Abstract Ecosystems are rarely at steady state, yet most theory predicting universal biodiversity patterns assumes they are. Here, we test whether and how eco-evolutionary dynamics drive departures from steady state by combining arthropod community data from the geologic chronosequence of the Hawaiian Archipelago with the Maximum Entropy Theory of Ecology (METE), a minimalist steady-state framework that simultaneously predicts species abundance distributions (SADs) and individual metabolic rate distributions (IPDs). The chronosequence of the Hawaiian Archipelago has yielded insights into eco-evolutionary processes because ecosystems growing on different aged substrates offer snapshots of community assembly with different histories. We find that deviations from METE peak at geologically middle-aged sites (150 Kya–1.4 Mya), consistent with active adaptive radiation pushing communities away from statistical steady state. Within-site β -diversity, which also peaks at middle-aged sites, robustly predicts deviations from METE across all sites, while the proportion of non-native species predicts deviations only after excluding the geologically youngest site. Partitioning β -diversity between native and non-native species resolves this discrepancy: at the youngest site, non-native species are distributed homogeneously and do not elevate β -diversity despite their high proportional representation. Together, these results are consistent with a trajectory from young, dispersal-assembled communities near statistical steady state, through an eco-evolutionary non-steady-state transition driven by diversification, to a new stable steady state at the oldest sites. Our findings suggest that periods of active diversification create windows of ecological instability that may facilitate biological invasion, with implications for understanding invasion dynamics in biodiversity hotspots.

Abstract Why do some lineages diversify while others do not? This remains a central question in evolutionary ecology. A long-standing assumption, dating to Darwin and embedded in the Unified Neutral Theory of Biodiversity, holds that abundant species should speciate at higher rates. Conversely, theoretical and empirical work highlights the possibility that rare and dispersal-limited clades might be more prone to speciation. Using a birth-death-immigration model with protracted speciation in a multi-population landscape connected by limited dispersal, we show that abundance has a hump-shaped effect on probability of speciation. Our model reveals that intermediate abundance maximizes speciation probability because larger populations disperse more, swamping regional differentiation and inhibiting speciation completion, while smaller populations lack the persistence and incipient speciation needed to diversify. We find empirical support for this prediction with an analysis of data from arthropods endemic to Hawai’i, where genus-level species richness shows a significant hump-shaped relationship with mean genus abundance. These findings provide a mechanistic explanation for a nuanced relationship between abundance and diversification.

Nutrient enrichment typically causes local plant diversity declines. A common but untested expectation is that nutrient enrichment also reduces variation in nutrient conditions among localities and selects for a smaller pool of species, causing greater diversity declines at larger than local scales and thus biotic homogenization. Here we apply a framework that links changes in species richness across scales to changes in the numbers of spatially restricted and widespread species for a standardized nutrient addition experiment across 72 grasslands on six continents. Overall, we find proportionally similar species loss at local and larger scales, suggesting similar declines of spatially restricted and widespread species, and no biotic homogenization after 4 years and up to 14 years of treatment. These patterns of diversity changes are generally consistent across species groups. Thus, nutrient enrichment poses threats to plant diversity, including for widespread species that are often critical for ecosystem functions.

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.

Global change is associated with variable shifts in the annual production of aboveground plant biomass, suggesting localized sensitivities with unclear causal origins. Combining remotely sensed normalized difference vegetation index data since the 1980s with contemporary field data from 84 grasslands on 6 continents, we show a widening divergence in site-level biomass ranging from +51% to -34% globally. Biomass generally increased in warmer, wetter and species-rich sites with longer growing seasons and declined in species-poor arid areas. Phenological changes were widespread, revealing substantive transitions in grassland seasonal cycling. Grazing, nitrogen deposition and plant invasion were prevalent in some regions but did not predict overall trends. Grasslands are undergoing sizable changes in production, with implications for food security, biodiversity and carbon storage especially in arid regions where declines are accelerating. Remotely sensed NDVI data and contemporary field data from 84 grasslands on 6 continents show increasing divergence in aboveground plant biomass between sites in different bioclimatic regions.

