Bright spots among the world's coral reefs.

Ongoing declines in the structure and function of the world's coral reefs require novel approaches to sustain these ecosystems and the millions of people who depend on them3. A presently unexplored approach that draws on theory and practice in human health and rural development is to systematically identify and learn from the 'outliers'­places where ecosystems are substantially better ('bright spots') or worse ('dark spots') than expected, given the environmental conditions and socioeconomic drivers they are exposed to. Here we compile data from more than 2,500 reefs worldwide and develop a Bayesian hierarchical model to generate expectations of how standing stocks of reef fish biomass are related to 18 socioeconomic drivers and environmental conditions. We identify 15 bright spots and 35 dark spots among our global survey of coral reefs, defined as sites that have biomass levels more than two standard deviations from expectations. Importantly, bright spots are not simply comprised of remote areas with low fishing pressure; they include localities where human populations and use of ecosystem resources is high, potentially providing insights into how communities have successfully confronted strong drivers of change. Conversely, dark spots are not necessarily the sites with the lowest absolute biomass and even include some remote, uninhabited locations often considered near pristine6. We surveyed local experts about social, institutional, and environmental conditions at these sites to reveal that bright spots are characterized by strong sociocultural institutions such as customary taboos and marine tenure, high levels of local engagement in management, high dependence on marine resources, and beneficial environmental conditions such as deep-water refuges. Alternatively, dark spots are characterized by intensive capture and storage technology and a recent history of environmental shocks. Our results suggest that investments in strengthening fisheries governance, particularly aspects such as participation and property rights, could facilitate innovative conservation actions that help communities defy expectations of global reef degradation.

Gravity of human impacts mediates coral reef conservation gains.

Coral reefs provide ecosystem goods and services for millions of people in the tropics, but reef conditions are declining worldwide. Effective solutions to the crisis facing coral reefs depend in part on understanding the context under which different types of conservation benefits can be maximized. Our global analysis of nearly 1,800 tropical reefs reveals how the intensity of human impacts in the surrounding seascape, measured as a function of human population size and accessibility to reefs ("gravity"), diminishes the effectiveness of marine reserves at sustaining reef fish biomass and the presence of top predators, even where compliance with reserve rules is high. Critically, fish biomass in high-compliance marine reserves located where human impacts were intensive tended to be less than a quarter that of reserves where human impacts were low. Similarly, the probability of encountering top predators on reefs with high human impacts was close to zero, even in high-compliance marine reserves. However, we find that the relative difference between openly fished sites and reserves (what we refer to as conservation gains) are highest for fish biomass (excluding predators) where human impacts are moderate and for top predators where human impacts are low. Our results illustrate critical ecological trade-offs in meeting key conservation objectives: reserves placed where there are moderate-to-high human impacts can provide substantial conservation gains for fish biomass, yet they are unlikely to support key ecosystem functions like higher-order predation, which is more prevalent in reserve locations with low human impacts.

Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes.

Difficulties in scaling up theoretical and experimental results have raised controversy over the consequences of biodiversity loss for the functioning of natural ecosystems. Using a global survey of reef fish assemblages, we show that in contrast to previous theoretical and experimental studies, ecosystem functioning (as measured by standing biomass) scales in a non-saturating manner with biodiversity (as measured by species and functional richness) in this ecosystem. Our field study also shows a significant and negative interaction between human population density and biodiversity on ecosystem functioning (i.e., for the same human density there were larger reductions in standing biomass at more diverse reefs). Human effects were found to be related to fishing, coastal development, and land use stressors, and currently affect over 75% of the world's coral reefs. Our results indicate that the consequences of biodiversity loss in coral reefs have been considerably underestimated based on existing knowledge and that reef fish assemblages, particularly the most diverse, are greatly vulnerable to the expansion and intensity of anthropogenic stressors in coastal areas.

Development and analyses of stakeholder driven conceptual models to support the implementation of ecosystem-based fisheries management in the U.S. Caribbean.

