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Foundation species facilitate communities, modulate energy flow, and define ecosystems, but their ecological roles after death are frequently overlooked. Here, we reveal the widespread importance of their dead structures as unique, interacting components of ecosystems that are vulnerable to global change. Key metabolic activity, mobility, and morphology traits of foundation species either change or persist after death with important consequences for ecosystem functions, biodiversity, and subsidy dynamics. Dead foundation species frequently mediate ecosystem stability, resilience, and transitions, often through feedbacks, and harnessing their structural and trophic roles can improve restoration outcomes. Enhanced recognition of dead foundation species and their incorporation into habitat monitoring, ecological theory, and ecosystem forecasting can help solve the escalating conservation challenges of the Anthropocene., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Habitat heterogeneity is considered a primary causal driver underpinning patterns of diversity, yet the universal role of heterogeneity in structuring biodiversity is unclear due to a lack of coordinated experiments testing its effects across geographic scales and habitat types. Furthermore, key species interactions that can enhance heterogeneity, such as facilitation cascades of foundation species, have been largely overlooked in general biodiversity models. Here, we performed 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which variation in biodiversity is explained by three axes of habitat heterogeneity: the amount of habitat, its morphological complexity, and capacity to provide ecological resources (e.g. food) within and between co-occurring foundation species. We show that positive and additive effects across the three axes of heterogeneity are common, providing a compelling mechanistic insight into the universal importance of habitat heterogeneity in promoting biodiversity via cascades of facilitative interactions. Because many aspects of habitat heterogeneity can be controlled through restoration and management interventions, our findings are directly relevant to biodiversity conservation., (© 2022. The Author(s).)

Foundation species enhance biodiversity and multifunctionality across many systems; however, whether foundation species patch configuration mediates their ecological effects is unknown. In a 6-month field experiment, we test which attributes of foundation species patch configuration - i.e. patch size, total patch area, perimeter, area-perimeter ratio, or connectivity - control biodiversity, stability and multifunctionality by adding a standardised density of mussel foundation species in patches of 1, 5, 10, 30, 60, 90 or 180 individuals to a southeastern US salt marsh. Over 67% of response variables increased with clustering of mussels, responses that were driven by increases in area-perimeter ratio (33%), decreases in perimeter (29%), or increases in patch size (5%), suggesting sensitivity to external stressors and/or dependence on foundation species-derived niche availability and segregation. Thus, mussel configuration - by controlling the relative distribution of multidimensional patch interior and edge niche space - critically modulates this foundation species' effects on ecosystem structure, stability and function., (© 2018 The Authors Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Food webs are an integral part of every ecosystem on the planet, yet understanding the mechanisms shaping these complex networks remains a major challenge. Recently, several studies suggested that non-trophic species interactions such as habitat modification and mutualisms can be important determinants of food web structure. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups. Here, we combine analyses of 58 food webs from seven terrestrial, freshwater and coastal systems to test (1) the general hypothesis that non-trophic facilitation by habitat-forming foundation species enhances food web complexity, and (2) whether these enhancements have either random or targeted effects on particular trophic levels, functional groups, and linkages throughout the food web. Our empirical results demonstrate that foundation species consistently enhance food web complexity in all seven ecosystems. Further analyses reveal that 15 out of 19 food web properties can be well-approximated by assuming that foundation species randomly facilitate species throughout the trophic network. However, basal species are less strongly, and carnivores are more strongly facilitated in foundation species' food webs than predicted based on random facilitation, resulting in a higher mean trophic level and a longer average chain length. Overall, we conclude that foundation species strongly enhance food web complexity through non-trophic facilitation of species across the entire trophic network. We therefore suggest that the structure and stability of food webs often depends critically on non-trophic facilitation by foundation species., Competing Interests: The authors have declared that no competing interests exist.