The dynamic structure of ecological communities results from interactions among taxa that change with shifts in species composition in space and time. However, our ability to study the interplay of ecological and evolutionary processes on community assembly remains relatively unexplored due to the difficulty of measuring community structure over long temporal scales. Here, we made use of a geological chronosequence across the Hawaiian Islands, representing 50 years to 4.15 million years of ecosystem development, to sample 11 communities of arthropods and their associated plant taxa using semiquantitative DNA metabarcoding. We then examined how ecological communities changed with community age by calculating quantitative network statistics for bipartite networks of arthropod-plant associations. The average number of interactions per species (linkage density), ratio of plant to arthropod species (vulnerability) and uniformity of energy flow (interaction evenness) increased significantly in concert with community age. The index of specialization H 2 ' has a curvilinear relationship with community age. Our analyses suggest that younger communities are characterized by fewer but stronger interactions, while biotic associations become more even and diverse as communities mature. These shifts in structure became especially prominent on East Maui (~0.5 million years old) and older volcanos, after enough time had elapsed for adaptation and specialization to act on populations in situ. Such natural progression of specialization during community assembly is probably impeded by the rapid infiltration of non-native species, with special risk to younger or more recently disturbed communities that are composed of fewer specialized relationships.

Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change.

Human activities are altering ecological communities around the globe. Understanding the implications of these changes requires that we consider the composition of those communities. However, composition can be summarized by many metrics which in turn are influenced by different ecological processes. For example, incidence-based metrics strongly reflect species gains or losses, while abundance-based metrics are minimally affected by changes in the abundance of small or uncommon species. Furthermore, metrics might be correlated with different predictors. We used a globally distributed experiment to examine variation in species composition within 60 grasslands on six continents. Each site had an identical experimental and sampling design: 24 plots x 4 years. We expressed compositional variation within each site-not across sites-using abundance- and incidence-based metrics of the magnitude of dissimilarity (Bray-Curtis and Sorensen, respectively), abundance- and incidence-based measures of the relative importance of replacement (balanced variation and species turnover, respectively), and species richness at two scales (per plot-year [alpha] and per site [gamma]). Average compositional variation among all plot-years at a site was high and similar to spatial variation among plots in the pretreatment year, but lower among years in untreated plots. For both types of metrics, most variation was due to replacement rather than nestedness. Differences among sites in overall within-site compositional variation were related to several predictors. Environmental heterogeneity (expressed as the CV of total aboveground plant biomass in unfertilized plots of the site) was an important predictor for most metrics. Biomass production was a predictor of species turnover and of alpha diversity but not of other metrics. Continentality (measured as annual temperature range) was a strong predictor of Sorensen dissimilarity. Metrics of compositional variation are moderately correlated: knowing the magnitude of dissimilarity at a site provides little insight into whether the variation is driven by replacement processes. Overall, our understanding of compositional variation at a site is enhanced by considering multiple metrics simultaneously. Monitoring programs that explicitly incorporate these implications, both when designing sampling strategies and analyzing data, will have a stronger ability to understand the compositional variation of systems and to quantify the impacts of human activities.

Forest structural diversity and its spatiotemporal variability are constrained by environmental and biological factors, including species pools, climate, land-use history, and legacies of disturbance regimes. These factors influence forest responses to disturbances and their interactions with structural diversity, potentially creating structurally mediated emergent properties at local to continental spatial scales and over evolutionary time. Here, we present a conceptual framework for exploring the emergent properties that arise from interactions between forest structural diversity and disturbances. We synthesize and present definitions for key terms, including emergent property, disturbance, and resilience, and highlight various types and examples of emergent properties, such as (1) interactions with species composition, (2) interactions with disturbance frequency and intensity, and (3) evolutionary changes to communities. Although emergent properties in forest ecosystems remain poorly understood, we describe a foundation for study and applied management of forest structural diversity to enhance forest restoration and resilience.

