Ecosystem‐based management of coral reefs under climate change

Coral reefs provide food and livelihoods for hundreds of millions of people as well as harbour some of the highest regions of biodiversity in the ocean. However, overexploitation, land‐use change and other local anthropogenic threats to coral reefs have left many degraded. Additionally, coral reefs are faced with the dual emerging threats of ocean warming and acidification due to rising CO₂ emissions, with dire predictions that they will not survive the century. This review evaluates the impacts of climate change on coral reef organisms, communities and ecosystems, focusing on the interactions between climate change factors and local anthropogenic stressors. It then explores the shortcomings of existing management and the move towards ecosystem‐based management and resilience thinking, before highlighting the need for climate change‐ready marine protected areas (MPAs), reduction in local anthropogenic stressors, novel approaches such as human‐assisted evolution and the importance of sustainable socialecological systems. It concludes that designation of climate change‐ready MPAs, integrated with other management strategies involving stakeholders and participation at multiple scales such as marine spatial planning, will be required to maximise coral reef resilience under climate change. However, efforts to reduce carbon emissions are critical if the long‐term efficacy of local management actions is to be maintained and coral reefs are to survive.     

Responses of algal communities to gradients in herbivore biomass and water quality in Marovo Lagoon, Solomon Islands

Settlement tiles were used to characterise and quantify coral reef associated algal communities along water quality and herbivory gradients from terrestrial influenced near shore sites to oceanic passage sites in Marovo Lagoon, the Solomon Islands. After 6 months, settlement tile communities from inshore reefs were dominated by high biomass algal turfs (filamentous algae and cyanobacteria) whereas tiles located on offshore reefs were characterised by a mixed low biomass community of calcareous crustose algae, fleshy crustose algae and bare tile. The exclusion of macrograzers, via caging of tiles, on the outer reef sites resulted in the development of an algal turf community similar to that observed on inshore reefs. Caging on the inshore reef tiles had a limited impact on community composition or biomass. Water quality and herbivorous fish biomass were quantified at each site to elucidate factors that might influence algal community structure across the lagoon. Herbivore biomass was the dominant driver of algal community structure. Algal biomass on the other hand was controlled by both herbivory and water quality (particularly dissolved nutrients). This study demonstrates that algal communities on settlement tiles are an indicator capable of integrating the impacts of water quality and herbivory over a small spatial scale (kilometres) and short temporal scale (months), where other environmental drivers (current, light, regional variability) are constant.     

Coral diversity across a disturbance gradient in the Pulau Seribu reef complex off Jakarta, Indonesia

Very few coral reefs are located close enough to metropolitan cities to study the influence of large urban populations on reef communities. Here, we compare the impact of a large-scale disturbance gradient with local-scale disturbance on coral richness, cover, and composition in the Jakarta Bay and Pulau Seribu reef complex off Jakarta, Indonesia. We found no effect of local land-use type of coral reef islands on richness, composition or cover, nor did taxon richness differ among zones at the large-scale. There was, however, a pronounced difference in composition and coral cover among zones. Cover was very low and composition differed markedly in the near-shore zone 1 (Jakarta Bay) where human-induced disturbance is most intense. Cover was highest in the outlying reefs of zone 3. The highly perturbed zone 1 reefs were, furthermore, distinguished by the virtual absence of otherwise abundant coral taxa such as Acropora hyacinthus and Porites rus and the prevalence of taxa such as Oulastrea crispata and Favia maxima. Almost 60% of the spatial variation in composition was related to variation in shelf depth and island size. The importance of shelf depth was related to the prevalence of a strong environmental gradient in reef depth, pollution, and mechanical reef disturbance and salinity from Jakarta Bay to the outlying reefs. Although there was a significant univariate relationship between spatial variation in composition and distance, this did not enter into the multivariate model, except when presence–absence data was used, indicating that environmental processes are the primary structuring forces in determining local coral assemblage composition across the Pulau Seribu complex.     

