Vertical structure of reef fish assemblages and light penetration reveal new boundaries of mesophotic ecosystems in the subtropical Southwestern Atlantic.

In the Southwest Atlantic, fisheries and in situ observations suggest that mesophotic reefs occur further south than has been reported in the literature, and a description of these subtropical regions is still lacking. We used Baited Remote Underwater stereo-Videos to explore unrevealed patterns in the vertical structure of fish assemblages on subtropical Atlantic reefs, contrasting shallow and mesophotic habitats. Our data on species turnover and light penetration reveal that in the subtropical Atlantic, the boundaries between shallow and mesophotic habitats occur at depths of 18 m, which is shallower than most previous studies have shown. Generalized additive mixed models identified different environmental and management factors as important predictor variables that explain distribution of fish assemblages' attributes. Besides adding new evidence about mesophotic limits, we described variations in the vertical structure of fish assemblages, providing important information for the description and conservation of mesophotic environments in subtropical regions.

Dominance of photo over chromatic acclimation strategies by habitat-forming mesophotic red algae.

Red coralline algae are the deepest living macroalgae, capable of creating spatially complex reefs from the intertidal to 100+ m depth with global ecological and biogeochemical significance. How these algae maintain photosynthetic function under increasingly limiting light intensity and spectral availability is key to explaining their large depth distribution. Here, we investigated the photo- and chromatic acclimation and morphological change of free-living red coralline algae towards mesophotic depths in the Fernando do Noronha archipelago, Brazil. From 13 to 86 m depth, thalli tended to become smaller and less complex. We observed a dominance of the photo-acclimatory response, characterized by an increase in photosynthetic efficiency and a decrease in maximum electron transport rate. Chromatic acclimation was generally stable across the euphotic-mesophotic transition with no clear depth trend. Taxonomic comparisons suggest these photosynthetic strategies are conserved to at least the Order level. Light saturation necessitated the use of photoprotection to 65 m depth, while optimal light levels were met at 86 m. Changes to the light environment (e.g. reduced water clarity) due to human activities therefore places these mesophotic algae at risk of light limitation, necessitating the importance of maintaining good water quality for the conservation and protection of mesophotic habitats.

Modeling risks in marine protected areas: Mapping of habitats, biodiversity, and cultural ecosystem services in the southernmost atlantic coral reef.

Marine Protected Area (MPA) is a fundamental strategy for the maintenance of ocean ecological processes worldwide and, consequently, their associated ecosystem services. Nevertheless, the quality of the services provided by MPAs, including cultural services such as recreational activities, depends on the effective management of marine habitats and biodiversity. Here, we performed an ecosystemic assessment in reef environments within a subtropical MPA, modeling the potential risks for their habitats and their recreational activities. The Queimada Grande Island (QGI), southeastern Brazil, was used as the model area since this island encompasses a unique and irreplaceable marine habitat, the Southernmost Atlantic coral reef. We firstly assessed and mapped the habitats, the biodiversity, and the recreational activities associated with QGI reefs. Next, we considered different scenarios of management for the modeling risks across the study area. We found that the coral reef and its adjacent habitats, such as the rhodolith bed, make the sheltered face of the island an important area for the provision of the cultural ecosystem services and overlapping uses such as onboard recreational fishing, spearfishing, and recreational diving. This area was also evaluated as the one under the highest risk of impact, considering the current scenario of management. The most successful scenario modeling to reduce these risks was the hypothetical implementation of a 66% reduction of all activities over all QGI habitats. Despite that, the scenario simulating the application of the regulations present in the MPA management plan was enough to reduce almost half the maximum risk value. Therefore, we concluded that to provide a balance among conservation, uses, and the local economy, the application of these regulations is the better management scenario modeled for the study area. Such results provided useful information and tools for local management and decision-making in this singular marine environment, also being an example for mapping ecosystem services and modeling risks in MPAs worldwide.

Effects of the COVID-19 pandemic on SCUBA diving experience in marine protected areas.

