Morpho-physiological traits and tissue burdens of Ecklonia radiata linked to environmental variation in an urban estuary.

Organisms respond to their environment in various ways, including moving, adapting, acclimatising or a combination of responses. Within estuarine habitats, organisms are exposed to naturally variable environmental conditions. In urbanised estuaries, these natural variations can interact with human stressors such as habitat modification and pollution. Here, we investigated trait variation in the golden kelp Ecklonia radiata across an urban estuary - Sydney Harbour, Australia. We found that kelp morphology differed significantly between the more human-modified inner and the less modified outer harbour. Kelp individuals were smaller, had fewer laminae, and lacked spines in the inner harbour where it was warmer, more contaminated and less light was available. Inner harbour populations were characterised by lower tissue nitrogen and higher lead concentrations. These findings provide insights into how environmental variation could affect kelp morphology and physiology, and the high trait variation suggests adaptive capacity in E. radiata.

Influence of habitat features on the colonisation of native and non-indigenous species.

Marine artificial structures provide substrates on which organisms can settle and grow. These structures facilitate establishment and spread of non-indigenous species, in part due to their distinct physical features (substrate material, movement, orientation) compared to natural habitat analogues such as rocky shores, and because following construction, they have abundant resources (space) for species to colonise. Despite the perceived importance of these habitat features, few studies have directly compared distributions of native and non-indigenous species or considered how functional identity and associated environmental preferences drive associations. We undertook a meta-analysis to investigate whether colonisation of native and non-indigenous species varies between artificial structures with features most closely resembling natural habitats (natural substrates, fixed structures, surfaces oriented upwards) and those least resembling natural habitats (artificial materials, floating structures, downfacing or vertical surfaces), or whether functional identity is the primary driver of differences. Analyses were done at global and more local (SE Australia) scales to investigate if patterns held regardless of scale. Our results suggest that functional group (i.e., algae, ascidians. barnacles, bryozoans, polychaetes) rather than species classification (i.e., native or non-indigenous) are the main drivers of differences in communities between different types of artificial structures. Specifically, there were differences in the abundance of ascidians, barnacles, and polychaetes between (1) upfacing and downfacing/vertical surfaces, and (2) floating and fixed substrates. When differences were detected, taxa were most abundant on features least resembling natural habitats. Results varied between global and SE Australian analyses, potentially due to reduced variability across studies in the SE Australian dataset. Thus, the functional group and associated preferences of the highest threat NIS in the area should be considered in design strategies (e.g., ecological engineering) to limit their establishment on newly built infrastructure.

Variable effects of substrate colour and microtexture on sessile marine taxa in Australian estuaries.

Concrete infrastructure in coastal waters is increasing. While adding complex habitat and manipulating concrete mixtures to enhance biodiversity have been studied, field investigations of sub-millimetre-scale complexity and substrate colour are lacking. Here, the interacting effects of 'colour' (white, grey, black) and 'microtexture' (smooth, 0.5 mm texture) on colonisation were assessed at three sites in Australia. In Townsville, no effects of colour or microtexture were observed. In Sydney, spirorbid polychaetes occupied more space on smooth than textured tiles, but there was no effect of microtexture on serpulid polychaetes, bryozoans and algae. In Melbourne, barnacles were more abundant on black than white tiles, while serpulid polychaetes showed opposite patterns and ascidians did not vary with treatments. These results suggest that microtexture and colour can facilitate colonisation of some taxa. The context-dependency of the results shows that inclusion of these factors into marine infrastructure designs needs to be carefully considered.

Complexity-functioning relationships differ across different environmental conditions.