Fisheries management agencies in the U.S. Caribbean are currently taking steps into transitioning from a single species approach to one that includes Ecosystem Based Fisheries Management (EBFM) considerations. In this study, we developed and analyzed stakeholder-driven conceptual models with seven different stakeholder groups in Puerto Rico and the US Virgin Islands to assess and compare their perceptions of the fishery ecosystem. Conceptual models were developed for each stakeholder group during 29 separate workshops involving a total of 236 participants representing Commercial Fishers, Managers, Academics, Local Businesses, Environmental NGOs, and the Caribbean Fishery Management Council (CFMC) District Advisory Panels (DAPs) and Scientific and Statistical Committee (SSC). Non-Metric Multidimensional Scaling (nMDS) and two-mode social network analysis were used to investigate differences and similarities between stakeholder groups as well as to identify priority ecosystem elements and threats. Results show important variations between stakeholders and islands in terms of their perceived importance of ecosystem components and relationships, which supports the need for collaborative approaches and co-production of knowledge in the United States (U.S.) Caribbean region. Despite this variation, important areas of common concern among stakeholders were identified such as: habitat integrity (e.g., coral reefs), water quality, and influence of recreational fisheries and tourism on marine ecosystems. Findings of this study support the use of stakeholder-driven conceptual models as effective tools to guide decision-making, aid prioritization of data collection, and increase collaboration and cooperation among stakeholders in the context of fisheries management.

Growth dynamics in Acropora cervicornis and A. prolifera in southwest Puerto Rico.

Natural population recovery of Acropora palmata, A. cervicornis and their hybrid, Acropora prolifera, have fluctuated significantly after their Caribbean-wide, disease-induced mass mortality in the early 1980s. Even though significant recovery has been observed in a few localities, recurrent disease outbreaks, bleaching, storm damage, local environmental deterioration, algae smothering, predation, low sexual recruitment and low survivorship have affected the expected, quick recovery of these weedy species. In this study, the status of three recovering populations of A. cervicornis and two of A. prolifera were assessed over one year using coral growth and mortality metrics, and changes in their associated algae and fish/invertebrate communities in three localities in the La Parguera Natural Reserve (LPNR), southwest coast of Puerto Rico. Five branches were tagged in each of 29, medium size (1-2 m in diameter) A. cervicornis and 18 A. prolifera colonies in the Media Luna, Mario and San Cristobal reefs off LPNR. Branches were measured monthly, together with observations to evaluate associated disease(s), algae accumulation and predation. A. cervicornis grew faster [3.1 ± 0.44 cm/month (= 37.2 cm/y)] compared to A. prolifera [2.6 ± 0.41 cm/month (= 31.2 cm/y)], and growth was significantly higher during Winter-Spring compared to Summer-Fall for both taxa (3.5 ± 0.58 vs. 0.53 ± 0.15 cm/month in A. cervicornis, and 2.43 ± 0.71 vs. 0.27 ± 0.20 cm/month in A. prolifera, respectively). Algal accumulation was only observed in A. cervicornis, and was higher during Spring-Summer compared to Fall-Winter (6.1 ± 0.91 cm/month and 3.8 ± 0.29 cm/month, respectively, (PERMANOVA, df = 2, MS = 10.2, p = 0.37)). Mortality associated with white band disease, algae smothering and fish/invertebrate predation was also higher in A. cervicornis and varied among colonies within sites, across sites and across season. The balance between tissue grow and mortality determines if colonies survive. This balance seems to be pushed to the high mortality side often by increasing frequency of high thermal anomalies, inducing bleaching and disease outbreaks and other factors, which have historically impacted the natural recovery of these taxa in the La Parguera Natural Reserve in Puerto Rico and possibly other areas in the region. Overall, results indicate variability in both growth and mortality rates in both taxa across localities and seasons, with A. cervicornis showing overall higher mortalities compared to A. prolifera.

Meeting fisheries, ecosystem function, and biodiversity goals in a human-dominated world.

The worldwide decline of coral reefs necessitates targeting management solutions that can sustain reefs and the livelihoods of the people who depend on them. However, little is known about the context in which different reef management tools can help to achieve multiple social and ecological goals. Because of nonlinearities in the likelihood of achieving combined fisheries, ecological function, and biodiversity goals along a gradient of human pressure, relatively small changes in the context in which management is implemented could have substantial impacts on whether these goals are likely to be met. Critically, management can provide substantial conservation benefits to most reefs for fisheries and ecological function, but not biodiversity goals, given their degraded state and the levels of human pressure they face.

