Forbs ("wildflowers") are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.

Large herbivore communities are changing globally, with populations of wild herbivores generally declining while domestic herbivore populations are increasing, influencing ecosystem function along with the impacts of climate change. Manipulative experiments have rarely captured the interaction between patterns of large herbivore assemblage change and climatic conditions. This interaction may affect the functional traits and functional diversity of herbaceous communities; this requires investigation, as these metrics have been useful proxies for ecosystem function. We used a large herbivore exclosure experiment replicated along a topo-climatic gradient to explore the interaction between climate and herbivore assemblage on community-level functional traits and the functional diversity of herbaceous plant understories. Our findings demonstrate interacting effects between large herbivore assemblages and climate. We found a shift from drought-tolerant traits to drought-avoidant traits with increasing aridity, specifically with regard to plant leaf area and specific leaf area. We also determined that plant community responses to grazing changed from an herbivore avoidance strategy at drier sites to a more herbivore-tolerant strategy at wetter sites. We observed that the effects of herbivores on community-level traits can sometimes counteract those of climate. Finally, we found that cattle and large wild herbivores can differ in the magnitude and direction of effects on functional traits and diversity.

Large-bodied wild ungulates are declining worldwide, while domestic livestock continue to increase in abundance. Such changes in large herbivore communities should have strong effects on the control of ticks and tick-borne disease as they can indirectly modify habitat and directly serve as final hosts for ticks' lifecycles. Numerous studies have now linked changing ungulate communities to changes in tick populations and disease risk. However, the effects of changing large herbivore communities are variable across studies, and the effect of climate as a mediating factor of this variation remains poorly understood. Also, studies to date have largely focused on wildlife loss without considering the extent to which livestock additions may alter tick populations, even though livestock replacement of wildlife is the global norm. In this study, we used a large-scale exclosure experiment replicated along a topo-climatic gradient to examine the effects on tick populations of both large herbivore removal and livestock additions. We found that while questing ticks increased modestly, by 21%, when large herbivores were removed from a system they decreased more substantially, by 50%, when livestock (in the form of cattle) were added. Importantly, in addition to the direct effects of climate on tick populations, climate also mediates the effect of ungulates on questing tick density. Particularly, the addition of livestock under the most arid conditions decreased tick presence, likely due to changes in ground-level microclimates away from those beneficial to ticks. Overall, the work contributes to our understanding of tick population responses to globally common human-induced rangeland alterations under the concurrent effects of climate change.

Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them. Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure. We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments. Synthesis. Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change.

Wildfire activity is accelerating on many rangelands worldwide, yet the potential for grazing to be used as a fire management tool remains largely unknown. Particularly, little is known about the influence of grazing on ignition and initial spread of fire, as well as how these vary by differences in grazing management. We investigated effects of grazing intensity (light, moderate, high) on fuel characteristics, fire ignition, and initial spread during the wildfire season in a native-dominated shrub steppe in eastern Oregon. We found that differences in grazing intensity have differential effects on fuel profiles (cover, height, moisture, biomass) with resulting impacts on fire behavior, but these relationships varied across study years. In particular, grazing had a stronger effect on ignition probability in drier years. Fire behavior in lightly grazed plots were similar to ungrazed plots, while moderate grazing was similar to high-intensity grazing. Results of this study highlight that grazing can be useful as a tool for wildfire management, and grazing at moderate and high intensities can reduce the probability of fire propagation in native-dominated sagebrush ecosystems. Further, the effects of grazing are context dependent and therefore may depend on specific objectives and environmental conditions.