It has long been recognized that primary foundation species (FS), such as trees and seagrasses, enhance biodiversity. Among the species facilitated are secondary FS, including mistletoes and epiphytes. Case studies have demonstrated that secondary FS can further modify habitat-associated organisms ('inhabitants'), but their net effects remain unknown. Here we assess how inhabitants, globally, are affected by secondary FS. We extracted and calculated 2,187 abundance and 397 richness Hedges' g effect sizes from 91 and 50 publications, respectively. A weighted meta-analysis revealed that secondary FS significantly enhanced the abundance and richness of inhabitants compared to the primary FS alone. This indirect facilitation arising through sequential habitat formation was consistent across environmental and experimental conditions. Complementary unweighted analyses on log response ratios revealed that the magnitude of these effects was similar to the global average strength of direct facilitation from primary foundation species and greater than the average strength of trophic cascades, a widely recognized type of indirect facilitation arising through sequential consumption. The finding that secondary FS enhance the abundance and richness of inhabitants has important implications for understanding the mechanisms that regulate biodiversity. Integrating secondary FS into conservation practice will improve our ability to protect biodiversity and ecosystem function.

Although there is mounting evidence that biodiversity is an important and widespread driver of ecosystem multifunctionality, much of this research has focused on small-scale biodiversity manipulations. Hence, which mechanisms maintain patches of enhanced biodiversity in natural systems and if these patches elevate ecosystem multifunctionality at both local and landscape scales remain outstanding questions. In a 17 month experiment conducted within southeastern United States salt marshes, we found that patches of enhanced biodiversity and multifunctionality arise only where habitat-forming foundation species overlap--i.e. where aggregations of ribbed mussels (Geukensia demissa) form around cordgrass (Spartina alterniflora) stems. By empirically scaling up our experimental results to the marsh platform at 12 sites, we further show that mussels--despite covering only approximately 1% of the marsh surface--strongly enhance five distinct ecosystem functions, including decomposition, primary production and water infiltration rate, at the landscape scale. Thus, mussels create conditions that support the co-occurrence of high densities of functionally distinct organisms within cordgrass and, in doing so, elevate salt marsh multifunctionality from the patch to landscape scale. Collectively, these findings suggest that patterns in foundation species' overlap drive variation in biodiversity and ecosystem functioning within and across natural ecosystems.We therefore argue that foundation species should be integrated in our conceptual understanding of forces that moderate biodiversity--ecosystem functioning relationships, approaches for conserving species diversity and strategies to improve the multifunctionality of degraded ecosystems.

Facilitation cascades arise where primary foundation species facilitate secondary (dependent) foundation species, and collectively, they increase habitat complexity and quality to enhance biodiversity. Whether such phenomena occur in nonmarine systems and if secondary foundation species enhance food web structure (e.g., support novel feeding guilds) and ecosystem function (e.g., provide nursery for juveniles) remain unclear. Here we report on field experiments designed to test whether trees improve epiphyte survival and epiphytes secondarily increase the number and diversity of adult and juvenile invertebrates in a potential live oak-Tillandsia usneoides (Spanish moss) facilitation cascade. Our results reveal that trees reduce physical stress to facilitate Tillandsia, which, in turn, reduces desiccation and predation stress to facilitate invertebrates. In experimental removals, invertebrate total density, juvenile density, species richness and H' diversity were 16, 60, 1.7, and 1.5 times higher, and feeding guild richness and H' were 5 and 11 times greater in Tillandsia-colonized relative to Tillandsia-removal limb plots. Tillandsia enhanced communities similarly in a survey across the southeastern United States. These findings reveal that a facilitation cascade organizes this widespread terrestrial assemblage and expand the role of secondary foundation species as drivers of trophic structure and ecosystem function. We conceptualize the relationship between foundation species' structural attributes and associated species abundance and composition in a Foundation Species-Biodiversity (FSB) model. Importantly, the FSB predicts that, where secondary foundation species form expansive and functionally distinct structures that increase habitat availability and complexity within primary foundation species, they generate and maintain hot spots of biodiversity and trophic interactions.