Recovery when you are on your own: Slow population responses in an isolated marine reserve.

Geographic isolation is an important yet underappreciated factor affecting marine reserve performance. Isolation, in combination with other factors, may preclude recruit subsidies, thus slowing recovery when base populations are small and causing a mismatch between performance and stakeholder expectations. Mona Island is a small, oceanic island located within a partial biogeographic barrier-44 km from the Puerto Rico shelf. We investigated if Mona Island's no-take zone (MNTZ), the largest in the U.S. Caribbean, was successful in increasing mean size and density of a suite of snapper and grouper species 14 years after designation. The La Parguera Natural Reserve (LPNR) was chosen for evaluation of temporal trends at a fished location. Despite indications of fishing within the no-take area, a reserve effect at Mona Island was evidenced from increasing mean sizes and densities of some taxa and mean total density 36% greater relative to 2005. However, the largest predatory species remained rare at Mona, preventing meaningful analysis of population trends. In the LPNR, most commercial species (e.g., Lutjanus synagris, Lutjanus apodus, Lutjanus mahogoni) did not change significantly in biomass or abundance, but some (Ocyurus chrysurus, Lachnolaimus maximus), increased in abundance owing to strong recent recruitment. This study documents slow recovery in the MNTZ that is limited to smaller sized species, highlighting both the need for better compliance and the substantial recovery time required by commercially valuable, coral reef fishes in isolated marine reserves.

Differences in rocky reef habitats related to human disturbances across a latitudinal gradient.

This study tested for differences in the composition of intertidal and shallow subtidal rocky reef habitats subjected to a range of human pressures across ∼1000 km of coastline in New South Wales, Australia over 5 years. Percentage covers of habitats were sampled using aerial photography and a large grain size (20 m2 intertidal; 800 m2 subtidal) in a nested hierarchical design. Results were consistent with anthropogenic impacts on habitat structure only around estuaries with the most heavily urbanised or agriculturally-intense catchments. The most convincing relationships documented here related to environmental variables such as SST, latitude, reef width and proximity to large estuaries irrespective of human disturbance levels. Moreover, there were suggestions that any influences of estuarine waters (be they anthropogenic or natural) on reef assemblages could potentially extend 10s of kilometres from major estuaries. In general, our results supported those of studies that utilised smaller grain sizes (greatest variability often at smallest spatial scales), but we found that variability over scales of 100s of km can be similar to or greater than variability over scales of 10s of metres.

Large-scale spatial distribution patterns of gastropod assemblages in rocky shores.

Gastropod assemblages from nearshore rocky habitats were studied over large spatial scales to (1) describe broad-scale patterns in assemblage composition, including patterns by feeding modes, (2) identify latitudinal pattern of biodiversity, i.e., richness and abundance of gastropods and/or regional hotspots, and (3) identify potential environmental and anthropogenic drivers of these assemblages. Gastropods were sampled from 45 sites distributed within 12 Large Marine Ecosystem regions (LME) following the NaGISA (Natural Geography in Shore Areas) standard protocol (www.nagisa.coml.org). A total of 393 gastropod taxa from 87 families were collected. Eight of these families (9.2%) appeared in four or more different LMEs. Among these, the Littorinidae was the most widely distributed (8 LMEs) followed by the Trochidae and the Columbellidae (6 LMEs). In all regions, assemblages were dominated by few species, the most diverse and abundant of which were herbivores. No latitudinal gradients were evident in relation to species richness or densities among sampling sites. Highest diversity was found in the Mediterranean and in the Gulf of Alaska, while highest densities were found at different latitudes and represented by few species within one genus (e.g. Afrolittorina in the Agulhas Current, Littorina in the Scotian Shelf, and Lacuna in the Gulf of Alaska). No significant correlation was found between species composition and environmental variables (r≤0.355, p>0.05). Contributing variables to this low correlation included invasive species, inorganic pollution, SST anomalies, and chlorophyll-a anomalies. Despite data limitations in this study which restrict conclusions in a global context, this work represents the first effort to sample gastropod biodiversity on rocky shores using a standardized protocol across a wide scale. Our results will generate more work to build global databases allowing for large-scale diversity comparisons of rocky intertidal assemblages.