AbstractMobile animals that traverse ecosystem boundaries can fundamentally reshape environments by providing critical nutrient and energy inputs to the ecosystems they inhabit. In particular, aggregations of seabirds often transform coastal and island ecosystems through large amounts of nutrient‐rich guano deposition. Anthropogenically driven losses of these subsidies can occur through changes in abundance of mobile species, including seabirds, and have been shown to drive whole‐scale ecosystem state change on islands. However, even though many species that forage on anthropogenic food sources are highly mobile and may thus play important roles in moving nutrients from urban systems to otherwise conserved ecosystems, the impacts of anthropogenic supplements on spatial subsidies have been largely ignored. Here we examine the effects of large nesting colonies of Western Gulls (Larus occidentalis), a generalist carnivore known to forage on human refuse, on the Channel Islands of California. Specifically, we explore how their foraging on human subsidies may change nutrient deposition patterns at their relatively remote and protected breeding islands. We equipped gulls with GPS loggers to assess the frequency of urban foraging, and we partnered this tracking data with bird density data to estimate the rate of wild and urban‐derived guano deposition on two different islands. Consistent with research on other gull species, we found high (up to 40%) but island‐specific rates of urban foraging, resulting between 66 and 93 kg ha−1 of guano in these two sites during the breeding season, a level greater than half the amount of fertilizer applied annually in typical commercial agricultural settings and likely the primary source of nitrogen and phosphorus inputs to this system. Specifically, we estimate that 27 kg ha−1 year−1 of nutrient‐rich guano is shuttled to these otherwise isolated islands from anthropogenic sources. This research highlights the large shadow (i.e., footprint) that human activity can cast on even remote ecosystems by driving significant nutrient enrichment through impacts on animal behavior and connectivity.

Despite wide recognition of the importance of anthropogenically driven changes in large herbivore communities-including both declines in wildlife and increases in livestock-there remain large gaps in our knowledge about the impacts of these changes on plant communities, particularly when combined with concurrent changes in climate. Considering these prominent forms of global change in tandem enables us to better understand controls on savanna vegetation structure and diversity under real-world conditions. We conducted a field experiment using complete and semi-permeable herbivore exclosures to explore the difference in plant communities among sites with wild herbivores only, with cattle in addition to wild herbivores, and with no large herbivores. To understand variation in effects across climatic contexts, the experiment was replicated at three locations along a topoclimatic gradient in California. Critically, this is the first such experiment to compare cattle and wildlife impacts along an environmental gradient within a single controlled experiment. Vegetation structure responded strongly to herbivore treatment regardless of climate. Relative to the isolated effects of wildlife, exclusion of all large herbivores generally increased structural components related to cover and above-ground biomass while the addition of cattle led to reductions in vegetation cover, litter, shading and standing biomass. Furthermore, wildlife had a consistent neutral or positive effect on plant diversity, while the effect of livestock addition was context dependent. Cattle had a neutral to strongly negative effect at low aridity, but a positive effect at high aridity. These results suggest that (a) herbivore effects can override climate effects on vegetation structure, (b) cattle addition can drive different effects on diversity and (c) herbivore effects on diversity are modulated by climate. Synthesis. Our results illustrate very distinctive shifts in plant communities between two realistic forms of change in ungulate herbivore assemblages-livestock addition and large herbivore losses-particularly for plant diversity responses, and that these responses vary across climatic contexts. This finding has important implications for the management and protection of plant biodiversity given that over a quarter of the Earth's land area is managed for livestock and climate regimes are changing globally.

Predator-prey interactions shape ecosystem stability and are influenced by changes in ecosystem productivity. However, because multiple biotic and abiotic drivers shape the trophic responses of predators to productivity, we often observe patterns, but not mechanisms, by which productivity drives food web structure. One way to capture mechanisms shaping trophic responses is to quantify trophic interactions among multiple trophic groups and by using complementary metrics of trophic ecology. In this study, we combine two diet-tracing methods: diet DNA and stable isotopes, for two trophic groups (top predators and intermediate predators) in both low- and high-productivity habitats to elucidate where in the food chain trophic structure shifts in response to changes in underlying ecosystem productivity. We demonstrate that while top predators show increases in isotopic trophic position (δ15N) with productivity, neither their isotopic niche size nor their DNA diet composition changes. Conversely, intermediate predators show clear turnover in DNA diet composition towards a more predatory prey base in high-productivity habitats. Taking this multi-trophic approach highlights how predator identity shapes responses in predator-prey interactions across environments with different underlying productivity, building predictive power for understanding the outcomes of ongoing anthropogenic change.

