salinity, plant productivity, and SOM. Low precipitation leads to high salinity, which limits plant productivity and appears to constrain SOM accumulation. Our analyses use data from the Gulf of Mexico, but our results can be related to coastal wetlands across the globe and provide a foundation for predicting the ecological effects of future reductions in precipitation and freshwater availability. Coastal wetlands provide many ecosystem services that are SOM dependent and highly vulnerable to climate change. Collectively, our results indicate that future changes in SOM and plant productivity, regulated by cascading effects of precipitation on freshwater availability and salinity, could impact wetland stability and affect the supply of some wetland ecosystem services.

Herbivory and eutrophication mediate grassland plant nutrient responses across a global climatic gradient.

Plant stoichiometry, the relative concentration of elements, is a key regulator of ecosystem functioning and is also being altered by human activities. In this paper we sought to understand the global drivers of plant stoichiometry and compare the relative contribution of climatic vs. anthropogenic effects. We addressed this goal by measuring plant elemental (C, N, P and K) responses to eutrophication and vertebrate herbivore exclusion at eighteen sites on six continents. Across sites, climate and atmospheric N deposition emerged as strong predictors of plot-level tissue nutrients, mediated by biomass and plant chemistry. Within sites, fertilization increased total plant nutrient pools, but results were contingent on soil fertility and the proportion of grass biomass relative to other functional types. Total plant nutrient pools diverged strongly in response to herbivore exclusion when fertilized; responses were largest in ungrazed plots at low rainfall, whereas herbivore grazing dampened the plant community nutrient responses to fertilization. Our study highlights (1) the importance of climate in determining plant nutrient concentrations mediated through effects on plant biomass, (2) that eutrophication affects grassland nutrient pools via both soil and atmospheric pathways and (3) that interactions among soils, herbivores and eutrophication drive plant nutrient responses at small scales, especially at water-limited sites.

Associations between Urban Sprawl and Life Expectancy in the United States.

In recent years, the United States has had a relatively poor performance with respect to life expectancy compared to the other developed nations. Urban sprawl is one of the potential causes of the high rate of mortality in the United States. This study investigated cross-sectional associations between sprawl and life expectancy for metropolitan counties in the United States in 2010. In this study, the measure of life expectancy in 2010 came from a recently released dataset of life expectancies by county. This study modeled average life expectancy with a structural equation model that included five mediators: annual vehicle miles traveled (VMT) per household, average body mass index, crime rate, and air quality index as mediators of sprawl, as well as percentage of smokers as a mediator of socioeconomic status. After controlling for sociodemographic characteristics, this study found that life expectancy was significantly higher in compact counties than in sprawling counties. Compactness affects mortality directly, but the causal mechanism is unclear. For example, it may be that sprawling areas have higher traffic speeds and longer emergency response times, lower quality and less accessible health care facilities, or less availability of healthy foods. Compactness affects mortality indirectly through vehicle miles traveled, which is a contributor to traffic fatalities, and through body mass index, which is a contributor to many chronic diseases. This study identified significant direct and indirect associations between urban sprawl and life expectancy. These findings support further research and practice aimed at identifying and implementing changes to urban planning designed to support health and healthy behaviors.

Context-dependent interactions and the regulation of species richness in freshwater fish.

Species richness is regulated by a complex network of scale-dependent processes. This complexity can obscure the influence of limiting species interactions, making it difficult to determine if abiotic or biotic drivers are more predominant regulators of richness. Using integrative modeling of freshwater fish richness from 721 lakes along an 11o latitudinal gradient, we find negative interactions to be a relatively minor independent predictor of species richness in lakes despite the widespread presence of predators. Instead, interaction effects, when detectable among major functional groups and 231 species pairs, were strong, often positive, but contextually dependent on environment. These results are consistent with the idea that negative interactions internally structure lake communities but do not consistently 'scale-up' to regulate richness independently of the environment. The importance of environment for interaction outcomes and its role in the regulation of species richness highlights the potential sensitivity of fish communities to the environmental changes affecting lakes globally.