Aspects of benthic decapod diversity and distribution from rocky nearshore habitat at geographically widely dispersed sites.

Relationships of diversity, distribution and abundance of benthic decapods in intertidal and shallow subtidal waters to 10 m depth are explored based on data obtained using a standardized protocol of globally-distributed samples. Results indicate that decapod species richness overall is low within the nearshore, typically ranging from one to six taxa per site (mean = 4.5). Regionally the Gulf of Alaska decapod crustacean community structure was distinguishable by depth, multivariate analysis indicating increasing change with depth, where assemblages of the high and mid tide, low tide and 1 m, and 5 and 10 m strata formed three distinct groups. Univariate analysis showed species richness increasing from the high intertidal zone to 1 m subtidally, with distinct depth preferences among the 23 species. A similar depth trend but with peak richness at 5 m was observed when all global data were combined. Analysis of latitudinal trends, confined by data limitations, was equivocal on a global scale. While significant latitudinal differences existed in community structure among ecoregions, a semi-linear trend in changing community structure from the Arctic to lower latitudes did not hold when including tropical results. Among boreal regions the Canadian Atlantic was relatively species poor compared to the Gulf of Alaska, whereas the Caribbean and Sea of Japan appeared to be species hot spots. While species poor, samples from the Canadian Atlantic were the most diverse at the higher infraordinal level. Linking 11 environmental variables available for all sites to the best fit family-based biotic pattern showed a significant relationship, with the single best explanatory variable being the level of organic pollution and the best combination overall being organic pollution and primary productivity. While data limitations restrict conclusions in a global context, results are seen as a first-cut contribution useful in generating discussion and more in-depth work in the still poorly understood field of biodiversity distribution.

Spatial relationships between polychaete assemblages and environmental variables over broad geographical scales.

This study examined spatial relationships between rocky shore polychaete assemblages and environmental variables over broad geographical scales, using a database compiled within the Census of Marine Life NaGISA (Natural Geography In Shore Areas) research program. The database consisted of abundance measures of polychaetes classified at the genus and family levels for 74 and 93 sites, respectively, from nine geographic regions. We tested the general hypothesis that the set of environmental variables emerging as potentially important drivers of variation in polychaete assemblages depend on the spatial scale considered. Through Moran's eigenvector maps we indentified three submodels reflecting spatial relationships among sampling sites at intercontinental (>10,000 km), continental (1000-5000 km) and regional (20-500 km) scales. Using redundancy analysis we found that most environmental variables contributed to explain a large and significant proportion of variation of the intercontinental submodel both for genera and families (54% and 53%, respectively). A subset of these variables, organic pollution, inorganic pollution, primary productivity and nutrient contamination was also significantly related to spatial variation at the continental scale, explaining 25% and 32% of the variance at the genus and family levels, respectively. These variables should therefore be preferably considered when forecasting large-scale spatial patterns of polychaete assemblages in relation to ongoing or predicted changes in environmental conditions. None of the variables considered in this study were significantly related to the regional submodel.

Marine biodiversity in the Caribbean: regional estimates and distribution patterns.