Faced with the need for isolation of most people to reduce the transmission of COVID-19, a great concern for the well-being of the population has resurfaced. Considering the numerous benefits of outdoor activities to human health and well-being, we assessed whether the quality of the experience of recreational divers in a marine protected area (MPA) was impacted during the pandemic. We applied a semi-structured questionnaire to divers, collected their socioeconomic profile, preferences, motivations, experiences, and compared the results with those obtained in the previous season. We found that the quality of the recreational diver's experience remained high even during one of the world's biggest health crises. This suggests that the combination of contact with a preserved environment and the adaptation of regulations in favor of visitor safety was enough to stimulate positive experiences. Therefore, we highlight the importance of investing in adaptive management so that MPAs continue to promote ecosystem services such as human health and subjective well-being. Management implications: •Quality of experience for recreational divers remained high and was unaffected after the first major wave of COVID-19 cases.•Through adaptive tourism management, MPAs can promote positive experiences even in challenging contexts.•High impact of environmental attributes on the quality of divers' experience reinforces that threats to ecosystems are social impacts.•Socially unequal access to benefits generated by MPAs may negatively influence population's support for these areas.•Efforts are needed to strengthen the equity of distribution of positive social impacts in MPAs.•Potential to provide positive experiences for users must be seen beyond economic value, but as an investment in the population's subjective well-being.

Levelling-up rhodolith-bed science to address global-scale conservation challenges.

Global marine conservation remains fractured by an imbalance in research efforts and policy actions, limiting progression towards sustainability. Rhodolith beds represent a prime example, as they have ecological importance on a global scale, provide a wealth of ecosystem functions and services, including biodiversity provision and potential climate change mitigation, but remain disproportionately understudied, compared to other coastal ecosystems (tropical coral reefs, kelp forests, mangroves, seagrasses). Although rhodolith beds have gained some recognition, as important and sensitive habitats at national/regional levels during the last decade, there is still a notable lack of information and, consequently, specific conservation efforts. We argue that the lack of information about these habitats, and the significant ecosystem services they provide, is hindering the development of effective conservation measures and limiting wider marine conservation success. This is becoming a pressing issue, considering the multiple severe pressures and threats these habitats are exposed to (e.g., pollution, fishing activities, climate change), which may lead to an erosion of their ecological function and ecosystem services. By synthesizing the current knowledge, we provide arguments to highlight the importance and urgency of levelling-up research efforts focused on rhodolith beds, combating rhodolith bed degradation and avoiding the loss of associated biodiversity, thus ensuring the sustainability of future conservation programs.

Ecosystem engineer morphological traits and taxon identity shape biodiversity across the euphotic-mesophotic transition.

The euphotic-mesophotic transition is characterized by dramatic changes in environmental conditions, which can significantly alter the functioning of ecosystem engineers and the structure of their associated communities. However, the drivers of biodiversity change across the euphotic-mesophotic transition remain unclear. Here, we investigated the mechanisms affecting the biodiversity-supporting potential of free-living red coralline algae-globally important habitat creators-towards mesophotic depths. Across a 73 m depth gradient, we observed a general decline in macrofaunal biodiversity (fauna abundance, taxon richness and alpha diversity), but an increase in beta-diversity (i.e. variation between assemblages) at the deepest site (86 m depth, where light levels were less than 1% surface irradiance). We identified a gradient in abundance decline rather than distinct ecological shifts, driven by a complex interaction between declining light availability, declining size of the coralline algal host individuals and a changing host taxonomy. However, despite abundance declines, high between-assemblage variability at deeper depths allowed biodiversity-supporting potential to be maintained, highlighting their importance as coastal refugia.

Calcium carbonate production in the southernmost subtropical Atlantic coral reef.