Coral Reef Community Composition in the Context of Disturbance History on the Great Barrier Reef,Australia

Much research on coral reefs has documented differential declines in coral and associated organisms. In order to contextualise this general degradation, research on community composition is necessary in the context of varied disturbance histories and the biological processes and physical features thought to retard or promote recovery. We conducted a spatial assessment of coral reef communities across five reefs of the central Great Barrier Reef, Australia, with known disturbance histories, and assessed patterns of coral cover and community composition related to a range of other variables thought to be important for reef dynamics. Two of the reefs had not been extensively disturbed for at least 15 years prior to the surveys. Three of the reefs had been severely impacted by crown-of-thorns starfish outbreaks and coral bleaching approximately a decade before the surveys, from which only one of them was showing signs of recovery based on independent surveys. We incorporated wave exposure (sheltered and exposed) and reef zone (slope, crest and flat) into our design, providing a comprehensive assessment of the spatial patterns in community composition on these reefs. Categorising corals into life history groupings, we document major coral community differences in the unrecovered reefs, compared to the composition and covers found on the undisturbed reefs. The recovered reef, despite having similar coral cover, had a different community composition from the undisturbed reefs, which may indicate slow successional processes, or a different natural community dominance pattern due to hydrology and other oceanographic factors. The variables that best correlated with patterns in the coral community among sites included the density of juvenile corals, herbivore fish biomass, fish species richness and the cover of macroalgae. Given increasing impacts to the Great Barrier Reef, efforts to mitigate local stressors will be imperative to encouraging coral communities to persist into the future.     

Assembly rules of reef corals are flexible along a steep climatic gradient

Coral reefs, one of the world's most complex and vulnerable ecosystems, face an uncertain future in coming decades as they continue to respond to anthropogenic climate change, overfishing, pollution, and other human impacts [1, 2]. Traditionally, marine macroecology is based on presence/absence data from taxonomic checklists or geographic ranges, providing a qualitative overview of spatial shifts in species richness that treats rare and common species equally [3, 4]. As a consequence, regional and long-term shifts in relative abundances of individual taxa are poorly understood. Here we apply a more rigorous quantitative approach to examine large-scale spatial variation in the species composition and abundance of corals on midshelf reefs along the length of Australia's Great Barrier Reef, a biogeographic region where species richness is high and relatively homogeneous [5]. We demonstrate that important functional components of coral assemblages "sample" space differently at 132 sites separated by up to 1740 km, leading to complex latitudinal shifts in patterns of absolute and relative abundance. The flexibility in community composition that we document along latitudinal environmental gradients indicates that climate change is likely to result in a reassortment of coral reef taxa rather than wholesale loss of entire reef ecosystems.     

Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.     

Millennial-scale change in the structure of a Caribbean reef ecosystem and the role of human and natural disturbance

Caribbean coral reefs have transformed into algal-dominated habitats over the past half-century, but the role of specific anthropogenic drivers is unresolved due to the lack of ecosystem-level data predating human disturbance. To better understand the extent and causes of long-term Caribbean reef declines, we produced a continuous 3000-yr record of the ecosystem state of three reefs in Bocas del Toro, Caribbean Panama. From fossils and sediments obtained from reef matrix cores, we tracked changes in reef accretion rates and the taxonomic and functional group composition of fish, coral, urchin, bivalve and benthic foraminifera. This dataset provided a comprehensive picture of reef community and environmental change. At all sites, reefs shifted from systems with greater relative abundance of herbivorous fish, epifaunal suspension feeding bivalves and Diadema urchins to systems with greater relative abundance of micropredator fish, infaunal bivalves and Echinometra urchins. These transitions were initiated a millennium ago at two less-degraded reefs fringing offshore islands and ~250 yr ago at a degraded patch reef near the continental coast. Ecosystem shifts were accompanied by a decline in reef accretion rates, and at the patch reef, a decline in water quality since the 18th century. Within all cores, synchronous increases in infaunal bivalves and declines in herbivorous fish regardless of water quality suggest a loss of hard substrate and increasingly hypoxic sediment conditions related to herbivore loss. While the early timing of ecosystem transitions at the fringing reefs implicates large-scale hydrological change, the more recent timing of change and loss of water quality at the patch reef implicates terrigenous runoff from land-clearing. Our whole-ecosystem reconstruction reveals that reef ecosystem deterioration appears to follow a predictable trajectory whether driven by natural or anthropogenic disturbances and that historical local human activities have quickly unraveled reefs at a scale similar to longer-term natural environmental change.     