Fatal Attraction? Intraguild Facilitation and Suppression among Predators.

Competition and suppression are recognized as dominant forces that structure predator communities. Facilitation via carrion provisioning, however, is a ubiquitous interaction among predators that could offset the strength of suppression. Understanding the relative importance of these positive and negative interactions is necessary to anticipate community-wide responses to apex predator declines and recoveries worldwide. Using state-sponsored wolf (Canis lupus) control in Alaska as a quasi experiment, we conducted snow track surveys of apex, meso-, and small predators to test for evidence of carnivore cascades (e.g., mesopredator release). We analyzed survey data using an integrative occupancy and structural equation modeling framework to quantify the strengths of hypothesized interaction pathways, and we evaluated fine-scale spatiotemporal responses of nonapex predators to wolf activity clusters identified from radio-collar data. Contrary to the carnivore cascade hypothesis, both meso- and small predator occupancy patterns indicated guild-wide, negative responses of nonapex predators to wolf abundance variations at the landscape scale. At the local scale, however, we observed a near guild-wide, positive response of nonapex predators to localized wolf activity. Local-scale association with apex predators due to scavenging could lead to landscape patterns of mesopredator suppression, suggesting a key link between occupancy patterns and the structure of predator communities at different spatial scales.

A decade of insights into grassland ecosystem responses to global environmental change.

Earth's biodiversity and carbon uptake by plants, or primary productivity, are intricately interlinked, underlie many essential ecosystem processes, and depend on the interplay among environmental factors, many of which are being changed by human activities. While ecological theory generalizes across taxa and environments, most empirical tests of factors controlling diversity and productivity have been observational, single-site experiments, or meta-analyses, limiting our understanding of variation among site-level responses and tests of general mechanisms. A synthesis of results from ten years of a globally distributed, coordinated experiment, the Nutrient Network (NutNet), demonstrates that species diversity promotes ecosystem productivity and stability, and that nutrient supply and herbivory control diversity via changes in composition, including invasions of non-native species and extinction of native species. Distributed experimental networks are a powerful tool for tests and integration of multiple theories and for generating multivariate predictions about the effects of global changes on future ecosystems.

Disentangling vegetation diversity from climate-energy and habitat heterogeneity for explaining animal geographic patterns.

Broad-scale animal diversity patterns have been traditionally explained by hypotheses focused on climate-energy and habitat heterogeneity, without considering the direct influence of vegetation structure and composition. However, integrating these factors when considering plant-animal correlates still poses a major challenge because plant communities are controlled by abiotic factors that may, at the same time, influence animal distributions. By testing whether the number and variation of plant community types in Europe explain country-level diversity in six animal groups, we propose a conceptual framework in which vegetation diversity represents a bridge between abiotic factors and animal diversity. We show that vegetation diversity explains variation in animal richness not accounted for by altitudinal range or potential evapotranspiration, being the best predictor for butterflies, beetles, and amphibians. Moreover, the dissimilarity of plant community types explains the highest proportion of variation in animal assemblages across the studied regions, an effect that outperforms the effect of climate and their shared contribution with pure spatial variation. Our results at the country level suggest that vegetation diversity, as estimated from broad-scale classifications of plant communities, may contribute to our understanding of animal richness and may be disentangled, at least to a degree, from climate-energy and abiotic habitat heterogeneity.

Comment on "Worldwide evidence of a unimodal relationship between productivity and plant species richness".

Fraser et al. (Reports, 17 July 2015, p. 302) report a unimodal relationship between productivity and species richness at regional and global scales, which they contrast with the results of Adler et al. (Reports, 23 September 2011, p. 1750). However, both data sets, when analyzed correctly, show clearly and consistently that productivity is a poor predictor of local species richness.

Integrative modelling reveals mechanisms linking productivity and plant species richness.