This paper provides an analysis of the distribution patterns of marine biodiversity and summarizes the major activities of the Census of Marine Life program in the Caribbean region. The coastal Caribbean region is a large marine ecosystem (LME) characterized by coral reefs, mangroves, and seagrasses, but including other environments, such as sandy beaches and rocky shores. These tropical ecosystems incorporate a high diversity of associated flora and fauna, and the nations that border the Caribbean collectively encompass a major global marine biodiversity hot spot. We analyze the state of knowledge of marine biodiversity based on the geographic distribution of georeferenced species records and regional taxonomic lists. A total of 12,046 marine species are reported in this paper for the Caribbean region. These include representatives from 31 animal phyla, two plant phyla, one group of Chromista, and three groups of Protoctista. Sampling effort has been greatest in shallow, nearshore waters, where there is relatively good coverage of species records; offshore and deep environments have been less studied. Additionally, we found that the currently accepted classification of marine ecoregions of the Caribbean did not apply for the benthic distributions of five relatively well known taxonomic groups. Coastal species richness tends to concentrate along the Antillean arc (Cuba to the southernmost Antilles) and the northern coast of South America (Venezuela-Colombia), while no pattern can be observed in the deep sea with the available data. Several factors make it impossible to determine the extent to which these distribution patterns accurately reflect the true situation for marine biodiversity in general: (1) highly localized concentrations of collecting effort and a lack of collecting in many areas and ecosystems, (2) high variability among collecting methods, (3) limited taxonomic expertise for many groups, and (4) differing levels of activity in the study of different taxa.

Patterns of spatial variation of assemblages associated with intertidal rocky shores: a global perspective.

Assemblages associated with intertidal rocky shores were examined for large scale distribution patterns with specific emphasis on identifying latitudinal trends of species richness and taxonomic distinctiveness. Seventy-two sites distributed around the globe were evaluated following the standardized sampling protocol of the Census of Marine Life NaGISA project (www.nagisa.coml.org). There were no clear patterns of standardized estimators of species richness along latitudinal gradients or among Large Marine Ecosystems (LMEs); however, a strong latitudinal gradient in taxonomic composition (i.e., proportion of different taxonomic groups in a given sample) was observed. Environmental variables related to natural influences were strongly related to the distribution patterns of the assemblages on the LME scale, particularly photoperiod, sea surface temperature (SST) and rainfall. In contrast, no environmental variables directly associated with human influences (with the exception of the inorganic pollution index) were related to assemblage patterns among LMEs. Correlations of the natural assemblages with either latitudinal gradients or environmental variables were equally strong suggesting that neither neutral models nor models based solely on environmental variables sufficiently explain spatial variation of these assemblages at a global scale. Despite the data shortcomings in this study (e.g., unbalanced sample distribution), we show the importance of generating biological global databases for the use in large-scale diversity comparisons of rocky intertidal assemblages to stimulate continued sampling and analyses.

Large-scale spatial distribution patterns of echinoderms in nearshore rocky habitats.

This study examined echinoderm assemblages from nearshore rocky habitats for large-scale distribution patterns with specific emphasis on identifying latitudinal trends and large regional hotspots. Echinoderms were sampled from 76 globally-distributed sites within 12 ecoregions, following the standardized sampling protocol of the Census of Marine Life NaGISA project (www.nagisa.coml.org). Sample-based species richness was overall low (<1-5 species per site), with a total of 32 asteroid, 18 echinoid, 21 ophiuroid, and 15 holothuroid species. Abundance and species richness in intertidal assemblages sampled with visual methods (organisms >2 cm in 1 m(2) quadrats) was highest in the Caribbean ecoregions and echinoids dominated these assemblages with an average of 5 ind m(-2). In contrast, intertidal echinoderm assemblages collected from clearings of 0.0625 m(2) quadrats had the highest abundance and richness in the Northeast Pacific ecoregions where asteroids and holothurians dominated with an average of 14 ind 0.0625 m(-2). Distinct latitudinal trends existed for abundance and richness in intertidal assemblages with declines from peaks at high northern latitudes. No latitudinal trends were found for subtidal echinoderm assemblages with either sampling technique. Latitudinal gradients appear to be superseded by regional diversity hotspots. In these hotspots echinoderm assemblages may be driven by local and regional processes, such as overall productivity and evolutionary history. We also tested a set of 14 environmental variables (six natural and eight anthropogenic) as potential drivers of echinoderm assemblages by ecoregions. The natural variables of salinity, sea-surface temperature, chlorophyll a, and primary productivity were strongly correlated with echinoderm assemblages; the anthropogenic variables of inorganic pollution and nutrient contamination also contributed to correlations. Our results indicate that nearshore echinoderm assemblages appear to be shaped by a network of environmental and ecological processes, and by the differing responses of various echinoderm taxa, making generalizations about the patterns of nearshore rocky habitat echinoderm assemblages difficult.