Habitat complexity is widely considered an important determinant of biodiversity, and enhancing complexity can play a key role in restoring degraded habitats. However, the effects of habitat complexity on ecosystem functioning - as opposed to biodiversity and community structure - are relatively poorly understood for artificial habitats, which dominate many coastlines. With Greening of Grey Infrastructure (GGI) approaches, or eco-engineering, increasingly being applied around the globe, it is important to understand the effects that modifying habitat complexity has on both biodiversity and ecological functioning in these highly modified habitats. We assessed how manipulating physical (primary substrate) and/or biogenic habitat (bivalves) complexity on intertidal artificial substrata affected filtration rates, net and gross primary productivity (NPP and GPP, respectively) and community respiration (CR) - as well as abundance of filter feeders and macro-algae and habitat use by cryptobenthic fish across six locations in three continents. We manipulated both physical and biogenic complexity using 1) flat or ridged (2.5 cm or 5 cm) settlement tiles that were either 2) unseeded or seeded with oysters or mussels. Across all locations, increasing physical and biogenic complexity (5 cm seeded tiles) had a significant effect on most ecological functioning variables, increasing overall filtration rates and community respiration of the assemblages on tiles but decreasing productivity (both GPP and NPP) across all locations. There were no overall effects of increasing either type of habitat complexity on cryptobenthic fish MaxN, total time in frame or macro-algal cover. Within each location, there were marked differences in the effects of habitat complexity. In Hobart, we found higher filtration, filter feeder biomass and community respiration on 5 cm tiles compared to flat tiles. However, at this location, both macro-algae cover and GPP decreased with increasing physical complexity. Similarly in Dublin, filtration, filter feeder biomass and community respiration were higher on 5 cm tiles compared to less complex tiles. In Sydney, filtration and filter feeder biomass were higher on seeded than unseeded tiles, and fish MaxN was higher on 5 cm tiles compared to flat tiles. On unseeded tiles in Sydney, filter feeder biomass also increased with increasing physical complexity. Our findings suggest that GGI solutions via increased habitat complexity are likely to have trade-offs among potentially desired functions, such as productivity and filtration rates, and variable effects on cryptobenthic fish communities. Importantly, our results show that the effects of GGI practices can vary markedly according to the environmental context and therefore should not be blindly and uniformly applied across the globe.

Current extent and future opportunities for living shorelines in Australia.

Living shorelines aim to enhance the resilience of coastlines to hazards while simultaneously delivering co-benefits such as carbon sequestration. Despite the potential ecological and socio-economic benefits of living shorelines over conventional engineered coastal protection structures, application is limited globally. Australia has a long and diverse coastline that provides prime opportunities for living shorelines using beaches and dunes, vegetation, and biogenic reefs, which may be either natural ('soft' approach) or with an engineered structural component ('hybrid' approach). Published scientific studies, however, have indicated limited use of living shorelines for coastal protection in Australia. In response, we combined a national survey and interviews of coastal practitioners and a grey and peer-reviewed literature search to (1) identify barriers to living shoreline implementation; and (2) create a database of living shoreline projects in Australia based on sources other than scientific literature. Projects included were those that had either a primary or secondary goal of protection of coastal assets from erosion and/or flooding. We identified 138 living shoreline projects in Australia through the means sampled starting in 1970; with the number of projects increasing through time particularly since 2000. Over half of the total projects (59 %) were considered to be successful according to their initial stated objective (i.e., reducing hazard risk) and 18 % of projects could not be assessed for their success based on the information available. Seventy percent of projects received formal or informal monitoring. Even in the absence of peer-reviewed support for living shoreline construction in Australia, we discovered local and regional increases in their use. This suggests that coastal practitioners are learning on-the-ground, however more generally it was stated that few examples of living shorelines are being made available, suggesting a barrier in information sharing among agencies at a broader scale. A database of living shoreline projects can increase knowledge among practitioners globally to develop best practice that informs technical guidelines for different approaches and helps focus attention on areas for further research.

Impacts of Climate Change on Marine Foundation Species.

Marine foundation species are the biotic basis for many of the world's coastal ecosystems, providing structural habitat, food, and protection for myriad plants and animals as well as many ecosystem services. However, climate change poses a significant threat to foundation species and the ecosystems they support. We review the impacts of climate change on common marine foundation species, including corals, kelps, seagrasses, salt marsh plants, mangroves, and bivalves. It is evident that marine foundation species have already been severely impacted by several climate change drivers, often through interactive effects with other human stressors, such as pollution, overfishing, and coastal development. Despite considerable variation in geographical, environmental, and ecological contexts, direct and indirect effects of gradual warming and subsequent heatwaves have emerged as the most pervasive drivers of observed impact and potent threat across all marine foundation species, but effects from sea level rise, ocean acidification, and increased storminess are expected to increase. Documented impacts include changes in the genetic structures, physiology, abundance, and distribution of the foundation species themselves and changes to their interactions with other species, with flow-on effects to associated communities, biodiversity, and ecosystem functioning. We discuss strategies to support marine foundation species into the Anthropocene, in order to increase their resilience and ensure the persistence of the ecosystem services they provide.

