Unveiling the transition from niche to dispersal assembly in ecology.

A central goal in ecology is to understand what maintains species diversity in local communities. Classic ecological theory1,2 posits that niches dictate the maximum number of species that can coexist in a community and that the richness of observed species will be below this maximum only where immigration is very low. A new alternative theory3,4 is that niches, instead, dictate the minimum number of coexisting species and that the richness of observed species will usually be well above this because of ongoing immigration. We conducted an experimental test to discriminate between these two unified theories using a manipulative field experiment with tropical intertidal communities. We found, consistent with the new theory, that the relationship of species richness to immigration rate stabilized at a low value at low immigration rates and did not saturate at high immigration rates. Our results suggest that tropical intertidal communities have low niche diversity and are typically in a dispersal-assembled regime where immigration is high enough to overfill the niches. Observational data from other studies3,5 suggest that these conclusions may generalize to other ecological systems. Our new experimental approach can be adapted for other systems and be used as a 'niche detector' and a tool for assessing when communities are niche versus dispersal assembled.

Clarifications on habitat complexity: A response to technical note by Madin et al.

In a critique of our recent review on measuring habitat complexity in ecology, Madin et al. (2023) advocate the use of fractal dimension in ecology and defend their geometric constraint theory of habitat complexity. We explain the flaws in their arguments and highlight points where they misinterpreted our statements.

Measuring habitat complexity and spatial heterogeneity in ecology.

Habitat complexity has been considered a key driver of biodiversity and other ecological phenomena for nearly a century. However, there is still no consensus over the definition of complexity or how to measure it. Up-to-date and clear guidance on measuring complexity is urgently needed, particularly given the rise of remote sensing and advent of technologies that allow environments to be scanned at unprecedented spatial extents and resolutions. Here we review how complexity is measured in ecology. We provide a framework for metrics of habitat complexity, and for the related concept of spatial heterogeneity. We focus on the two most commonly used complexity metrics in ecology: fractal dimension and rugosity. We discuss the pros and cons of these metrics using practical examples from our own empirical data and from simulations. Fractal dimension is particularly widely used, and we provide a critical examination of it drawing on research from other scientific fields. We also discuss informational metrics of complexity and their potential benefits. We chart a path forward for research on measuring habitat complexity by presenting, as a guide, sets of essential and desirable criteria that a metric of complexity should possess. Lastly, we discuss the applied significance of our review.

Behaviour broadens thermal safety margins on artificial coastal defences in the tropics.

Tropical species are predicted to be among the most vulnerable to climate change as they often live close to their upper limits to thermal tolerance and in many cases, behavioural thermoregulation is required to persist in the thermal extremes of tropical latitudes. In concert with warming temperatures, near-shore species are faced with the additional threat of shoreline hardening, leading to a reduction in microhabitats that can provide thermal refuges. This situation is exemplified in Singapore, which lies almost on the equator and so experiences year-round hot temperatures, and much of its coastline is now seawall. To investigate the thermal ecology of a common intertidal gastropod, Nerita undata, on these artificial structures, we measured thermal conditions on two seawalls, the temperatures of habitats occupied by the snail, and compared these with the snail's thermal tolerance by measuring heart rate and behavioural thermoregulation (as preferred temperature, Tpref). At one of the two seawalls (Tanjong Rimau), temperatures experienced by N. undata exceeded all measures of thermal tolerance in the sun, while at the other (Palawan Beach), they did not. Temperatures in habitats occupied by the snails on the seawalls were similar to their measured Tpref in the laboratory and were lower than all measures of thermal tolerance. Behavioural thermoregulation by the snails, therefore, significantly increased the thermal safety margins of N. undata on the relatively homogenous seawalls in Singapore, and at one of the two seawalls were necessary to allow snails to survive. Accordingly, to facilitate motile species to maintain broad thermal safety margins through behavioural regulation, the provision of additional refuges from thermal stress is recommended on artificial coastal defences such as seawalls.

Human-engineered hydrodynamic regimes as a driver of cryptic microinvertebrate assemblages on urban artificial shorelines.

Urban shorelines undergo substantial hydrodynamic changes as a result of coastal engineering and shoreline armouring that can alter sedimentation, turbidity, and other factors. These changes often coincide with major shifts in the composition and distribution of marine biota, however, rarely are hydrodynamic-mediated factors confirmed experimentally as the mechanism underpinning these shifts. This study first characterized hydrodynamic-related distribution patterns among epilithic and epiphytic microinvertebrates on urban seawalls in Singapore. We found reduced microinvertebrate abundances and distinct microinvertebrate community structure within benthic turf algae in areas where coastal defences had reduced wave energy and increased sediment deposition, among other hydrodynamic-related abiotic changes. Low-exposure areas also had reduced densities of macroinvertebrate grazers and less dense turf algae (lower mass per cm2) than adjacent high-exposure areas. Using harpacticoid copepods as a model taxon, we performed a reciprocal transplant experiment to discern between the effects of exposure-related conditions and grazing. Results from the experiment indicate that conditions associated with restricted wave energy from shoreline engineering limit harpacticoid population densities, as transplantation to low-exposure areas led to rapid reductions in abundance. At the same time, we found no effect from grazer exclusion cages, suggesting harpacticoids are minimally impacted by exposure-related gradients in gastropod macrograzer densities over short time scales. Given the key role of intertidal microinvertebrates, particularly harpacticoids, in nearshore food webs, we postulate that human-engineered hydrodynamic regimes are an important factor shaping marine ecosystem functioning in urban areas.

