Many studies have documented habitat cascades where two co‐occurring habitat‐forming species control biodiversity. However, more than two habitat‐formers could theoretically co‐occur. We here documented a sixth‐level habitat cascade from the Avon‐Heathcote Estuary, New Zealand, by correlating counts of attached inhabitants to the size and accumulated biomass of their biogenic hosts. These data revealed predictable sequences of habitat‐formation (=attachment space). First, the bivalve Austrovenus provided habitat for green seaweeds (Ulva) that provided habitat for trochid snails in a typical estuarine habitat cascade. However, the trochids also provided habitat for the nonnative bryozoan Conopeum that provided habitat for the red seaweed Gigartina that provided habitat for more trochids, thereby resetting the sequence of the habitat cascade, theoretically in perpetuity. Austrovenus is here the basal habitat‐former that controls this “long” cascade. The strength of facilitation increased with seaweed frond size, accumulated seaweed biomass, accumulated shell biomass but less with shell size. We also found that Ulva attached to all habitat‐formers, trochids attached to Ulva and Gigartina, and Conopeum and Gigartina predominately attached to trochids. These “affinities” for different habitat‐forming species probably reflect species‐specific traits of juveniles and adults. Finally, manipulative experiments confirmed that the amount of seaweed and trochids was important and consistent regulators of the habitat cascade in different estuarine environments. We also interpreted this cascade as a habitat‐formation network that describes the likelihood of an inhabitant being found attached to a specific habitat‐former. We conclude that the strength of the cascade increased with the amount of higher‐order habitat‐formers, with differences in form and function between higher and lower‐order habitat‐formers, and with the affinity of inhabitants for higher‐order habitat‐formers. We suggest that long habitat cascades are common where species traits allow for physical attachment to other species, such as in marine benthic systems and old forest.

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.

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Recent research has shown that co-occurring primary and secondary habitat-forming species typically support higher biodiversity than do monocultures of the primary habitat-former alone. However, these ‘habitat cascades’ may not be universal and it is important to know whether, when and where positive effects on biodiversity from secondary habitat-forming species change to negative effects. Here, we tested how anthropogenic stressors (fertilisation and sedimentation) and unattached secondary habitat-forming Ulva seaweeds affected the primary habitat-forming seagrass, Zostera muelleri, and its associated invertebrates in the Avon–Heathcote Estuary, New Zealand. We experimentally stressed Zostera by adding different fertilisation and sediment levels. Fertilisation had little impact, whereas even low sedimentation levels had strong negative effects on Zostera and its associated fauna. In a second experiment, sediments and Ulva were added to seagrass beds and unvegetated mudflats to test whether sediment stress modifies habitat cascades. We found again strong negative effects of sediments on Zostera, irrespective of spatio-temporal conditions, and that negative effects of sediments on invertebrates were enhanced in the presence of the secondary habitat former. These results highlighted that anthropogenic stressors can destabilise habitat cascades; processes that may be of particular importance in estuaries that are characterised by low biodiversity and stressful environmental conditions.

Oceanography and Marine Research J o u rn al of Oc e a n o g ra phy an d M a ri ne Res e a r c h