Topographic heterogeneity triggers complementary cascades that enhance ecosystem multifunctionality.

Topographic heterogeneity sets the stage for community assembly, but its effects on ecosystem functioning remain poorly understood. Here, we test the hypothesis that topographic heterogeneity underpins multiple cascading species interactions and functional pathways that indirectly control multifunctionality. To do so, we combined experimental manipulation of a form of topographic heterogeneity on rocky shores (holes of various sizes) with a comprehensive assessment of naturally assembled communities and multifunctionality. Structural equation modeling indicated that heterogeneity: (1) enhanced biodiversity by supporting filter feeder richness; (2) triggered a facilitation cascade via reef-forming (polychaete) and biomass-dominant (macroalga) foundation species, which in turn broadly supported functionally diverse epibiotic and understory assemblages; and (3) inhibited a key consumer (limpet). The model supported that these mechanisms exerted complementary positive effects on individual functions (e.g., water filtration, ecosystem metabolism, nutrient uptake) and, in turn, collectively enhanced multifunctionality. Topographic heterogeneity may therefore serve as a cornerstone physical attribute by initiating multiple cascades that propagate through ecological communities via foundation species, ultimately manifesting disproportionate effects on ecosystem multifunctionality.

Establishing complexity targets to enhance artificial reef designs.

Artificial reefs (AR), which are integral tools for fish management, ecological reconciliation and restoration efforts, require non-polluting materials and intricate designs that mimic natural habitats. Despite their three-dimensional complexity, current designs nowadays rely on empirical methods that lack standardised pre-immersion assessment. To improve ecosystem integration, we propose to evaluate 3-dimensional Computer-aided Design (3D CAD) models using a method inspired by functional ecology principles. Based on existing metrics, we assess geometric (C-convexity, P-packing, D-fractal dimension) and informational complexity (R-specific richness, H- diversity, J-evenness). Applying these metrics to different reefs constructed for habitat protection, biomass production and bio-mimicry purposes, we identify potential complexity target points (CTPs). This method provides a framework for improving the effectiveness of artificial reef design by allowing for the adjustment of structural properties. These CTPs represent the first step in enhancing AR designs. We can refine them by evaluating complexity metrics derived from 3D reconstructions of natural habitats to advance bio-mimicry efforts. In situ, post-immersion studies can help make the CTPs more specific for certain species of interest by exploring complexity-diversity or complexity-species distribution relationships at the artificial reef scale.

Top-down effects of intraspeciflic predator behavioral variation.

Among-individual variation in predator traits is ubiquitous in nature. However, variation among populations in this trait variation has been seldom considered in trophic dynamics. This has left unexplored (a) to what degree does among-individual variation in predator traits regulate prey populations and (b) to what degree do these effects vary spatially. We address these questions by examining how predator among-individual variation in functional traits shapes communities across habitats of varying structural complexity, in field conditions. We manipulated Chinese mantis (Tenodera sinensis) density (six or twelve individuals) and behavioral trait variability (activity level by movement on an open field) in experimental patches of old fields with varying habitat complexity (density of plant material). Then, we quantified their impacts on lower trophic levels, specifically prey (arthropods > 4 mm) and plant biomass. Predator behavioral variability only altered prey biomass in structurally complex plots, and this effect depended on mantis density. In the plots with the highest habitat complexity and mantis density, behaviorally variable groups decreased prey biomass by 40.3%. In complex plots with low mantis densities, low levels of behavioral variability decreased prey biomass by 32.2%. Behavioral variability and low habitat complexity also changed prey community composition, namely by increasing ant biomass by 881%. Our results demonstrate that among-individual trait variation can shape species-rich prey communities. Moreover, these effects depend on both predator density and habitat complexity. Incorporating this important facet of ecological diversity revealed normally unnoticed effects of functional traits on the structure and function of food webs.

Molecular plasticity to ocean warming and habitat loss in a coral reef fish.