Once every 13 or 17 years within eastern North American deciduous forests, billions of periodical cicadas concurrently emerge from the soil and briefly satiate a diverse array of naive consumers, offering a rare opportunity to assess the cascading impacts of an ecosystem-wide resource pulse on a complex food web. We quantified the effects of the 2021 Brood X emergence and report that more than 80 bird species opportunistically switched their foraging to include cicadas, releasing herbivorous insects from predation and essentially doubling both caterpillar densities and accumulated herbivory levels on host oak trees. These short-lived but massive emergence events help us to understand how resource pulses can rewire interaction webs and disrupt energy flows in ecosystems, with potentially long-lasting effects.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

BACKGROUND AND AIMS:How well plants reproduce near their geographic range edge can determine whether distributions will shift in response to changing climate. Reproduction at the range edge can be limiting if pollinator scarcity leads to pollen limitation, or if abiotic stressors affect allocation to reproduction. For many animal-pollinated plants with expanding ranges, the mechanisms by which they have overcome these barriers are poorly understood. METHODS:In this study, we examined plant-pollinator interactions hypothesized to impact reproduction of the black mangrove, Avicennia germinans, which is expanding northward in coastal Florida, USA. We monitored insects visiting A. germinans populations varying in proximity to the geographic range edge, measured the pollen loads of the most common insect taxa and pollen receipt by A. germinans stigmas, and quantified flower and propagule production. KEY RESULTS:We found that despite an 84 % decline in median floral visits by insects at northernmost versus southernmost sites, range-edge pollen receipt remained high. Notably, local floral visitor assemblages exhibited substantial turnover along the study's latitudinal gradient, with large-bodied bees and hover flies increasingly common at northern sites. We also observed elevated flower production in northern populations and higher per capita reproductive output at the range edge. Furthermore, mean propagule mass in northern populations was 18 % larger than that from the southernmost populations. CONCLUSIONS:These findings reveal no erosion of fecundity in A. germinans populations at range limits, allowing rapid expansion of mangrove cover in the region. These results also illustrate that substantial turnover in the assemblage of flower-visiting insects can occur at an expanding range edge without altering pollen receipt.

Islands support unique plants, animals, and human societies found nowhere else on the Earth. Local and global stressors threaten the persistence of island ecosystems, with invasive species being among the most damaging, yet solvable, stressors. While the threat of invasive terrestrial mammals on island flora and fauna is well recognized, recent studies have begun to illustrate their extended and destructive impacts on adjacent marine environments. Eradication of invasive mammals and restoration of native biota are promising tools to address both island and ocean management goals. The magnitude of the marine benefits of island restoration, however, is unlikely to be consistent across the globe. We propose a list of six environmental characteristics most likely to affect the strength of land–sea linkages: precipitation, elevation, vegetation cover, soil hydrology, oceanographic productivity, and wave energy. Global databases allow for the calculation of comparable metrics describing each environmental character across islands. Such metrics can be used today to evaluate relative potential for coupled land–sea conservation efforts and, with sustained investment in monitoring on land and sea, can be used in the future to refine science-based planning tools for integrated land–sea management. As conservation practitioners work to address the effects of climate change, ocean stressors, and biodiversity crises, it is essential that we maximize returns from our management investments. Linking efforts on land, including eradication of island invasive mammals, with marine restoration and protection should offer multiplied benefits to achieve concurrent global conservation goals.

Emerald ash borer (Agrilus planipennis Fairmaire (Coleoptera: Buprestidae)), an invasive phloem-feeding beetle native to Asia, has devastated North American ash forests since its detection in Michigan, United States in 2002. As the emerald ash borer has continued to spread, the potential for successful long-term management hinges upon the release, establishment, and spread of introduced larval and egg parasitoids for biological control. Here, we focus on the establishment and evidence for spatial spread of introduced larval parasitoid, Spathius agrili Yang and Spathius galinae Belokobylskij & Strazanac (Hymenoptera: Braconidae) in the state of Maryland. To assess each species, we analyzed historical release and recovery data and resampled previous release sites and nonrelease sites for establishment. We found little evidence of establishment or spread for S. agrili, despite a comparatively large number of release locations, events, and individuals. By contrast, despite much lower propagule pressure and shorter history of releases, we detected multiple established populations of S. galinae at release sites and at sites up to 90 km from the nearest release point approximately 3 yr after its most current release. Our findings show that S. galinae has established and spread rapidly following field releases whereas its congener, S. agrili has not. Although it may still be too early to evaluate the level of population control and ash protection afforded by S. galinae, these findings indicate the need for continued investment in S. galinae for emerald ash borer classical biological control efforts.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory. A NutNet experiment in 57 grasslands across six continents shows that when herbivores are excluded from grasslands with a long coevolutionary history of grazing plant diversity is reduced, while in grasslands without a long grazing history the evolutionary history of the plant species regulates the response of plant diversity.