Facilitation of non-indigenous ascidian by marine eco-engineering interventions at an urban site.

Marine artificial structures often support lower native species diversity and more non-indigenous species (NIS), but adding complex habitat and using bioreceptive materials have the potential to mitigate these impacts. Here, the interacting effects of structural complexity (flat, complex with pits) and concrete mixture (standard, or with oyster shell or vermiculite aggregate) on recruitment were assessed at two intertidal levels at an urban site. Complex tiles had less green algal cover, oyster shell mixtures had less brown (Ralfsia sp.) algal cover. At a low tidal elevation, the non-indigenous ascidian Styela plicata dominated complex tiles. Additionally, mixtures with oyster shell supported higher total cover of sessile species, and a higher cover of S. plicata. There were no effects of complexity or mixture on biofilm communities and native and NIS richness. Overall, these results suggest that habitat complexity and some bioreceptive materials may facilitate colonisation by a dominant invertebrate invader on artificial structures.

Oyster reef restoration fails to recoup global historic ecosystem losses despite substantial biodiversity gain.

Human activities have led to degradation of ecosystems globally. The lost ecosystem functions and services accumulate from the time of disturbance to the full recovery of the ecosystem and can be quantified as a "recovery debt," providing a valuable tool to develop better restoration practices that accelerate recovery and limit losses. Here, we quantified the recovery of faunal biodiversity and abundance toward a predisturbed state following structural restoration of oyster habitats globally. We found that while restoration initiates a rapid increase in biodiversity and abundance of reef-associated species within 2 years, recovery rate then decreases substantially, leaving a global shortfall in recovery of 35% below a predisturbed state. While efficient restoration methods boost recovery and minimize recovery shortfalls, the time to full recovery is yet to be quantified. Therefore, potential future coastal development should weigh up not only the instantaneous damage to ecosystem functions but also the potential for generational loss of services.

A function-based typology for Earth's ecosystems.

As the United Nations develops a post-2020 global biodiversity framework for the Convention on Biological Diversity, attention is focusing on how new goals and targets for ecosystem conservation might serve its vision of 'living in harmony with nature'1,2. Advancing dual imperatives to conserve biodiversity and sustain ecosystem services requires reliable and resilient generalizations and predictions about ecosystem responses to environmental change and management3. Ecosystems vary in their biota4, service provision5 and relative exposure to risks6, yet there is no globally consistent classification of ecosystems that reflects functional responses to change and management. This hampers progress on developing conservation targets and sustainability goals. Here we present the International Union for Conservation of Nature (IUCN) Global Ecosystem Typology, a conceptually robust, scalable, spatially explicit approach for generalizations and predictions about functions, biota, risks and management remedies across the entire biosphere. The outcome of a major cross-disciplinary collaboration, this novel framework places all of Earth's ecosystems into a unifying theoretical context to guide the transformation of ecosystem policy and management from global to local scales. This new information infrastructure will support knowledge transfer for ecosystem-specific management and restoration, globally standardized ecosystem risk assessments, natural capital accounting and progress on the post-2020 global biodiversity framework.

Remnant oyster reefs as fish habitat within the estuarine seascape.

Interest in oyster reef conservation and restoration is growing globally, but particularly in Australia, it is unclear the extent to which oyster reefs complement (versus replicate) habitat provisioning by other structured habitats in the seascape. Remote underwater video surveys of two east Australian estuaries revealed that at high tide, oyster reefs not only supported distinct fish communities to bare sediments but also to adjacent seagrass beds and mangrove forests. Fish observations in oyster reefs were close to double that of mangroves and seagrass, with species richness, abundance, feeding and wandering behaviours similar. Several species of blenny and goby were unique to oyster reefs and oyster-containing mangroves, whilst recreationally fished species such as bream and mullet were more abundant on oyster reefs than in other habitats. Resolving the association between oyster reefs and fish species within the broader seascape will assist in developing restoration and management strategies that maximise fisheries benefit.