Sea surface temperatures are rising at unprecedented rates, leading to a progressive degradation of complex habitats formed by coral reefs. In parallel, acute thermal stress can lead to physiological challenges for ectotherms that inhabit coral reefs, including fishes. Warming and habitat simplification could push marine fishes beyond their physiological limits in the near future. Specifically, questions remain on how warming and habitat structure influence the brain of marine fishes. Here we evaluated how thermal stress and habitat loss are acting independently and synergistically as stressors in a damselfish of the Western Atlantic, Abudefduf saxatilis. For this experiment, 40 individuals were exposed to different combinations of temperature (27°C or 31°C) and habitat complexity (complex vs simple) for 10 days, and changes in brain gene expression and oxidative stress of liver and muscle were evaluated. The results indicate that warming resulted in increased oxidative damage in the liver (p=0.007) and changes in gene expression of the brain including genes associated with neurotransmission, immune function, and tissue repair. Individuals from simplified habitats showed higher numbers of differentially expressed genes, and changes for genes associated with synaptic plasticity and spatial memory. In addition, a reference transcriptome of A. saxatilis is presented here for the first time, serving as a resource for future molecular studies. This project enhances our understanding of how fishes are responding to the combination of coral reef degradation and thermal stress, while elucidating the plastic mechanisms that will enable generalists to persist in a changing world.

Habitat complexity influences neuron number in six species of Puerto Rican Anolis.

Elucidating the selective forces shaping the diversity of vertebrate brains continues to be a major area of inquiry, particularly as it relates to cognition. Historically brain evolution was interpreted through the lens of relative brain size; however, recent evidence has challenged this approach. Investigating neuroanatomy at a finer scale, such as neuron number, can provide new insights into the forces shaping brain evolution in the context of information processing capacity. Ecological factors, such as the complexity of a species' habitat, place demands on cognition that could shape neuroanatomy. In this study, we investigate the relationship between neuron number and habitat complexity in three brain regions across six closely related anole species from Puerto Rico. After controlling for brain mass, we found that the number of neurons increased with habitat complexity across species in the telencephalon and 'rest of the brain,' but not in the cerebellum. Our results demonstrate that habitat complexity has shaped neuroanatomy in the Puerto Rican anole radiation and provide further evidence of the role of habitat complexity in vertebrate brain evolution.

Unlocking ground-based imagery for habitat mapping.

Fine-grained environmental data across large extents are needed to resolve the processes that impact species communities from local to global scales. Ground-based images (GBIs) have the potential to capture habitat complexity at biologically relevant spatial and temporal resolutions. Moving beyond existing applications of GBIs for species identification and monitoring ecological change from repeat photography, we describe promising approaches to habitat mapping, leveraging multimodal data and computer vision. We illustrate empirically how GBIs can be applied to predict distributions of species at fine scales along Street View routes, or to automatically classify and quantify habitat features. Further, we outline future research avenues using GBIs that can bring a leap forward in analyses for ecology and conservation with this underused resource.

Ecological and morphological correlates of visual acuity in birds.

Birds use their visual systems for important tasks, such as foraging and predator detection, that require them to resolve an image. However, visual acuity (the ability to perceive spatial detail) varies by two orders of magnitude across birds. Prior studies indicate that eye size and aspects of a species' ecology may drive variation in acuity, but these studies have been restricted to small numbers of species. We used a literature review to gather data on acuity measured either behaviorally or anatomically for 94 species from 38 families. We then examined how acuity varies in relation to (1) eye size, (2) habitat spatial complexity, (3) habitat light level, (4) diet composition, (5) prey mobility and (6) foraging mode. A phylogenetically controlled model including all of the above factors as predictors indicated that eye size and foraging mode are significant predictors of acuity. Examining each ecological variable in turn revealed that acuity is higher in species whose diet comprises vertebrates or scavenged food and whose foraging modes require resolving prey from farther away. Additionally, species that live in spatially complex, vegetative habitats have lower acuity than expected for their eye sizes. Together, our results suggest that the need to detect important objects from far away - such as predators for species that live in open habitats, and food items for species that forage on vertebrate and scavenged prey - has likely been a key driver of higher acuity in some species, helping us to elucidate how visual capabilities may be adapted to an animal's visual needs.