Emerging evidence have been supporting the idea that the better known South Atlantic coral reefs (located between 18°S and 24°S) are now essentially senescent structures that have experienced little or no additional vertical reef growth over the past millennia. This has often coincided with a shift to a dominance of non-coral calcifying organisms becoming the main CaCO3 producers in these high latitude and marginal marine settings. Here, we used Calcification Accretion Units (CAUs) and census-based methods to measure non-coral rates of CaCO3 production on the geologically senescent reef and adjacent rhodolith beds within the southernmost subtropical Atlantic reef (i.e., Queimada Grande Reef, QGR). The reef habitat is currently producing CaCO3 at rates of ∼126 g m-2 yr-1. In contrast, fragments of dead corals skeletons deposited adjacent to the reef over the last ∼2000 years are now colonized by crustose coralline red algae. These form a rhodolith bed that produces CaCO3 at rates of 858 g m-2 yr-1. Our results indicate that, whilst not sufficient to promote active net framework accumulation, CaCO3 production by coralline algae and bryozoans on the QGR appears to be sufficient to at least limit net large-scale erosion of the underlying reef structure, allowing the reef structure to persist in a state close to budgetary stasis. Finally, our results are also of relevance for providing insights regarding the balance of CaCO3 production/dissolution/erosion processes in coral reefs, especially in these less understood marginal reefs.

Tropical rhodolith beds are a major and belittled reef fish habitat.

Understanding habitat-level variation in community structure provides an informed basis for natural resources' management. Reef fishes are a major component of tropical marine biodiversity, but their abundance and distribution are poorly assessed beyond conventional SCUBA diving depths. Based on a baited-video survey of fish assemblages in Southwestern Atlantic's most biodiverse region we show that species composition responded mainly to the two major hard-bottom megahabitats (reefs and rhodolith beds) and to the amount of light reaching the bottom. Both megahabitats encompassed typical reef fish assemblages but, unexpectedly, richness in rhodolith beds and reefs was equivalent. The dissimilar fish biomass and trophic structure in reefs and rhodolith beds indicates that these systems function based on contrasting energy pathways, such as the much lower herbivory recorded in the latter. Rhodolith beds, the dominant benthic megahabitat in the tropical Southwestern Atlantic shelf, play an underrated role as fish habitats, and it is critical that they are considered in conservation planning.

A New Species of Amphipod Pariphinotus Kunkel, 1910 (Amphipoda: Phliantidae) from the Southwestern Atlantic.

The genus Pariphinotus Kunkel, 1910 comprises a total of six species around the world, with circumtropical and subtropical distribution, commonly occurring in shallow waters, in soft bottoms and algae, including a previous record of P. seclusus (Shoemaker, 1933) from Brazil. Material examined was collected by SCUBA diving, from rhodolith beds, 12 meters deep, at the Fernando de Noronha Archipelago, off the northeastern Brazilian coast. The new Brazilian species described here can be diagnosed by a unique combination of character states present in other species of the genus. However, it has two exclusive characteristics, distinguishing it from the group: medial apex of pleopod 3 peduncle with two small hook setae, and uropod 1 peduncle not exceeding the apex of telson.

Long-term effects of competition and environmental drivers on the growth of the endangered coral Mussismilia braziliensis (Verril, 1867).

Most coral reefs have recently experienced acute changes in benthic community structure, generally involving dominance shifts from slow-growing hard corals to fast-growing benthic invertebrates and fleshy photosynthesizers. Besides overfishing, increased nutrification and sedimentation are important drivers of this process, which is well documented at landscape scales in the Caribbean and in the Indo-Pacific. However, small-scale processes that occur at the level of individual organisms remain poorly explored. In addition, the generality of coral reef decline models still needs to be verified on the vast realm of turbid-zone reefs. Here, we documented the outcome of interactions between an endangered Brazilian-endemic coral (Mussismilia braziliensis) and its most abundant contacting organisms (turf, cyanobacteria, corals, crustose coralline algae and foliose macroalgae). Our study was based on a long (2006-2016) series of high resolution data (fixed photoquadrats) acquired along a cross-shelf gradient that includes coastal unprotected reefs and offshore protected sites. The study region (Abrolhos Bank) comprises the largest and richest coralline complex in the South Atlantic, and a foremost example of a turbid-zone reef system with low diversity and expressive coral cover. Coral growth was significantly different between reefs. Coral-algae contacts predominated inshore, while cyanobacteria and turf contacts dominated offshore. An overall trend in positive coral growth was detected from 2009 onward in the inshore reef, whereas retraction in live coral tissue was observed offshore during this period. Turbidity (+) and cyanobacteria (-) were the best predictors of coral growth. Complimentary incubation experiments, in which treatments of Symbiodinium spp. from M. braziliensis colonies were subjected to cyanobacterial exudates, showed a negative effect of the exudate on the symbionts, demonstrating that cyanobacteria play an important role in coral tissue necrosis. Negative effects of cyanobacteria on living coral tissue may remain undetected from percent cover estimates gathered at larger spatial scales, as these ephemeral organisms tend to be rapidly replaced by longer-living macroalgae, or complex turf-like consortia. The cross-shelf trend of decreasing turbidity and macroalgae abundance suggests either a direct positive effect of turbidity on coral growth, or an indirect effect related to the higher inshore cover of foliose macroalgae, constraining cyanobacterial abundance. It is unclear whether the higher inshore macroalgal abundance (10-20% of reef cover) is a stable phase related to a long-standing high turbidity background, or a contemporary response to anthropogenic stress. Our results challenge the idea that high macroalgal cover is always associated with compromised coral health, as the baselines for turbid zone reefs may derive sharply from those of coral-dominated reefs that dwell under oligotrophic conditions.