Predator-prey interactions shape ecosystems and can help maintain biodiversity. However, for many of the earth's most biodiverse and abundant organisms, including terrestrial arthropods, these interactions are difficult or impossible to observe directly with traditional approaches. Based on previous theory, it is likely that predator-prey interactions for these organisms are shaped by a combination of predator traits, including body size and species-specific hunting strategies. In this study, we combined diet DNA metabarcoding data of 173 individual invertebrate predators from nine species (a total of 305 individual predator-prey interactions) with an extensive community body size data set of a well-described invertebrate community to explore how predator traits and identity shape interactions. We found that (1) mean size of prey families in the field usually scaled with predator size, with species-specific variation to a general size-scaling relationship (exceptions likely indicating scavenging or feeding on smaller life stages). We also found that (2) although predator hunting traits, including web and venom use, are thought to shape predator-prey interaction outcomes, predator identity more strongly influenced our indirect measure of the relative size of predators and prey (predator:prey size ratios) than either of these hunting traits. Our findings indicate that predator body size and species identity are important in shaping trophic interactions in invertebrate food webs and could help predict how anthropogenic biodiversity change will influence terrestrial invertebrates, the earth's most diverse animal taxonomic group.

Invasive rodent eradications are frequently undertaken to curb island biodiversity loss. However, the breadth of rodents’ ecological impact, even after eradication, is not always fully recognized. For example, the most widespread invasive rodent, the black rat (Rattus rattus), while omnivorous, eats predominantly seeds and fruit. Yet, the effects of seed predation release after eradication on plant communities and ecological functions are not well understood, posing a gap for island restoration. We examined the role of seed predation release following black rat eradication in changes to tree composition and aboveground biomass across an islet network (Palmyra Atoll) in the Central Pacific. We conducted repeated surveys of seed, juvenile, and adult tree biomass and survival in permanent vegetation plots before and after the eradication of rats. We observed a 95% reduction in seed predation for an introduced, previously cultivated tree population (Cocos nucifera). Juvenile tree biomass of all species increased 14‐fold, with C. nucifera increasing the most, suggesting that eradication increased this tree's competitive advantage. Indeed, based on stage‐structured demographic models, rat eradication led to a 10% increase in C. nucifera population growth rate. The effect of invasive rodent seed predation varies considerably among the plant species in a community and can shift competitive dynamics, sometimes in favor of invasive plants. These bottom‐up effects should be considered in evaluating the costs and benefits of eradication. Documenting the variation in invasive rodent diet items, along with long‐term surveys, can help prioritize island eradications where restoration is most likely to be successful.

The perpetuation of tree populations is dependent on successful seedling establishment, which is in turn controlled by biotic and climatic factors. California's endemic oak species face limited recruitment and shrinking ranges due to anthropogenic climate change and land use, both of which can negatively impact germination and seedling survival. In this study, we investigated Quercus seed germination and seedling establishment under three large herbivore regimes (no large herbivores, wild herbivores, and both wild herbivores and cattle) across three sites representing present, near-future, and far-future projections for climate in California's montane oak woodlands. Seed germination and living seedling abundance were highest in the intermediate and mesic climate levels, where total exclosures resulted in the best outcomes for seedlings. In contrast, in the arid level, the best large herbivore regime for establishment consisted of wild herbivores and no cattle. Seedling mortality was highest in the most arid climate level, and within that level, mortality was reduced by over 25% in both the wildlife and wildlife plus cattle treatments compared to the total exclosures. Our results suggest that increasing aridity will limit oak establishment and modulate the impact of large herbivores on oak seedlings, with wild large herbivores becoming a beneficial factor for seedling abundance, and both wild herbivores and cattle reducing mortality. Dynamic and contextually informed management of large herbivore populations and other restoration actions will be necessary to ensure the sustainability of California's oak populations.