Historical reconstruction reveals recovery in Hawaiian coral reefs

Coral reef ecosystems are declining worldwide, yet regional differences in the trajectories, timing and extent of degradation highlight the need for in-depth regional case studies to understand the factors that contribute to either ecosystem sustainability or decline. We reconstructed social-ecological interactions in Hawaiian coral reef environments over 700 years using detailed datasets on ecological conditions, proximate anthropogenic stressor regimes and social change. Here we report previously undetected recovery periods in Hawaiian coral reefs, including a historical recovery in the MHI (~AD 1400-1820) and an ongoing recovery in the NWHI (~AD 1950-2009+). These recovery periods appear to be attributed to a complex set of changes in underlying social systems, which served to release reefs from direct anthropogenic stressor regimes. Recovery at the ecosystem level is associated with reductions in stressors over long time periods (decades+) and large spatial scales (>10(3) km(2)). Our results challenge conventional assumptions and reported findings that human impacts to ecosystems are cumulative and lead only to long-term trajectories of environmental decline. In contrast, recovery periods reveal that human societies have interacted sustainably with coral reef environments over long time periods, and that degraded ecosystems may still retain the adaptive capacity and resilience to recover from human impacts.     

Short-term changes of fish assemblages observed in the near-pristine reefs of the Phoenix Islands

Climate change-related disturbances are increasingly recognized as critical threats to biodiversity and species abundance. On coral reefs, climate disturbances have known consequences for reef fishes, but it is often difficult to isolate the effect of coral bleaching from preceding or simultaneous disturbances such as fishing, pollution, and habitat loss. In this study, pre-bleaching surveys of fish family assemblages in the remote Phoenix Islands in 2002 are compared to post-bleaching in 2005, following severe thermal stress. Post-bleaching, total coral cover decreased substantially, as did the combined abundance of all fish families. Yet, changes in abundance for specific fish families were not uniform, and varied greatly from site to site. Of the 13 fish families examined, 3 exhibited significant changes in abundance from 2002 to 2005, regardless of site (Carangidae, Chaetodontidae, and serranid subfamily Epinephelinae). For these families, we explored whether changes in abundance were related to island type (island vs atoll) and/or declining coral cover (percent change). Carangidae on islands experienced larger changes in abundance than those on atolls, though declines in abundance over time were not associated with changes in live coral cover. In contrast, for Chaetodontidae, declines in abundance over time were most dramatic on atolls, and were also associated with changes in live coral cover. The remoteness of the Phoenix Islands excludes many typical local anthropogenic stressors as drivers of short-term changes; observed changes are instead more likely attributed to natural variation in fish populations, or associated with coral loss following the 2002–2003 major thermal stress event.     

Ecological limitations to the resilience of coral reefs

The decline of coral reefs has been broadly attributed to human stressors being too strong and pervasive, whereas biological processes that may render coral reefs fragile have been sparsely considered. Here we review several ecological factors that can limit the ability of coral reefs to withstand disturbance. These include: (1) Many species lack the adaptive capacity to cope with the unprecedented disturbances they currently face; (2) human disturbances impact vulnerable life history stages, reducing reproductive output and the supply of recruits essential for recovery; (3) reefs can be vulnerable to the loss of few species, as niche specialization or temporal and spatial segregation makes each species unique (i.e., narrow ecological redundancy); in addition, many foundation species have similar sensitivity to disturbances, suggesting that entire functions can be lost to single disturbances; and (4) feedback loops and extinction vortices may stabilize degraded states or accelerate collapses even if stressors are removed. This review suggests that the degradation of coral reefs is due to not only the severity of human stressors but also the “fragility” of coral reefs. As such, appropriate governance is essential to manage stressors while being inclusive of ecological process and human uses across transnational scales. This is a considerable but necessary upgrade in current management if the integrity, and delivery of goods and services, of coral reefs is to be preserved.     