Naturally Occurring Vegetation Connectivity Facilitates Ant-Mediated Coffee Berry Borer Removal.

Vegetation connectivity is an essential aspect of the habitat complexity that impacts species interactions at local scales. However, agricultural intensification reduces connectivity in agroforestry systems, including coffee agroecosystems, which may hinder the movement of natural enemies and reduce the ecosystem services that they provide. Ants play an important role in regulating the coffee berry borer (CBB), which is the most damaging coffee pest. For arboreal ant communities, the connections between trees are important structures that facilitate ant mobility, resource recruitment, foraging success, and pest control ability. To better understand how connectivity impacts arboreal ants in coffee agroecosystems, we conducted an experiment to assess the impact of artificial (string) and naturally occurring vegetation (vines, leaves, branches) connectivity on Azteca sericeasur behavior on coffee plants. We compared ant activity, resource recruitment, and CBB removal rates across three connectivity treatments connecting coffee plants to A. sericeasur nest trees: vegetation connectivity, string, and control (not connected) treatments. We found higher rates of ant activity, resource recruitment, and CBB removal on plants with naturally occurring vegetation connections to A. sericeasur nest trees. Artificial connectivity (string) increased the rates of resource recruitment and CBB removal but to a lesser extent than vegetation connectivity. Moreover, vegetation connectivity buffered reductions in ant activity with distance from the ant nest tree. These results reinforce how habitat complexity in the form of vegetation connectivity impacts interspecific interactions at the local scale. Our results also suggest that leaving some degree of vegetation connectivity between coffee plants and shade trees can promote ant-mediated biological pest control in coffee systems.

Recent and rapid reef recovery around Koh Phangan Island, Gulf of Thailand, driven by plate-like hard corals.

Mass bleaching events and local anthropogenic influences have changed the benthic communities of many coral reefs with pronounced spatial differences that are linked to resilience patterns. The Gulf of Thailand is an under-investigated region with only few existing datasets containing long-term developments of coral reef communities using the same method at fixed sites. We thus analyzed benthic community data from seven reefs surrounding the island of Koh Phangan collected between 2014 and 2022. Findings revealed that the average live hard coral cover around Koh Phangan increased from 37% to 55% over the observation period, while turf algae cover decreased from 52% to 29%, indicating some recovery of local reefs. This corresponds to a mean increased rate of coral cover by 2.2% per year. The increase in live hard coral cover was mainly driven by plate-like corals, which quadrupled in proportion over the last decade from 7% to 28% while branching corals decreased in proportion from 9% to 2%. Furthermore, the hard coral genus richness increased, indicating an increased hard coral diversity. While in other reefs, increasing live hard coral cover is often attributed to fast-growing, branching coral species, considered more susceptible to bleaching and other disturbances, the reefs around Koh Phangan recovered mainly via growth of plate-like corals, particularly of the genus Montipora. Although plate-like morphologies are not necessarily more bleaching tolerant, they are important for supporting reef fish abundance and structural complexity on reefs, aiding reef recovery and sturdiness. Hence, our findings indicate that the intensity of local stressors around Kho Phangan allows reef recovery driven by some hard coral species.

Dead but not forgotten: complexity of Acropora palmata colonies increases with greater composition of dead coral.