The Farther the Better: Effects of Multiple Environmental Variables on Reef Fish Assemblages along a Distance Gradient from River Influences.

The conservation and management of site-attached assemblages of coastal reefs are particularly challenging because of the tremendous environmental variation that exists at small spatial scales. In this sense, understanding the primary sources of variation in spatial patterns of the biota is fundamental for designing effective conservation policies. We investigated spatial variation in fish assemblages around the windward and leeward sides of coastal islands situated across a gradient of riverine influence (13 km in length). Specifically, relationships between rocky reef fish assemblages and benthic, topographic and physical predictors were assessed. We hypothesized that river induced disturbances may overcome local habitat features in modeling spatial patterns of fish distribution. Fish assemblages varied primarily due to the strong directional gradient of riverine influence (22.6% of the estimated components of variation), followed by topographic complexity (15%), wave exposure (9.9%), and benthic cover (8%). The trophic structure of fish assemblages changed from having a high abundance of invertebrate feeders in macroalgae-dominated reefs close to river mouths to a high proportion of herbivores, planktivores and invertebrate feeder species in reefs with large boulders covered by epilithic algal matrices, as the distance from rivers increased. This gradient led to an increase of 4.5-fold in fish richness and fish trophic group diversity, 11-fold in fish biomass and 10-fold in fish abundance. Our results have implications for the conservation and monitoring of assemblages patchily distributed at small spatial scales. The major role of distance from river influences on fish assemblages rather than benthic cover and topographic complexity suggest that managing land-based activities should be a conservation priority toward reef restoration.

Baseline Assessment of Mesophotic Reefs of the Vitória-Trindade Seamount Chain Based on Water Quality, Microbial Diversity, Benthic Cover and Fish Biomass Data.

Seamounts are considered important sources of biodiversity and minerals. However, their biodiversity and health status are not well understood; therefore, potential conservation problems are unknown. The mesophotic reefs of the Vitória-Trindade Seamount Chain (VTC) were investigated via benthic community and fish surveys, metagenomic and water chemistry analyses, and water microbial abundance estimations. The VTC is a mosaic of reef systems and includes fleshy algae dominated rhodolith beds, crustose coralline algae (CCA) reefs, and turf algae dominated rocky reefs of varying health levels. Macro-carnivores and larger fish presented higher biomass at the CCA reefs (4.4 kg per frame) than in the rhodolith beds and rocky reefs (0.0 to 0.1 kg per frame). A larger number of metagenomic sequences identified as primary producers (e.g., Chlorophyta and Streptophyta) were found at the CCA reefs. However, the rocky reefs contained more diseased corals (>90%) than the CCA reefs (~40%) and rhodolith beds (~10%). Metagenomic analyses indicated a heterotrophic and fast-growing microbiome in rocky reef corals that may possibly lead to unhealthy conditions possibly enhanced by environmental features (e.g. light stress and high loads of labile dissolved organic carbon). VTC mounts represent important hotspots of biodiversity that deserve further conservation actions.

Dynamics of coral reef benthic assemblages of the Abrolhos Bank, eastern Brazil: inferences on natural and anthropogenic drivers.