The Reef Environment Centralized InFormation System (RECIFS): An integrated geo-environmental database for coral reef research and conservation

Motivation

Host to intricate networks of marine species, coral reefs are among the most biologically diverse ecosystems on Earth. Over the past few decades, major degradations of coral reefs have been observed worldwide, which is largely attributed to the effects of climate change and local stressors related to human activities. Now more than ever, characterizing how the environment shapes the dynamics of the reef ecosystem (e.g., shifts in species abundance, community changes, emergence of locally adapted populations) is key to uncovering the environmental drivers of reef degradation, and developing efficient conservation strategies in response. To achieve these objectives, it is pivotal that environmental data describing the processes driving such ecosystem dynamics, which occur across specific spatial and temporal scales, are easily accessible to coral reef researchers and conservation stakeholders alike.

Main types of variable contained

Multiple environmental variables characterizing various facets of the reef environment, including water chemistry and physics (e.g., temperature, pH, chlorophyll concentration), local anthropogenic pressures (e.g., boat traffic, distance from agricultural or urban areas) and sea currents patterns.

Spatial location and grain

Worldwide reef cells of 5 by 5 km.

Time period and grain

Last 3–4 decades, monthly and yearly resolution.

Major taxa and level of measurement

Environmental data important for coral reefs and associated biodiversity.

Software format

Interactive web application available at https://recifs.epfl.ch .     

Ecological condition of coral reef assemblages in the Cuban Archipelago

The condition of coral reefs in the Cuban Archipelago is poorly known. We aimed to analyse coral assemblages across 199 reef sites belonging to 12 localities. Crest and fore reefs were assessed using six metrics: species richness, density, coral cover, mortality, coral size and reef complexity. The condition of reefs varied across the archipelago from healthy to depleted reefs. The localities with best scores were Cienfuegos, Bahía de Cochinos and Cazones. These reefs have values of living coral cover (>20%) and complexity (>50?cm) similar to the best preserved Caribbean reefs. However, the majority of crest biotopes suffered important deterioration with old mortality of Acropora palmata populations and moderate coral cover (15%); although crest reefs still maintained their structural complexity. Despite moderate levels of coral cover in fore reefs (18%), their condition was alarming because 25% of the sites had cover below the recovery threshold of 10%, accumulated mortality and structural flattening. Compared with the 1980s, the species richness was roughly the same (42) for crest and fore reefs, although dominance has changed to widespread tolerant species. Coral reef assemblages varied at local and regional scales in similar magnitude, suggesting the combined effects of natural and anthropogenic drivers.     

Simulations of Long-Term Community Dynamics in Coral Reefs - How Perturbations Shape Trajectories

Tropical coral reefs feature extraordinary biodiversity and high productivity rates in oligotrophic waters. Due to increasing frequencies of perturbations – anthropogenic and natural – many reefs are under threat. Such perturbations often have devastating effects on these unique ecosystems and especially if they occur simultaneously and amplify each other''s impact, they might trigger a phase shift and create irreversible conditions.We developed a generic, spatially explicit, individual-based model in which competition drives the dynamics of a virtual benthic reef community – comprised of scleractinian corals and algae – under different environmental settings. Higher system properties, like population dynamics or community composition arise through self-organization as emergent properties. The model was parameterized for a typical coral reef site at Zanzibar, Tanzania and features coral bleaching and physical disturbance regimes as major sources of perturbations. Our results show that various types and modes (intensities and frequencies) of perturbations create diverse outcomes and that the switch from high diversity to single species dominance can be evoked by small changes in a key parameter.Here we extend the understanding of coral reef resilience and the identification of key processes, drivers and respective thresholds, responsible for changes in local situations. One future goal is to provide a tool which may aid decision making processes in management of coral reefs.     

The effects of top–down versus bottom–up control on benthic coral reef community structure

While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral–algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.     