Effects of habitat area and spatial configuration on biodiversity in an experimental intertidal community.

Isolating the effects of fragmentation per se (i.e., spatial configuration of habitat patches) on species richness is an ongoing challenge as habitat configuration often covaries with the amount of habitat. Consequently, there is a lack of experimental evidence for configurational effects on species richness in the whole landscape. Here, we developed a novel experimental system for testing the independent and interactive effects of habitat area and configuration on tropical intertidal species richness. Our results confirmed the expectation that average species richness would increase monotonically with habitat area. More intriguingly, we found mixed evidence for a non-monotonic relationship between species richness and fragmentation per se, with the highest richness at intermediate fragmentation configuration, that is, when habitat tiles were placed in a "several-small" configuration. The effect of habitat configuration was not due to passive sampling (since area was controlled for), variation in total individual abundance, or niche specialization of species to different landscape configurations. We postulate that a combination of processes, including local negative density dependence and dispersal limitation, could give rise to the observed pattern. We emphasize the importance of considering configurational effects on biodiversity at broader spatial scales and for more experimental research to delve into the mechanisms driving the patterns seen here.

Urban coral reefs: Degradation and resilience of hard coral assemblages in coastal cities of East and Southeast Asia.

Given predicted increases in urbanization in tropical and subtropical regions, understanding the processes shaping urban coral reefs may be essential for anticipating future conservation challenges. We used a case study approach to identify unifying patterns of urban coral reefs and clarify the effects of urbanization on hard coral assemblages. Data were compiled from 11 cities throughout East and Southeast Asia, with particular focus on Singapore, Jakarta, Hong Kong, and Naha (Okinawa). Our review highlights several key characteristics of urban coral reefs, including "reef compression" (a decline in bathymetric range with increasing turbidity and decreasing water clarity over time and relative to shore), dominance by domed coral growth forms and low reef complexity, variable city-specific inshore-offshore gradients, early declines in coral cover with recent fluctuating periods of acute impacts and rapid recovery, and colonization of urban infrastructure by hard corals. We present hypotheses for urban reef community dynamics and discuss potential of ecological engineering for corals in urban areas.

Structural complexity and component type increase intertidal biodiversity independently of area.

Complexity is well accepted as one of the primary drivers of biodiversity, however, empirical support for such positive associations is often confounded with surface area and undermined by other potential explanatory factors, especially the type of structural component (e.g., pits, crevices, overhangs, etc.). In the present study, sample units (artificial substrates) of equal surface area (± 0.2%) were used to simultaneously examine the independent effects of complexity and different structural component types on species richness (S), abundance (N), and community composition. We created simple and complex concrete substrates of four different geometric designs using novel software. The substrates (n = 8) were mounted onto granite seawalls (at two tidal heights) on two islands south of Singapore Island. After 13 months of colonization, all 384 tiles were collected and their assemblages compared. A total of 53 744 individuals of 70 species/morphospecies were collected and identified. Our results show that greater complexity can support greater species richness and different communities that are independent of surface area. Furthermore, the type of structure (e.g., "pits," "grooves," "towers") can have an effect on richness and community composition that is independent of complexity.

Complexity for artificial substrates (CASU): software for creating and visualising habitat complexity.

Physical habitat complexity regulates the structure and function of biological communities, although the mechanisms underlying this relationship remain unclear. Urbanisation, pollution, unsustainable resource exploitation and climate change have resulted in the widespread simplification (and loss) of habitats worldwide. One way to restore physical complexity to anthropogenically simplified habitats is through the use of artificial substrates, which also offer excellent opportunities to explore the effects of different components (variables) of complexity on biodiversity and community structure that would be difficult to separate in natural systems. Here, we describe a software program (CASU) that enables users to visualise static, physical complexity. CASU also provides output files that can be used to create artificial substrates for experimental and/or restoration studies. It has two different operational modes: simple and advanced. In simple mode, users can adjust the five main variables of informational complexity (i.e. the number of object types, relative abundance of object types, density of objects, variability and range in the objects' dimensions, and their spatial arrangement) and visualise the changes as they do so. The advanced mode allows users to design artificial substrates by fine-tuning the complexity variables as well as alter object-specific parameters. We illustrate how CASU can be used to create tiles of different designs for application in a marine environment. Such an ability to systematically influence physical complexity could greatly facilitate ecological restoration by allowing conservationists to rebuild complexity in degraded and simplified habitats.