Complexity-biodiversity relationships on marine urban structures: reintroducing habitat heterogeneity through eco-engineering.

Urbanization is leading to biodiversity loss through habitat homogenization. The smooth, featureless surfaces of many marine urban structures support ecological communities, often of lower biodiversity, distinct from the complex natural habitats they replace. Eco-engineering (design for ecological co-benefits) seeks to enhance biodiversity and ecological functions on urban structures. We assessed the benefits to biodiversity of retrofitting four types of complex habitat panels to an intertidal seawall at patch (versus flat control panels) and site (versus unmodified control seawalls and reference rocky shores) scales. Two years after installation, patch-scale effects of complex panels on biodiversity ranged from neutral to positive, depending on the protective features they provided, though all but one design (honeycomb) supported unique species. Water-retaining features (rockpools) and crevices, which provided moisture retention and cooling, increased biodiversity and supported algae and invertebrates otherwise absent. At the site scale, biodiversity benefits ranged from neutral at the high- and mid-intertidal to positive at the low-intertidal elevation. The results highlight the importance of matching eco-engineering interventions to the niche of target species, and environmental conditions. While species richness was greatest on rockpool and crevice panels, the unique species supported by other panel designs highlights that to maximize biodiversity, habitat heterogeneity is essential. This article is part of the theme issue 'Ecological complexity and the biosphere: the next 30 years'.

Turning a lost reef ecosystem into a national restoration program.

Achieving a sustainable socioecological future now requires large-scale environmental repair across legislative borders. Yet, enabling large-scale conservation is complicated by policy-making processes that are disconnected from socioeconomic interests, multiple sources of knowledge, and differing applications of policy. We considered how a multidisciplinary approach to marine habitat restoration generated the scientific evidence base, community support, and funding needed to begin the restoration of a forgotten, functionally extinct shellfish reef ecosystem. The key actors came together as a multidisciplinary community of researchers, conservation practitioners, recreational fisher communities, and government bodies that collaborated across sectors to rediscover Australia's lost shellfish reefs and communicate the value of its restoration. Actions undertaken to build a case for large-scale marine restoration included synthesizing current knowledge on Australian shellfish reefs and their historical decline, using this history to tell a compelling story to spark public and political interest, integrating restoration into government policy, and rallying local support through community engagement. Clearly articulating the social, economic, and environmental business case for restoration led to state and national funding for reef restoration to meet diverse sustainability goals (e.g., enhanced biodiversity and fisheries productivity) and socioeconomic goals (e.g., job creation and recreational opportunities). A key lesson learned was the importance of aligning project goals with public and industry interests so that projects could address multiple political obligations. This process culminated in Australia's largest marine restoration initiative and shows that solutions for large-scale ecosystem repair can rapidly occur when socially valued science acts on political opportunities.

Transformación de un Ecosistema Arrecifal Perdido en un Programa Nacional de Restauración Resumen Actualmente se requiere una reparación ambiental a gran escala que atraviese fronteras legislativas para lograr un futuro socio-ecológico sustentable. Aun así, habilitar la conservación a gran escala es complicado debido a los procesos de elaboración de políticas que están desconectadas de los intereses socio-económicos, las múltiples fuentes de conocimiento y las diferentes aplicaciones de las políticas. Consideramos cómo una estrategia multidisciplinaria para la restauración de hábitats marinos generó una base de evidencia científica, apoyo comunitario y el financiamiento necesario para así iniciar la restauración de un ecosistema arrecifal de conchas funcionalmente extinto. Los actores clave formaron una comunidad multidisciplinaria de investigadores, practicantes de la conservación, comunidades de pescadores recreativos y órganos gubernamentales que colaboró con varios sectores para redescubrir los arrecifes perdidos de Australia y comunicó el valor de su restauración. Las acciones realizadas para armar el caso para la restauración marina a gran escala incluyeron la síntesis del conocimiento actual sobre los arrecifes de conchas en Australia y su declinación histórica, el uso de esta historia para contar una narración convincente que active el interés público y político, la integración de la restauración a la política gubernamental y la movilización del apoyo local por medio de la participación comunitaria. Claramente, la articulación del caso del negocio social, económico y ambiental para la restauración llevó al financiamiento estatal y nacional para la restauración arrecifal a cumplir diversos objetivos socio-económicos (p. ej.: creación de empleos, oportunidades recreativas) y de restauración (p. ej.: una productividad realzada de la biodiversidad y las pesquerías). Una lección clave que aprendimos fue lo importante que es alinear los objetivos del proyecto con los intereses públicos y de la industria, de tal manera que los proyectos aborden las múltiples obligaciones políticas. Este proceso culminó con la iniciativa de restauración marina más grande en Australia y demuestra que las soluciones para la reparación de los ecosistemas a gran escala pueden ocurrir rápidamente cuando la ciencia con valor social actúa sobre las oportunidades políticas.