Operationalizing resilience for adaptive coral reef management under global environmental change

Cumulative pressures from global climate and ocean change combined with multiple regional and local‐scale stressors pose fundamental challenges to coral reef managers worldwide. Understanding how cumulative stressors affect coral reef vulnerability is critical for successful reef conservation now and in the future. In this review, we present the case that strategically managing for increased ecological resilience (capacity for stress resistance and recovery) can reduce coral reef vulnerability (risk of net decline) up to a point. Specifically, we propose an operational framework for identifying effective management levers to enhance resilience and support management decisions that reduce reef vulnerability. Building on a system understanding of biological and ecological processes that drive resilience of coral reefs in different environmental and socio‐economic settings, we present an Adaptive Resilience‐Based management (ARBM) framework and suggest a set of guidelines for how and where resilience can be enhanced via management interventions. We argue that press‐type stressors (pollution, sedimentation, overfishing, ocean warming and acidification) are key threats to coral reef resilience by affecting processes underpinning resistance and recovery, while pulse‐type (acute) stressors (e.g. storms, bleaching events, crown‐of‐thorns starfish outbreaks) increase the demand for resilience. We apply the framework to a set of example problems for Caribbean and Indo‐Pacific reefs. A combined strategy of active risk reduction and resilience support is needed, informed by key management objectives, knowledge of reef ecosystem processes and consideration of environmental and social drivers. As climate change and ocean acidification erode the resilience and increase the vulnerability of coral reefs globally, successful adaptive management of coral reefs will become increasingly difficult. Given limited resources, on‐the‐ground solutions are likely to focus increasingly on actions that support resilience at finer spatial scales, and that are tightly linked to ecosystem goods and services.     

Baselines and degradation of coral reefs in the Northern Line Islands

Effective conservation requires rigorous baselines of pristine conditions to assess the impacts of human activities and to evaluate the efficacy of management. Most coral reefs are moderately to severely degraded by local human activities such as fishing and pollution as well as global change, hence it is difficult to separate local from global effects. To this end, we surveyed coral reefs on uninhabited atolls in the northern Line Islands to provide a baseline of reef community structure, and on increasingly populated atolls to document changes associated with human activities. We found that top predators and reef-building organisms dominated unpopulated Kingman and Palmyra, while small planktivorous fishes and fleshy algae dominated the populated atolls of Tabuaeran and Kiritimati. Sharks and other top predators overwhelmed the fish assemblages on Kingman and Palmyra so that the biomass pyramid was inverted (top-heavy). In contrast, the biomass pyramid at Tabuaeran and Kiritimati exhibited the typical bottom-heavy pattern. Reefs without people exhibited less coral disease and greater coral recruitment relative to more inhabited reefs. Thus, protection from overfishing and pollution appears to increase the resilience of reef ecosystems to the effects of global warming.     

Diversity among Macroalgae-Consuming Fishes on Coral Reefs: A Transcontinental Comparison

Despite high diversity and abundance of nominally herbivorous fishes on coral reefs, recent studies indicate that only a small subset of taxa are capable of removing dominant macroalgae once these become established. This limited functional redundancy highlights the potential vulnerability of coral reefs to disturbance and stresses the need to assess the functional role of individual species of herbivores. However, our knowledge of species-specific patterns in macroalgal consumption is limited geographically, and there is a need to determine the extent to which patterns observed in specific reefs can be generalised at larger spatial scales. In this study, video cameras were used to quantify rates of macroalgae consumption by fishes in two coral reefs located at a similar latitude in opposite sides of Australia: the Keppel Islands in the Great Barrier Reef (eastern coast) and Ningaloo Reef (western coast). The community of nominally herbivorous fish was also characterised in both systems to determine whether potential differences in the species observed feeding on macroalgae were related to spatial dissimilarities in herbivore community composition. The total number of species observed biting on the dominant brown alga Sargassum myriocystum differed dramatically among the two systems, with 23 species feeding in Ningaloo, compared with just 8 in the Keppel Islands. Strong differences were also found in the species composition and total biomass of nominally herbivorous fish, which was an order of magnitude higher in Ningaloo. However, despite such marked differences in the diversity, biomass, and community composition of resident herbivorous fishes, Sargassum consumption was dominated by only four species in both systems, with Naso unicornis and Kyphosus vaigiensis consistently emerging as dominant feeders of macroalgae.     