Aims: Latitudinal gradients in plant communities are well studied, yet how these fundamental ecological patterns influence ecosystem recovery after extreme weather events remains largely unknown. In coastal foredunes, we investigated how the cover of a key dune‐building grass (Uniola paniculata), vegetation diversity and vegetation cover vary along a short latitudinal gradient during recovery from hurricane disturbance. Location: Southeastern USA. Methods: We surveyed 24 sites, from central Florida to north Georgia (>400 km), four times over 18 months. General linear mixed‐effect models were used to unravel patterns of vegetation responses across latitude. Results: Vegetation properties showed countervailing patterns across the latitudinal gradient. While vegetation richness, functional diversity and total cover generally declined, Uniola cover increased with increasing latitude. Further, the latitude–richness relationship strengthened while the latitude–functional diversity relationship was invariant with increasing time since the hurricane disturbance. Meanwhile, the latitude–Uniola association was seasonally dependent and strongest in the summer. Latitude also influenced diversity–cover relationships: vegetation cover was positively related to species richness at lower latitudes, while it was positively associated with functional diversity only at northern sites. We found no relationship between species richness or functional diversity and increases in cover between time steps; however, recruitment of new species and functional groups was associated with increases in vegetation cover between time steps at northern sites. Conclusions: Our study highlights the temporal dynamism and contrasting patterns along latitudinal gradients exhibited by key engineering species and overall plant diversity in foredunes — a crucial line of coastal protection — exposed to hurricane disturbances. These results suggest a need for greater integration of latitudinal and diversity effects into our understanding of coastal dune resilience. They also highlight the potential benefits of enhancing dune plant biodiversity, particularly in areas where the dune‐building grasses that are classically employed in restoration (e.g., Uniola) are unfavoured, to accelerate the re‐establishment of well‐vegetated dunes. [ABSTRACT FROM AUTHOR]

Many wetlands around the world that occur at the base of watersheds are under threat from land‐use change, hydrological alteration, nutrient pollution, and invasive species. A relevant measure of whether the ecological character of these ecosystems has changed is the species diversity of wetland‐dependent waterbirds, especially those of conservation value. Here, we evaluate the potential mechanisms controlling variability over time and space in avian species diversity of the wetlands in the Palo Verde National Park, a Ramsar Site of international importance in Costa Rica. To do so, we assessed the relative importance of several key wetland condition metrics (i.e., surface water depth, wetland extent, and vegetation greenness), and temporal fluctuations in these metrics, in predicting the abundance of five waterbirds of high conservation value as well as overall waterbird diversity over a 9‐yr period. Generalized additive models revealed that mean NDVI, an indicator of vegetation greenness, combined with a metric used to evaluate temporal fluctuations in the wetland extent best predicted four of the five waterbird species of high conservation value as well as overall waterbird species richness and diversity. Black‐bellied Whistling‐ducks, which account for over one‐half of all waterbird individuals, and all waterbird species together were better predicted by including surface water depth along with wetland extent and its fluctuations. Our calibrated species distribution model confidently quantified monthly averages of the predicted total waterbird abundances in seven of the 10 sub‐wetlands making up the Ramsar Site and confirmed that the biophysical diversity of this entire wetland system is important to supporting waterbird populations both as a seasonal refuge and more permanently. This work further suggests that optimizing the timing and location of ongoing efforts to reduce invasive vegetation cover may be key to avian conservation by increasing waterbird habitat. [ABSTRACT FROM AUTHOR]

Keystone species have large ecological effects relative to their abundance and have been identified in many ecosystems. However, global change is pervasively altering environmental conditions, potentially elevating new species to keystone roles. Here, we reveal that a historically innocuous grazer-the marsh crab Sesarma reticulatum-is rapidly reshaping the geomorphic evolution and ecological organization of southeastern US salt marshes now burdened by rising sea levels. Our analyses indicate that sea-level rise in recent decades has widely outpaced marsh vertical accretion, increasing tidal submergence of marsh surfaces, particularly where creeks exhibit morphologies that are unable to efficiently drain adjacent marsh platforms. In these increasingly submerged areas, cordgrass decreases belowground root:rhizome ratios, causing substrate hardness to decrease to within the optimal range for Sesarma burrowing. Together, these bio-physical changes provoke Sesarma to aggregate in high-density grazing and burrowing fronts at the heads of tidal creeks (hereafter, creekheads). Aerial-image analyses reveal that resulting "Sesarma-grazed" creekheads increased in prevalence from 10 ± 2% to 29 ± 5% over the past <25 y and, by tripling creek-incision rates relative to nongrazed creekheads, have increased marsh-landscape drainage density by 8 to 35% across the region. Field experiments further demonstrate that Sesarma-grazed creekheads, through their removal of vegetation that otherwise obstructs predator access, enhance the vulnerability of macrobenthic invertebrates to predation and strongly reduce secondary production across adjacent marsh platforms. Thus, sea-level rise is creating conditions within which Sesarma functions as a keystone species that is driving dynamic, landscape-scale changes in saltmarsh geomorphic evolution, spatial organization, and species interactions. [ABSTRACT FROM AUTHOR]