Coral reefs are highly biodiverse ecosystems that have declined due to natural and anthropogenic stressors. Researchers often attribute reef ecological processes to corals' complex structure, but effective conservation requires disentangling the contributions of coral versus reef structures. Many studies assessing the relationships between reef structure and ecological dynamics commonly use live coral as a proxy for reef complexity, disregarding the contribution of dead coral skeletons to reef habitat provision or other biogeochemical reef dynamics. This study aimed to assess the contribution of dead coral to reef complexity by examining structural variations in live and dead Acropora palmata colonies. We used photogrammetry to reconstruct digital elevation models (DEMs) and orthomosaics of the benthic region immediately surrounding 10 A. palmata colonies. These reconstructions were used to quantify structural metrics, including surface rugosity, fractal dimension, slope, planform curvature, and profile curvature, as a function of benthic composition (i.e., live A. palmata, dead A. palmata, or non-A. palmata substrate). The results revealed that dead coral maintained more varied profile curvatures and higher fractal dimensions than live or non-coral substrate. Conversely, A. palmata colonies with a higher proportion of live coral displayed more uniform structure, with lower fractal dimensions and less variability in profile curvature measures. Other metrics showed no significant difference among substrate types. These findings provide novel insights into the structural differences between live and dead coral, and an alternative perspective on the mechanisms driving the observed structural complexity on reefs. Overall, our results highlight the overlooked potential contributions of dead coral to reef habitat provision, ecological processes, and other biogeochemical reef dynamics, and could have important implications for coral reef conservation.

Habitat complexity drives food web structure along a dynamic mangrove coast.

Structurally complex habitats, such as mangrove forests, allow for rich assemblages of species that benefit from the provided space, volume and substrate. Changes in habitat complexity can affect species abundance, diversity and resilience. In this study, we explored the effects of habitat complexity on food web networks in four developmental stages of mangrove forests with differing structural complexities: climax > degrading > colonizing > bare, by analyzing food web structure, stable isotopes and habitat complexity. We found that food webs became gradually more biodiverse (species richness: +119 %), complex (link density: +39 %), and robust (connectance: -35 %) in climax versus bare stages with increasing complexity of the mangrove forest (i.e., number of trees, leaf cover, and pneumatophore densities). This study shows that habitat complexity drives food web network structure in dynamic mangrove forests. We recommend restoration practitioners to use this food web network approach to quantify habitat restoration successes complementary to traditional biodiversity metrics.

Microhabitat conditions drive uncertainty of risk and shape neophobic responses in Trinidadian guppies, Poecilia reticulata.

In response to uncertain risks, prey may rely on neophobic phenotypes to reduce the costs associated with the lack of information regarding local conditions. Neophobia has been shown to be driven by information reliability, ambient risk and predator diversity, all of which shape uncertainty of risk. We similarly expect environmental conditions to shape uncertainty by interfering with information availability. In order to test how environmental variables might shape neophobic responses in Trinidadian guppies (Poecilia reticulata), we conducted an in situ field experiment of two high-predation risk guppy populations designed to determine how the 'average' and 'variance' of several environmental factors might influence the neophobic response to novel predator models and/or novel foraging patches. Our results suggest neophobia is shaped by water velocity, microhabitat complexity, pool width and depth, as well as substrate diversity and heterogeneity. Moreover, we found differential effects of the 'average' and 'variance' environmental variables on food- and predator-related neophobia. Our study highlights that assessment of neophobic drivers should consider predation risk, various microhabitat conditions and neophobia being tested. Neophobic phenotypes are expected to increase the probability of prey survival and reproductive success (i.e. fitness), and are therefore likely linked to population health and species survival. Understanding the drivers and consequences of uncertainty of risk is an increasingly pressing issue, as ecological uncertainty increases with the combined effects of climate change, anthropogenic disturbances and invasive species.

Complex battlefields favor strong soldiers over large armies in social animal warfare.