Heterogeneity within and among co-occurring foundation species increases biodiversity.

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.

Material type influences the abundance but not richness of colonising organisms on marine structures.

Urbanisation of coastal areas and growth in the blue economy drive the proliferation of artificial structures in marine environments. These structures support distinct ecological communities compared to natural hard substrates, potentially reflecting differences in the materials from which they are constructed. We undertook a meta-analysis of 46 studies to compare the effects of different material types (natural or eco-friendly vs. artificial) on the colonising biota on built structures. Neither the abundance nor richness of colonists displayed consistent patterns of difference between artificial and natural substrates or between eco-friendly and standard concrete. Instead, there were differences in the abundance of organisms (but not richness) between artificial and natural materials, that varied according to material type and by functional group. When compared to biogenic materials and rock, polymer and metal supported significantly lower abundances of total benthic species (in studies assessing sessile and mobile species together), sessile invertebrates and corals (in studies assessing these groups individually). In contrast, non-indigenous species were significantly more abundant on wood than metal. Concrete supported greater abundances of the general community, including habitat-forming species, compared to wood. Our results suggest that the ecological requirements of the biological community, alongside economic, logistic and engineering factors should be considered in material selection for multifunctional marine structures that deliver both engineering and ecological (enhanced abundance and diversity) benefits.

Temperature-buffering by oyster habitat provides temporal stability for rocky shore communities.

Intertidal rocky shores are considered among the most thermally stressful marine ecosystems, where many species live close to their upper thermal limit and depend on access to cool microclimates to persist through heat events. In such environments, the provision of cool microclimates by habitat-forming species enables persistence of associated species during high temperature events. We assessed whether, by maintaining cool microclimates through heat events, habitat formed by rock oysters (Saccostrea cucullata) provides temporal stability to associated invertebrate communities over periods of extreme temperatures. On three tropical rocky shores of Hong Kong, which experiences a monsoonal climate, we compared changes in microclimates and invertebrate communities associated with oyster and bare rock habitats between the cool and hot seasons. Oyster habitats were, across both seasons, consistently characterised by lower maximum temperatures and greater thermal stability than bare rock habitats. Invertebrate communities in the bare rock habitat were less diverse and abundant in the hot than the cool season, but communities in the cooler habitats provided by oysters did not display temporal change. These results demonstrate that microclimates formed by oysters provide temporal stability to associated communities across periods of temperature change and are key determinants of species distributions in thermally stressful environments. The conservation and restoration of oyster habitats may, therefore, build resilience in associated ecological communities subject to ongoing environmental change.

Comparison of wrack dynamics between mangrove forests with and without seawalls.

