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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.

Climate change is leading to novel species interactions and profoundly altering ecosystems. In marine systems, tropical and subtropical species are increasing in higher latitudes. This has been linked to the deforestation of temperate coastlines, as direct effects of ocean warming combine with increased herbivory from tropical and sub-tropical fishes and lead to the decline of canopy-forming kelp. Here, we tested the hypothesis that this deforestation may be facilitated by greater palatability of temperate kelp and other canopy seaweeds compared to tropical taxa. We used multiple-choice filmed feeding field experiments and chemical analyses to measure the palatability of temperate and tropical seaweeds from Tosa Bay (southeastern Japan) and we used single-species feeding assays to measure changes in consumption of the kelp Ecklonia cava throughout the year. We found no evidence that temperate seaweeds are more palatable to herbivorous fish. In the multiple-choice assays, consumption was concentrated on both tropical and temperate Sargassum species, which are ephemeral and peak in abundance in the spring/early summer. Consumption of the kelp Ecklonia cava peaked during the autumn, when Sargassum species are absent. The highest levels of kelp herbivory coincide with the reproductive season for E. cava and may contribute to the long-term decline of these kelp forests in southern Japan.

Communities structured via facilitation cascades whereby one habitat-forming species promotes a secondary habitat-forming species with synergistic effects on biodiversity are increasingly documented. Habitat-formers can also extend the realized niche of other habitat-forming species by facilitating their recruitment into novel environments across heterogeneous landscapes. However, further understanding of whether secondary habitat-formers in these novel environments differ in their structure or support different communities from those settled in their natal habitat is required. Here, we investigated whether abiotic and biotic conditions in a novel connected habitat influence biodiversity within a secondary habitat-former facilitated outside its natal habitat. We contrasted the morphology of the habitat-forming alga, Sirophysalis trinodis, and its associated fish and epifaunal community occurring on rocky reef (natal habitat) and on nearby a biogenic hard substrate, live clam shells (novel habitat) in soft sediments. The algae on clams had a different morphology to those on the rocky shore and supported increased abundance of some epifaunal species. A reciprocal transplant experiment of S. trinodis individuals among habitats found that differences in the abundance of epifauna were explained by the increased abundance and consumption rates of predatory fish on the reef habitat compared to soft-sediment habitats, and not by differences in algal morphology between habitats. We demonstrate that facilitation promotes a secondary habitat-former into a novel habitat, which then enhances the abundance of associated epifauna by providing a predation refuge. This study contributes to a small but growing body of research demonstrating landscape-scale effects of facilitation cascades, and that basal habitat-formers can extend the realized niche of secondary habitat-formers and their associated communities.

Coastal urbanization has led to large-scale transformation of estuaries, with artificial structures now commonplace. Boat moorings are known to reduce seagrass cover, but little is known about their effect on fish communities. We used underwater video to quantify abundance, diversity, composition and feeding behaviour of fish assemblages on two scales: with increasing distance from moorings on fine scales, and among locations where moorings were present or absent. Fish were less abundant in close proximity to boat moorings, and the species composition varied on fine scales, leading to lower predation pressure near moorings. There was no relationship at the location with seagrass. On larger scales, we detected no differences in abundance or community composition among locations where moorings were present or absent. These findings show a clear impact of moorings on fish and highlight the importance of fine-scale assessments over location-scale comparisons in the detection of the effects of artificial structures.

The impacts of novel habitat-forming organisms on associated fauna have been difficult to predict, and may affect the fauna of neighbouring habitats due to changes in the spatial configuration of habitat patches of differing quality. Here, we test whether the localised expansion of a native habitat-forming macroalga, Caulerpa filiformis, on subtidal reefs can affect the abundance of fauna associated with a neighbouring macroalgal habitat. C.filiformis was a functionally distinct habitat for fauna, and the total abundance of epifauna associated with the resident alga, Sargassum linearifolium, was reduced at some sites when in close proximity to or surrounded by C.filiformis. Experimental manipulation of habitat configuration demonstrated that the low abundance of gastropods on S.linearifolium when surrounded by C.filiformis was likely explained by C.filiformis acting as a physical dispersal barrier for mobile fauna. Changes to the spatial configuration of novel and resident habitats can thus affect the abundance of fauna in addition to the direct replacement of habitats by species undergoing range expansions or increasing in abundance.