Photo 1. Red mangrove (Rhizophora mangle) at the dock, West island of Twin Cays, Belize. Photo credit: Alexander J. Forde. Photo 2. Views from the benthos of interior ponds: (upper panel) upside-down is sunny-side up for Cassiopea jellies (Cassiopeidae), which partner with zooxanthellae to photosynthesize; (lower panel) flocculent microbial mats rise from the bottom and accumulate as a nutrient subsidy along pond margins. Photo credits: Daniel S. Gruner (upper panel) and Alexander J. Forde (lower panel). Photo 3. Herbivores and herbivory on red mangroves (clockwise from top left): Megalopyge dyari (Megalopygidae); leaf miner Marmara sp. (Gracillariidae); mangrove tree crab (Aratus pisonii); and terminal bud damage by Ecdytolopha sp. (Tortricidae). Photo credits: Ilka C. Feller (upper panels, lower right panel) and Alexander J. Forde (lower left panel). Photo 4. The mangrove yellow warbler (Setophaga petechia) is one of several resident arthropod-gleaning bird species. Photo credit: Ilka C. Feller. Photo 5. Experimental bird exclosures on red mangroves along the margin between dwarf plants in a phosphorus-limited interior pond (foreground) and trees enriched by flocculent detritus (background). Photo credit: Daniel S. Gruner. These photographs illustrate the article “Insectivorous birds reduce herbivory but do not increase mangrove growth across productivity zones” by Alexander J. Forde, Ilka C. Feller, John D. Parker, and Daniel S. Gruner published in Ecology. https://doi.org/10.1002/ecy.3768

Top-down effects of predators and bottom-up effects of resources are important drivers of community structure and function in a wide array of ecosystems. Fertilization experiments impose variation in resource availability that can mediate the strength of predator impacts, but the prevalence of such interactions across natural productivity gradients is less clear. We studied the joint impacts of top-down and bottom-up factors in a tropical mangrove forest system, leveraging fine-grained patchiness in resource availability and primary productivity on coastal cays of Belize. We excluded birds from canopies of red mangrove (Rhizophoraceae: Rhizophora mangle) for 13 months in zones of phosphorus-limited, stunted dwarf mangroves, and in adjacent zones of vigorous mangroves that receive detrital subsidies. Birds decreased total arthropod densities by 62%, herbivore densities more than fivefold, and reduced rates of leaf and bud herbivory by 45% and 52%, respectively. Despite similar arthropod densities across both zones of productivity, leaf and bud damage were 2.0 and 4.3 times greater in productive stands. Detrital subsidies strongly impacted a suite of plant traits in productive stands, potentially making leaves more nutritious and vulnerable to damage. Despite consistently strong impacts on herbivory, we did not detect top-down forcing that impacted mangrove growth, which was similar with and without birds. Our results indicated that both top-down and bottom-up forces drive arthropod community dynamics, but attenuation at the plant-herbivore interface weakens top-down control by avian insectivores.

To evaluate how increased anthropogenic nutrient inputs alter carbon cycling in grasslands, we conducted a litter decomposition study across 20 temperate grasslands on three continents within the Nutrient Network, a globally distributed nutrient enrichment experiment We determined the effects of addition of experimental nitrogen (N), phosphorus (P) and potassium plus micronutrient (K-mu) on decomposition of a common tree leaf litter in a long-term study (maximum of 7 years; exact deployment period varied across sites). The use of higher order decomposition models allowed us to distinguish between the effects of nutrients on early- versus late-stage decomposition. Across continents, the addition of N (but not other nutrients) accelerated early-stage decomposition and slowed late-stage decomposition, increasing the slowly decomposing fraction by 28% and the overall litter mean residence time by 58%. Synthesis. Using a novel, long-term cross-site experiment, we found widespread evidence that N enhances the early stages of above-ground plant litter decomposition across diverse and widespread temperate grassland sites but slows late-stage decomposition. These findings were corroborated by fitting the data to multiple decomposition models and have implications for N effects on soil organic matter formation. For example, following N enrichment, increased microbial processing of litter substrates early in decomposition could promote the production and transfer of low molecular weight compounds to soils and potentially enhance the stabilization of mineral-associated organic matter. By contrast, by slowing late-stage decomposition, N enrichment could promote particulate organic matter (POM) accumulation. Such hypotheses deserve further testing.