The supply and fate of wrack (dead organic matter) is a critical determinant of the structure and function of shoreline ecosystems, and their role as carbon repositories. The increasingly common practise of armouring urbanised shorelines with seawalls impacts wrack deposits of unvegetated estuarine and coastal shorelines by truncating the intertidal zone and/or by modifying the physical and biological processes that deliver and remove wrack. This study tested whether such effects also extend to mangrove forests. A survey of wrack deposits in mangrove forests with and without seawalls along the Parramatta River, Sydney, Australia, revealed that at sites with seawalls placed at a mid-intertidal elevation wrack deposits were shifted from the high- to mid-intertidal but were otherwise of similar cover and composition. Experiments tracking the fate of wrack determined that, as compared to the mid-intertidal zone of unarmoured shorelines, wrack deposits at sites with seawalls were more readily mobilised. This was in some instances countered by the reduction in Casuarina glauca litter on armoured shorelines, as experiments revealed that Avicennia marina leaf decomposition was slower in the absence than the presence of C. glauca. Overall, the results suggest that effects of armouring on wrack composition and dynamics may be weaker in mangrove forests than on unvegetated shorelines. This could reflect the predominantly autochthonous source of wrack in mangrove forests, the habitat structure of forests minimizing hydrodynamic impacts of seawalls, and/or the differing reasons for which hard structures are constructed in low hydrodynamic energy vegetated as opposed to high hydrodynamic energy unvegetated settings.

Backshore nourishment of a beach degraded by off-road vehicles: Ecological impacts and benefits.

Worldwide, spoil from maintenance dredging of navigation channels is increasingly used to opportunistically nourish beaches. This is often justified on the presumption that nourishment will improve public beach amenity and restore sandy beach habitat. However, this is not necessarily the case, especially for beaches that do not have an immediate threat of significant erosion. We addressed the ecological impacts and benefits of a backshore sand nourishment project conducted along an off-road vehicle (ORV) damaged section of Blacksmiths Beach, New South Wales, Australia. Sediment, sourced from dredging the inlet of nearby Lake Macquarie, was placed on the foredune, ORVs were excluded and low-density vegetation was planted. Sampling before and after the management interventions, at the Impact (nourished) site, two Control sites (with ORVs), and two Reference sites (without ORVs), assessed ecological impacts of nourishment and the efficacy of the interventions in rehabilitating vegetation and invertebrate communities degraded by ORVs. Nourishment initially had large negative impacts on vegetation cover, as well as on invertebrate abundance and richness. Recovery to a pre-nourished state was, however, observed for vegetation cover after 9 months and invertebrate communities after 21 months. Nevertheless, by the end of our study that extended 21 months post-nourishment and ORV exclusion, there was no evidence of change in the nourished site towards the state of Reference sites. Overall, our study suggests that small-scale backshore sand nourishments of ocean beaches may have only short-term negative impacts on foredune ecosystems when accompanied with some replanting. Nevertheless, where the frequency of sand disposals is greater than the required recovery time, or cumulative effects amass, longer-term or sustained impacts may occur. Our study does not support the efficacy of sand nourishment as a tool for ecological restoration, at least in the short term, without sustained replanting and weeding efforts aimed at reinstating the vegetation community.

The mechanisms by which oysters facilitate invertebrates vary across environmental gradients.

The effective use of ecosystem engineers to conserve biodiversity requires an understanding of the types of resources an engineer modifies, and how these modifications vary with biotic and abiotic context. In the intertidal zone, oysters engineer ecological communities by reducing temperature and desiccation stress, enhancing the availability of hard substrate for attachment, and by ameliorating biological interactions such as competition and predation. Using a field experiment manipulating shading, predator access and availability of shell substrate at four sites distributed over 900 km of east Australian coastline, we investigated how the relative importance of these mechanisms of facilitation vary spatially. At all sites, and irrespective of environmental conditions, the provision of hard substrate by oysters enhanced the abundance and richness of invertebrates, in particular epibionts (barnacles and oyster spat) and grazing gastropods. Mobile arthropods utilised the habitat provided by disarticulated dead oysters more than live oyster habitat, whereas the abundance of polychaetes and bivalves were much greater in live oysters, suggesting the oyster filter-feeding activity is important for these groups. In warmer estuaries, shading by oysters had a larger effect on biodiversity, whereas in cooler estuaries, the provision of a predation refuge by oysters played a more important role. Such knowledge of how ecosystem engineering effects vary across environmental gradients can help inform management strategies targeting ecosystem resilience via the amelioration of specific environmental stressors, or conservation of specific community assemblages.

Secondary foundation species enhance biodiversity.

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.