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

Beyond just sea-level rise: considering macroclimatic drivers within coastal wetland vulnerability assessments to climate change.

Due to their position at the land-sea interface, coastal wetlands are vulnerable to many aspects of climate change. However, climate change vulnerability assessments for coastal wetlands generally focus solely on sea-level rise without considering the effects of other facets of climate change. Across the globe and in all ecosystems, macroclimatic drivers (e.g., temperature and rainfall regimes) greatly influence ecosystem structure and function. Macroclimatic drivers have been the focus of climate change-related threat evaluations for terrestrial ecosystems, but largely ignored for coastal wetlands. In some coastal wetlands, changing macroclimatic conditions are expected to result in foundation plant species replacement, which would affect the supply of certain ecosystem goods and services and could affect ecosystem resilience. As examples, we highlight several ecological transition zones where small changes in macroclimatic conditions would result in comparatively large changes in coastal wetland ecosystem structure and function. Our intent in this communication is not to minimize the importance of sea-level rise. Rather, our overarching aim is to illustrate the need to also consider macroclimatic drivers within vulnerability assessments for coastal wetlands.

Landscape structure affects specialists but not generalists in naturally fragmented grasslands.

Understanding how biotic communities respond to landscape spatial structure is critically important for conservation management as natural habitats become increasingly fragmented. However, empirical studies of the effects of spatial structure on plant species richness have found inconsistent results, suggesting that more comprehensive approaches are needed. We asked how landscape structure affects total plant species richness and the richness of a guild of specialized plants in a multivariate context. We sampled herbaceous plant communities at 56 dolomite glades (insular, fire-adapted grasslands) across the Missouri Ozarks, USA, and used structural equation modeling (SEM) to analyze the relative importance of landscape structure, soil resource availability, and fire history for plant communities. We found that landscape spatial structure, defined as the area-weighted proximity of glade habitat surrounding study sites (proximity index), had a significant effect on total plant species richness, but only after we controlled for environmental covariates. Richness of specialist species, but not generalists, was positively related to landscape spatial structure. Our results highlight that local environmental filters must be considered to understand the influence of landscape structure on communities and that unique species guilds may respond differently to landscape structure than the community as a whole. These findings suggest that both local environment and landscape context should be considered when developing management strategies for species of conservation concern in fragmented habitats.

Temporal shifts in top-down vs. bottom-up control of epiphytic algae in a seagrass ecosystem.

In coastal marine food webs, small invertebrate herbivores (mesograzers) have long been hypothesized to occupy an important position facilitating dominance of habitat-forming macrophytes by grazing competitively superior epiphytic algae. Because of the difficulty of manipulating mesograzers in the field, however, their impacts on community organization have rarely been rigorously documented. Understanding mesograzer impacts has taken on increased urgency in seagrass systems due to declines in seagrasses globally, caused in part by widespread eutrophication favoring seagrass overgrowth by faster-growing algae. Using cage-free field experiments in two seasons (fall and summer), we present experimental confirmation that mesograzer reduction and nutrients can promote blooms of epiphytic algae growing on eelgrass (Zostera marina). In this study, nutrient additions increased epiphytes only in the fall following natural decline of mesograzers. In the summer, experimental mesograzer reduction stimulated a 447% increase in epiphytes, appearing to exacerbate seasonal dieback of eelgrass. Using structural equation modeling, we illuminate the temporal dynamics of complex interactions between macrophytes, mesograzers, and epiphytes in the summer experiment. An unexpected result emerged from investigating the interaction network: drift macroalgae indirectly reduced epiphytes by providing structure for mesograzers, suggesting that the net effect of macroalgae on seagrass depends on macroalgal density. Our results show that mesograzers can control proliferation of epiphytic algae, that top-down and bottom-up forcing are temporally variable, and that the presence of macroalgae can strengthen top-down control of epiphytic algae, potentially contributing to eelgrass persistence.