In social animals, success can depend on the outcome of group battles. Theoretical models of warfare predict that group fighting ability is proportional to two key factors: the strength of each soldier in the group and group size. The relative importance of these factors is predicted to vary across environments [F. W. Lanchester, Aircraft in Warfare, the Dawn of the Fourth Arm (1916)]. Here, we provide an empirical validation of the theoretical prediction that open environments should favor superior numbers, whereas complex environments should favor stronger soldiers [R. N. Franks, L. W. Partridge, Anim. Behav. 45, 197-199 (1993)]. We first demonstrate this pattern using simulated battles between relatively strong and weak soldiers in a computer-driven algorithm. We then validate this result in real animals using an ant model system: In battles in which the number of strong native meat ant Iridomyrmex purpureus workers is constant while the number of weak non-native invasive Argentine ant Linepithema humile workers increases across treatments, fatalities of I. purpureus are lower in complex than in simple arenas. Our results provide controlled experimental evidence that investing in stronger soldiers is more effective in complex environments. This is a significant advance in the empirical study of nonhuman warfare and is important for understanding the competitive balance among native and non-native invasive ant species.

Substrate Heterogeneity as a Trigger for Species Diversity in Marine Benthic Assemblages.

Many studies show that habitat complexity or habitat diversity plays a major role in biodiversity throughout different spatial scales: as structural heterogeneity increases, so does the number of available (micro-) habitats for the potential species inventory. The capability of housing species (even rare species) increases rapidly with increasing habitat heterogeneity. However, habitat complexity is not easy to measure in marine sublittoral sediments. In our study, we came up with a proposal to estimate sublittoral benthic habitat complexity using standard underwater video techniques. This tool was subsequently used to investigate the effect of habitat complexity on species richness in comparison to other environmental parameters in a marine protected area situated in the Fehmarn Belt, a narrow strait in the southwestern Baltic Sea. Our results show that species richness is significantly higher in heterogeneous substrates throughout all considered sediment types. Congruently, the presence of rare species increases with structural complexity. Our findings highlight the importance of the availability of microhabitats for benthic biodiversity as well as of the study area for regional ecosystem functioning.

Taxonomical and functional analyses of epifaunal polychaetes associated with Mussismilia spp.: the effects of coral growth morphology.

Background: The increasing habitat heterogeneity and complexity shows positive effects over different communities, leading to environmental diversity, access to resources and reducing the effectiveness of predation. In the present study we evaluate the structural and functional patterns of polychaete assemblages of three Mussismilia species with different coral morphology. Mussismilia hispida has a massive growth pattern; M. braziliensis also is a massive coral but forms a crevice in the corallum base; and M. harttii has a meandroid pattern. Methods: Ten individuals of the three Mussismilia species were sampled in two reefs in the Todos-os-Santos Bay, and we analyzed the differences in richness and abundance of polychaete species and the functional diversity metrics: Rao's quadratic entropy, functional dispersion, functional evenness, number of functional groups and functional richness, among Mussismilia species. Results: Two-way ANOVA with permutations showed significant differences for polychaete abundances and richness among Mussismilia species (higher values for M. harttii), but no differences were recorded when compared between the two coral reef areas studied. There was no statistical difference among coral species or between reefs in relation to the functional diversity components influenced by abundance, such as Rao quadratic entropy, functional dispersion, and functional evenness. Some individual polychaete functional traits presented differences among Mussismilia species, and that also helped us to build a picture about the effect of different growth structures over functional aspects of polychaete assemblages. Thus, the taxonomical approach, the analysis of individual functional traits and the functional diversity metrics are fundamental tools to characterize the assemblage of organisms associated with corals.

Review of the Effects of Enclosure Complexity and Design on the Behaviour and Physiology of Zoo Animals.

The complexity of the habitat refers to its physical geometry, which includes abiotic and biotic elements. Habitat complexity is important because it allows more species to coexist and, consequently, more interactions to be established among them. The complexity of the habitat links the physical structure of the enclosure to the biological interactions, which occur within its limits. Enclosure complexity should vary temporally, to be able to influence the animals in different ways, depending on the period of the day and season and throughout the year. In the present paper, we discuss how habitat complexity is important, and how it can positively influence the physical and mental states of zoo animals. We show how habitat complexity can ultimately affect educational projects. Finally, we discuss how we can add complexity to enclosures and, thus, make the lives of animals more interesting and functional.

Intertidal habitat complexity influences the density of the non-native crab Hemigrapsus sanguineus.