Ant communities on small tropical islands: effects of island size and isolation are obscured by habitat disturbance and ‘tramp’ ant species

Aim Comparisons among islands offer an opportunity to study the effects of biotic and abiotic factors on small, replicated biological communities. Smaller population sizes on islands accelerate some ecological processes, which may decrease the time needed for perturbations to affect community composition. We surveyed ants on 18 small tropical islands to determine the effects of island size, isolation from the mainland, and habitat disturbance on ant community composition. Location Thousand Islands Archipelago (Indonesian name: Kepulauan Seribu) off Jakarta, West Java, Indonesia. Methods Ants were sampled from the soil surface, leaf litter and vegetation in all habitat types on each island. Island size, isolation from the mainland, and land‐use patterns were quantified using GIS software. The presence of settlements and of boat docks were used as indicators of anthropogenic disturbance. The richness of ant communities and non‐tramp ant species on each island were analysed in relation to the islands’ physical characteristics and indicators of human disturbance. Results Forty‐eight ant species from 5 subfamilies and 28 genera were recorded from the archipelago, and approximately 20% of the ant species were well‐known human‐commensal ‘tramp’ species. Islands with boat docks or human settlements had significantly more tramp species than did islands lacking these indicators of anthropogenic disturbance, and the diversity of non‐tramp species decreased with habitat disturbance. Main conclusions Human disturbance on islands in the Thousand Islands Archipelago promotes the introduction and/or establishment of tramp species. Tramp species affect the composition of insular ant communities, and expected biogeographical patterns of ant richness are masked. The island with the greatest estimated species richness and the greatest number of unique ant species, Rambut Island, is a forested bird sanctuary, highlighting the importance of protected areas in preserving the diversity of species‐rich invertebrate faunas.     

Episodic Disturbance from Boat Anchoring Is a Major Contributor to,but Does Not Alter the Trajectory of,Long-Term Coral Reef Decline

Isolating the relative effects of episodic disturbances and chronic stressors on long-term community change is challenging. We assessed the impact of an episodic disturbance associated with human visitation (boat anchoring) relative to other drivers of long-term change on coral reefs. A one-time anchoring event at Crab Cove, British Virgin Islands, in 2004 caused rapid losses of coral and reef structural complexity that were equal to the cumulative decline over 23 years observed at an adjacent site. The abundance of small site-attached reef fishes dropped by approximately one quarter after the anchoring event, but this drop was not immediate and only fully apparent two years after the anchoring event. There was no obvious recovery from the impact, and no evidence that this episodic impact accelerated or retarded subsequent declines from other causes. This apparent lack of synergism between the effect of this episodic human impact and other chronic stressors is consistent with the few other long-term studies of episodic impacts, and suggests that action to mitigate anchor damage should yield predictable benefits.     

Spatial patterns of Yucatan reef fish communities: Testing models using a multi-scale survey design

Using a hierarchical multi-scale survey design, we examined the spatial patterns of reef fish communities and tested ecological models concerning the relative importance of reef geomorphology and anthropogenic pressure possibly driving community structure. Canonical redundancy analysis was used as a form of multivariate analysis of variance (MANOVA) to asses differences in reef fish community composition at two spatial scales: broad (10⁵ m) and intermediate (10⁴ m). Surveys were conducted on the east coast of the Yucatan Peninsula (Mexican Caribbean fringing reef), including regions and reefs which differed in geomophologic structure and human use. Seven hundred and fourteen line transects were distributed among 13 reef localities belonging to different regions established a priori. Transects covered four types of reef habitat: lagoon, front, slope, and terrace. Tests of significance were based on permutation procedures. Significant differences among regions were found for the lagoon, slope, and terrace fish communities, consistent with the geomorphologic model, but it is only in the reef lagoon that they were consistent with the anthropogenic model, which may indicate an effect of coastal human activities. Significant differences among reefs within regions were observed, which could be associated with local environmental gradients. Canonical nested MANOVA was an appropriate method for testing ecological hypotheses about the functioning of complex biological systems. The use of a surveying strategy that explicitly incorporated the spatial structure represents an important contribution of this paper to coral reef fish ecology.