Current patterns of macroalgal diversity and biomass in northern hemisphere rocky shores.

Latitudinal gradients in species abundance and diversity have been postulated for nearshore taxa but few analyses have been done over sufficiently broad geographic scales incorporating various nearshore depth strata to empirically test these gradients. Typically, gradients are based on literature reviews and species lists and have focused on alpha diversity across the entire nearshore zone. No studies have used a standardized protocol in the field to examine species density among sites across a large spatial scale while also focusing on particular depth strata. The present research used field collected samples in the northern hemisphere to explore the relationships between macroalgal species density and biomass along intertidal heights and subtidal depths and latitude. Results indicated no overall correlations between either estimates of species density or biomass with latitude, although the highest numbers of both were found at mid-latitudes. However, when strata were examined separately, significant positive correlations were found for both species numbers and biomass at particular strata, namely the intertidal ones. While the data presented in this paper have some limitations, we show that latitudinal macroalgal trends in species density and biomass do exist for some strata in the northern hemisphere with more taxa and biomass at higher latitudes.

Inventory of ascidians (Tunicata, Ascidiacea) from the National Park La Restinga, Isla Margarita, Venezuela / Inventário de ascídias (Tunicata, Ascidiacea) do Parque Nacional La Restinga, Isla Margarita, Venezuela

Although ascidians form a conspicuous part of sessile assemblages in the Caribbean, no specialized inventories have been developed in Venezuela, except for a list of 15 species reported from Margarita Island (1984). Here we present the results of a taxonomic workshop held in the Universidad de Oriente, Boca del Rio, Margarita Island during April 20-25 of 2009, sponsored by the NaGISA-Caribbean Sea program. La Restinga National park was surveyed and we found 29 species belonging to 19 genera and 10 families. The most abundant colonial species were Clavelina oblonga, Aplidium accarense, Polyclinum constellatum, Distaplia bermudensis, Symplegma rubra, S. brakenhielmi, Botrylloides nigrum and Ecteinascidia turbinata. Among the solitary ascidians Phallusia nigra, Ascidia curvata, Microcosmus exasperatus, Styela canopus, Styela sp.1 and Styela sp. 2 were the most abundant. The ascidian diversity in this lagoon is one of the highest for similar habitats in the Caribbean and seven species are new registers for Venezuela (Ascidia curvata, Ecteinascidia styeloides, Aplidium accarense, Distaplia stylifera, Trididemnum orbiculatum, Symplegma rubra, and S. brakenhielmi). One point of concern for the conservation of la Restinga National Park is the presence of possible introduced species, and some management procedures are suggested.

Apesar das ascídias formarem uma parte conspícua das comunidades incrustantes no Caribe, nenhum inventário especializado foi realizado na Venezuela, com exceção de uma lista de 15 espécies registradas na Ilha Margarita (1984). Aqui apresentamos os resultados de um workshop de taxonomia realizado na Universidad de Oriente, Boca del Rio, Margarita Island em 20-25 de abril de 2009, como parte do programa NaGISA-Caribbean Sea program. O Parque Nacional La Restinga foi estudado e encontradas 29 espécies pertencendo a 19 gêneros e 10 famílias. As espécies coloniais mais abundantes foram Clavelina oblonga, Aplidium accarense, Polyclinum constellatum, Distaplia bermudensis, Symplegma rubra, S. brakenhielmi, Botrylloides nigrum e Ecteinascidia turbinata. Entre as espécies solitárias, Phallusia nigra, Ascidia curvata, Microcosmus exasperatus, Styela canopus, Styela sp.1 e Styela sp. 2 foram as mais abundantes. A diversidade de ascídias nesta laguna é uma das maiores neste tipo de habitat no Caribe e sete espécies representam novos registros para a Venezuela (Ascidia curvata, Ecteinascidia styeloides, Aplidium accarense, Distaplia stylifera, Trididemnum orbiculatum, Symplegma rubra, e S. brakenhielmi). Uma questão de preocupação para a conservação do Parque Nacional La Restinga é a presença de possíveis espécies exóticas, portanto algumas medidas de manejo são sugeridas.