Invasive plants can have significant negative interactions with native flora and fauna, often decreasing the abundance and diversity of native plants and invertebrate and vertebrate herbivores. Less is known, however, about how invasive plants influence higher-order consumers, such as carnivores. Arundo donax is a globally distributed invasive grass that forms dense monotypic stands in the habitats where it successfully establishes. This study investigated the influence of Arundo invasion on mammal assemblages in California’s Santa Clara River Valley. Specifically, we aimed to determine whether Arundo alters habitat use for carnivores, and if so, assess the possible mechanisms driving these differences. We used remote cameras to evaluate carnivore habitat use and model occupancy, and live traps and remote cameras to assay abundance of small mammals, common prey of these larger predators. We found that detections of all carnivores were significantly lower in Arundo-dominated habitat patches, suggesting decreased preference for Arundo habitat. This was reflected in the occupancy model, which found the highest probability of coyote and bobcat habitat use in native patches and lowest in Arundo; however, habitat did not seem to be as important of a predictor as in the pooled species results. Small mammal abundance was similar if not higher in Arundo, suggesting the possibility of Arundo acting as a refuge for prey species and altered predator–prey dynamics in Arundo-dominated patches. This study improves our knowledge of the often-complex ways that invasive plants, like Arundo, influence the ecology and behavior of faunal communities in invaded ecosystems.

A large part of ecosystem function in woodland systems depends on soil fungal communities. However, global climate change has the potential to fundamentally alter these communities as fungal species are filtered with changing environmental conditions. In this study, we examined the potential effects of climate on host-associated (i.e. tree-associated) soil fungal communities at climatically distinct sites in the Tehachapi Mountains in California, where more arid conditions represent likely regional climate futures. We found that soil fungal community composition changes strongly across sites, with species richness and diversity being highest at the most arid site. However, host association may buffer the effects of climate on community composition, as host-associated fungal communities are more similar to each other across climatically distinct sites than the whole fungal community. Lastly, an examination of functional traits for ectomycorrhizal fungi, a well-studied guild of fungal mutualist species, showed that stress-tolerant traits were more abundant at arid sites than mesic sites, providing a mechanistic understanding of these community patterns. Taken together, our results indicate that fungal community composition will likely shift with future climate change but that host association may buffer these effects, with shifts in functional traits having implications for future ecosystem function.

Recent increases in human disturbance pose significant threats to migratory species using collective movement strategies. Key threats to migrants may differ depending on behavioural traits (e.g. collective navigation), taxonomy and the environmental system (i.e. freshwater, marine or terrestrial) associated with migration. We quantitatively assess how collective navigation, taxonomic membership and environmental system impact species' vulnerability by (i) evaluating population change in migratory and non-migratory bird, mammal and fish species using the Living Planet Database (LPD), (ii) analysing the role of collective navigation and environmental system on migrant extinction risk using International Union for Conservation of Nature (IUCN) classifications and (iii) compiling literature on geographical range change of migratory species. Likelihood of population decrease differed by taxonomic group: migratory birds were more likely to experience annual declines than non-migrants, while mammals displayed the opposite pattern. Within migratory species in IUCN, we observed that collective navigation and environmental system were important predictors of extinction risk for fishes and birds, but not for mammals, which had overall higher extinction risk than other taxa. We found high phylogenetic relatedness among collectively navigating species, which could have obscured its importance in determining extinction risk. Overall, outputs from these analyses can help guide strategic interventions to conserve the most vulnerable migrations. This article is part of the theme issue ‘Collective movement ecology'.

Table with all migratory species in GROMS and IUCN, with migratory coding, collective coding, realms, environmental systems, IUCN categories, and taxonomic information.

An ecologist reflects on the unexpected twists and turns that shaped his scientific career

In his new book, Serendipity , eminent ecologist James A. Estes deftly weaves rigorous science with personal reflection to create an absorbing and introspective read that is equal parts memoir, ecological textbook, and motivational guidebook for young ecologists. Emphasizing the role of chance encounters in creating career opportunities, the book serves as a revitalizing reminder of the self-doubt and exhilaration that go hand-in-hand with scientific discovery.