Habitat structural complexity can provide protection from predators, potentially affecting population density of native and non-native prey. The invasive Asian shore crab, Hemigrapsus sanguineus, occurs in variable densities in the rocky intertidal zone of eastern North America and northern Europe, often in densities greater than in its native range. The present study examined the influence of habitat complexity on the density of H. sanguineus. Artificial shelters of concrete pavers with stones arranged in increasing complexity were deployed in the intertidal zone along a rocky shore in southeastern Massachusetts, USA, for 21 consecutive weekly intervals in 2020. Crabs consistently reached the highest densities in the most complex shelters despite their lower internal surface area. In addition, crabs exhibited shelter selectivity based on body size, with large crabs occupying artificial shelters in greater numbers than adjacent natural substrate. In a subsequent lab study, crab activity over 1 h was observed in the presence of the same artificial shelters, under simulated tidal conditions. Shelter complexity had little influence on the number of crabs under the pavers although crabs were more active when submerged in water than exposed to air. These results show that crab density increases as habitat complexity increases, and complexity may serve as a predictor of H. sanguineus density but not short-term behavior.

A word on habitat complexity.

In their recent synopsis, Loke and Chisholm (Ecology Letters, 25, 2269-2288, 2022) present an overview of habitat complexity metrics for ecologists. They provide a review and some sound advice. However, we found several of their analyses and opinions misleading. This technical note provides a different perspective on the complexity metrics assessed.

Managing aquatic habitat structure for resilient trophic interactions.

Structural habitat (the three-dimensional arrangement of physical matter, abiotic and biotic, at a location) is a foundational element for the resilience and maintenance of biodiversity, yet anthropogenic development is driving the global simplification of aquatic environments. Resource managers regularly seek to conserve aquatic food webs by increasing structural habitat complexity with expected benefits to fisheries; however, the global effectiveness of such actions is unclear. Our synthesis and theoretical analyses found that the response of a consumer-resource interaction (predatory sportfish and forage fish prey) to the addition of prey refuge habitat differed among systems with low and high rates of biomass transfer from resource to consumer (i.e., biomass potential); stabilization was not the rule. Greater prey refuge habitat availability tended to stabilize systems characterized by high biomass potential while simultaneously increasing consumer densities. In contrast, increasing prey refuge habitat availability in systems characterized by low biomass potential tended to mute energy transfer and moved consumer densities toward local extinction. Importantly, biomass potential and prey refuge can have antagonistic effects on stability and relative consumer densities, and it is therefore important to consider the local conditions of a system when using habitat manipulation as a management measure. Further development of our context-dependent perspective to whole food webs, and across different environments, may help to guide structural habitat management to better restore and protect aquatic ecosystems.

Habitat attributes mediate herbivory and influence community development in algal metacommunities.

Understanding the drivers and impacts of spatiotemporal variation in species abundance on community trajectories is key to understanding the factors contributing to ecosystem resilience. Temporal variation in species trajectories across patches can provide compensation for species loss and can influence successional patterns. However, little is known about the underlying mechanisms that lead to patterns of species or spatial compensation and how those patterns may be mediated by consumer-resource relationships. Here we describe an experiment testing whether habitat attributes (e.g., structural complexity and spatial heterogeneity) mediate the effects of herbivory on tropical marine macroalgal communities by reducing accessibility and detectability, respectively, leading to variable trajectories among algal species at community (within patch) and metacommunity (i.e., among patch) scales. Reduced accessibility (greater habitat complexity) decreased the effects of herbivory (i.e., depressed consumption rate, increased algal species richness), and both accessibility and detectability (spatial heterogeneity) influenced algal community structure. Moreover, decreased accessibility at the community scale and a mosaic of accessibility at the metacommunity scale led to variation in community assembly. We suggest that habitat attributes can be important influencers of consumer-resource interactions on coral reefs, which in turn can increase species diversity, promote species succession, and enhance stability in algal metacommunities.