The Abrolhos Bank (eastern Brazil) encompasses the largest and richest coral reefs of the South Atlantic. Coral reef benthic assemblages of the region were monitored from 2003 to 2008. Two habitats (pinnacles' tops and walls) were sampled per site with 3-10 sites sampled within different reef areas. Different methodologies were applied in two distinct sampling periods: 2003-2005 and 2006-2008. Spatial coverage and taxonomic resolution were lower in the former than in the latter period. Benthic assemblages differed markedly in the smallest spatial scale, with greater differences recorded between habitats. Management regimes and biomass of fish functional groups (roving and territorial herbivores) had minor influences on benthic assemblages. These results suggest that local environmental factors such as light, depth and substrate inclination exert a stronger influence on the structure of benthic assemblages than protection from fishing. Reef walls of unprotected coastal reefs showed highest coral cover values, with a major contribution of Montastraea cavernosa (a sediment resistant species that may benefit from low light levels). An overall negative relationship between fleshy macroalgae and slow-growing reef-building organisms (i.e. scleractinians and crustose calcareous algae) was recorded, suggesting competition between these organisms. The opposite trend (i.e. positive relationships) was recorded for turf algae and the two reef-building organisms, suggesting beneficial interactions and/or co-occurrence mediated by unexplored factors. Turf algae cover increased across the region between 2006 and 2008, while scleractinian cover showed no change. The need of a continued and standardized monitoring program, aimed at understanding drivers of change in community patterns, as well as to subsidize sound adaptive conservation and management measures, is highlighted.

Rhodolith beds in Brazil: a new potential habitat for marine bioprospection

Rhodoliths are the free-living forms of a number of nongeniculate coralline algae. Rhodolith beds are a common feature of subtidal environments and have been recognized as important carbonate producers and paleoenvironmental indicators, as well as recognized as habitat-forming species. The rhodolith structure provides a hard three-dimensional substrate serving as microhabitat for a wide range of biodiversity, including commercially importance species. The largest known latitudinal occurrence range of rhodolith beds ais on the Brazilian coastal shelf from 2°N to 25°S. Despite their importance for the Brazilian benthic communities, only in the last decade sampling efforts allowed a more comprehensive understanding of the beds' distribution, their structure and associated communities, as well as data concerning the influence of environmental factors on rhodolith bed structure and dynamics. In this work, we review the available information on the biodiversity associated with the recently described Brazilian rhodolith beds of the continental shelf and oceanic islands, focusing on the associated organisms with potential for bioprospection research.

Rhodolith beds are major CaCO3 bio-factories in the tropical South West Atlantic.

Rhodoliths are nodules of non-geniculate coralline algae that occur in shallow waters (<150 m depth) subjected to episodic disturbance. Rhodolith beds stand with kelp beds, seagrass meadows, and coralline algal reefs as one of the world's four largest macrophyte-dominated benthic communities. Geographic distribution of rhodolith beds is discontinuous, with large concentrations off Japan, Australia and the Gulf of California, as well as in the Mediterranean, North Atlantic, eastern Caribbean and Brazil. Although there are major gaps in terms of seabed habitat mapping, the largest rhodolith beds are purported to occur off Brazil, where these communities are recorded across a wide latitudinal range (2°N-27°S). To quantify their extent, we carried out an inter-reefal seabed habitat survey on the Abrolhos Shelf (16°50'-19°45'S) off eastern Brazil, and confirmed the most expansive and contiguous rhodolith bed in the world, covering about 20,900 km(2). Distribution, extent, composition and structure of this bed were assessed with side scan sonar, remotely operated vehicles, and SCUBA. The mean rate of CaCO(3) production was estimated from in situ growth assays at 1.07 kg m(-2) yr(-1), with a total production rate of 0.025 Gt yr(-1), comparable to those of the world's largest biogenic CaCO(3) deposits. These gigantic rhodolith beds, of areal extent equivalent to the Great Barrier Reef, Australia, are a critical, yet poorly understood component of the tropical South Atlantic Ocean. Based on the relatively high vulnerability of coralline algae to ocean acidification, these beds are likely to experience a profound restructuring in the coming decades.