Coastal ecosystem restoration is accelerating globally as a means of enhancing shoreline protection, carbon storage, water quality, fisheries, and biodiversity. Among the most substantial of these efforts have been those focused on re‐establishing oyster reefs across the US Atlantic and Gulf coasts. Despite considerable investment, it is unclear how the scale of and approaches toward oyster restoration have evolved. A synthesis of 1768 projects undertaken since 1964 reveals that oyster substrate restoration efforts have primarily been concentrated in the Chesapeake Bay and the Gulf Coast, have been heavily reliant on oyster shell, and have re‐established 4.5% of the reef area that has been lost across all regions. By comparing costs to ecosystem service benefits, we discovered that the return‐on‐investment of oyster restoration varies widely, but generally increases with project size. To facilitate the recovery of coastal ecosystems and their services, scientists and resource managers must adopt a new restoration paradigm prioritizing investment in sites that maximize economic and ecological benefits and minimize construction costs. [ABSTRACT FROM AUTHOR]

Explaining variability in the strength and sign of trophic interactions between primary consumers and plants is a long-standing research challenge. Consumer density and body size vary widely in space and time and are predicted to have interactive effects on consumer-plant interactions. In a southern US salt marsh, we used replicate field enclosures to orthogonally manipulate the body size (mass) and density of a dominant consumer (a snail). We investigated impacts (leaf damage and biomass) on monocultures of cordgrass, the foundation species, over three months. Increasing consumer density and body size increased leaf damage additively and, as predicted, multiplicatively reduced plant biomass. Notably, size and density determined the sign of consumer impact on plants: low to medium densities of small consumers enhanced, while high densities of large consumers strongly suppressed, plant biomass. Finally, total consumer metabolic biomass (mass0.75) within an enclosure parsimoniously explained plant biomass response, supporting theoretical predictions and suggesting that multiplicative effects of density and body size resulted from their effects on total metabolic biomass. The consequences of changes in consumer density and body size resulting from anthropogenic perturbations may therefore be predicted based on metabolic biomass. [ABSTRACT FROM AUTHOR]

Although there is mounting evidence that biodiversity is an important and widespread driver of ecosystem multifunctionality, much of this research has focused on small-scale biodiversity manipulations. Hence, which mechanisms maintain patches of enhanced biodiversity in natural systems and if these patches elevate ecosystem multifunctionality at both local and landscape scales remain outstanding questions. In a 17 month experiment conducted within southeastern United States salt marshes, we found that patches of enhanced biodiversity and multifunctionality arise only where habitat-forming foundation species overlap—i.e. where aggregations of ribbed mussels (Geukensia demissa) form around cordgrass (Spartina alterniflora) stems. By empirically scaling up our experimental results to the marsh platform at 12 sites, we further show that mussels—despite covering only approximately 1% of the marsh surface—strongly enhance five distinct ecosystem functions, including decomposition, primary production and water infiltration rate, at the landscape scale. Thus, mussels create conditions that support the co-occurrence of high densities of functionally distinct organisms within cordgrass and, in doing so, elevate salt marsh multifunctionality from the patch to landscape scale. Collectively, these findings suggest that patterns in foundation species' overlap drive variation in biodiversity and ecosystem functioning within and across natural ecosystems. We therefore argue that foundation species should be integrated in our conceptual understanding of forces that moderate biodiversity–ecosystem functioning relationships, approaches for conserving species diversity and strategies to improve the multifunctionality of degraded ecosystems. [ABSTRACT FROM AUTHOR]