Distance decay 2.0 - A global synthesis of taxonomic and functional turnover in ecological communities.

Aim: Understanding the variation in community composition and species abundances (i.e., ß-diversity) is at the heart of community ecology. A common approach to examine ß-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments. Location: Global. Time period: 1990 to present. Major taxa studied: From diatoms to mammals. Method: We measured the strength of the decay using ranked Mantel tests (Mantel r) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features. Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances. Main conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments.

Drivers of ecological effectiveness of marine protected areas: A meta-analytic approach from the Southwestern Atlantic Ocean (Brazil).

With the rapid global increase in the number and extent of marine protected areas (MPAs), there is a need for methods that enable an assessment of their actual contribution to biodiversity conservation. In Brazil, where MPAs have been designated to replenish biodiversity, there is a lack of regional-scale analysis of MPA impacts and the factors related to positive ecological change. This study aims to quantify the magnitude of the ecological effects of Brazilian MPAs and test whether some study and MPA characteristics (e.g., taxonomic group studied, exploitation level of species, MPA area, protection time, management effectiveness, level of connectedness, etc.) were underlying factors associated with their performance. We conducted a structured search in a database of scientific articles, selecting comparative studies of direct biodiversity metrics inside and outside MPAs offering different protection levels (i.e., fully- or partially-protected MPAs) or within MPAs with distinct zones. We then carried out a meta-analysis based on 424 observations found in 18 articles. Averaged across all studies, we found that MPAs had a 17% increase in the abundance of species, length of individuals, and community diversity. When compared to open-access areas, fully-protected MPAs increased biodiversity by 45%. However, MPAs offering partial protection had variable effects, ranging from significant positive to significant negative effects. MPA effects depended on the taxonomic group and exploitation level of species, with the strongest positive effects seen on exploited fish species and benthic invertebrates. Partially-protected MPAs that reported strong positive effects required long time of protection (>15years) and high level of connectivity. Conversely, fully-protected MPAs (i.e., no-take ones) could be effective even when small, under intense fishing pressure in their surroundings, and regardless of their level of connectivity. We used the Brazilian MPAs as a case study, but these results can contribute to a more comprehensive assessment of the association between ecological impacts of MPAs and drivers of conservation success, and offer key information to consolidate MPA networks that sustain biodiversity.

A safe haven for potential reproductive aggregations of the critically endangered Brazilian guitarfish (Pseudobatos horkelii).

Brazilian endemic batoid elasmobranch populations have declined dramatically in the past 40 years due to anthropic activities (e.g., overfishing). The Brazilian guitarfish, Pseudobatos horkelii, included in the IUCN red list of endangered species [Critically Endangered (CR)], has been captured as by-catch by trawling fishing boats to the edge of extinction. Despite governmental conservation initiatives, the species is still caught and commercialized along the Brazilian coast. In this study, the authors report three rare aggregation events for the Brazilian coast of P. horkelii, inside the only nearshore no-entry Brazilian marine protected area. Strategies for its protection are also discussed.

Trophic interactions will expand geographically but be less intense as oceans warm.

Interactions among species are likely to change geographically due to climate-driven species range shifts and in intensity due to physiological responses to increasing temperatures. Marine ectotherms experience temperatures closer to their upper thermal limits due to the paucity of temporary thermal refugia compared to those available to terrestrial organisms. Thermal limits of marine ectotherms also vary among species and trophic levels, making their trophic interactions more prone to changes as oceans warm. We assessed how temperature affects reef fish trophic interactions in the Western Atlantic and modeled projections of changes in fish occurrence, biomass, and feeding intensity across latitudes due to climate change. Under ocean warming, tropical reefs will experience diminished trophic interactions, particularly herbivory and invertivory, potentially reinforcing algal dominance in this region. Tropicalization events are more likely to occur in the northern hemisphere, where feeding by tropical herbivores is predicted to expand from the northern Caribbean to extratropical reefs. Conversely, feeding by omnivores is predicted to decrease in this area with minor increases in the Caribbean and southern Brazil. Feeding by invertivores declines across all latitudes in future predictions, jeopardizing a critical trophic link. Most changes are predicted to occur by 2050 and can significantly affect ecosystem functioning, causing dominance shifts and the rise of novel ecosystems.

Spatiotemporal variations in density and biomass of rocky reef fish in a biogeographic climatic transition zone: trends over 9 years, inside and outside the only nearshore no-take marine-protected area on the southern Brazilian coast.

Biogeographical transition zones are important areas to investigate evolutionary ecological questions, but long-term population monitoring is needed to better understand ecological processes that govern population variations in such edge environments. The southernmost Brazilian rocky reefs are the southern limit of distribution for 96% of the tropical ichthyofauna of the western Atlantic. The Arvoredo Marine Biological Reserve is the only nearshore no-take marine-protected area (MPA) located in this transition zone. The main aim was to investigate how the populations of rocky reef fish species vary in density and biomass in space and over time, inside and outside the Arvoredo MPA. This study presents results based on a 9 year (2008-2017) underwater visual census monitoring study to evaluate the density and biomass of key fish species. Variations in density and biomass were detected for most species. Factors and mechanisms that may have influenced spatial variation are habitat structural complexity and protection from fisheries. Temporal variations, otherwise, may have been influenced by species proximity to their distributional limit, in synergy with density-dependent mechanisms and stochastic winter temperature oscillations. The MPAs harbour higher density and biomass for most species. Nonetheless, a prominent temporal decline in the recruitment of Epinephelus marginatus calls into question the continuous effectiveness of the MPA.

Population expansion of the invasive Pomacentridae Chromis limbata (Valenciennes, 1833) in southern Brazilian coast: long-term monitoring, fundamental niche availability and new records.

Human-mediated species invasions are recognized as a leading cause of global biotic homogenization and extinction. Studies on colonization events since early stages, establishment of new populations and range extension are scarce because of their rarity, difficult detection and monitoring. Chromis limbata is a reef-associated and non-migratory marine fish from the family Pomacentridae found in depths ranging between 3 and 45 m. The original distribution of the species encompassed exclusively the eastern Atlantic, including the Azores, Madeira and the Canary Islands. It is also commonly reported from West Africa between Senegal and Pointe Noire, Congo. In 2008, vagrant individuals of C. limbata were recorded off the east coast of Santa Catarina Island, South Brazil (27° 41' 44″ S, 48° 27' 53″ W). This study evaluated the increasing densities of C. limbata populations in Santa Catarina State shoreline. Two recent expansions, northwards to São Paulo State and southwards to Rio Grande do Sul State, are discussed, and a niche model of maximum entropy (MaxEnt) was performed to evaluate suitable C. limbata habitats. Brazilian populations are established and significantly increasing in most sites where the species has been detected. The distributional boundaries predicted by the model are clearly wider than their known range of occurrence, evidencing environmental suitability in both hemispheres from areas where the species still does not occur. Ecological processes such as competition, predation and specially habitat selectivity may regulate their populations and overall distribution range. A long-term monitoring programme and population genetics studies are necessary for a better understanding of this invasion and its consequences to natural communities.

An updated phylogeny of the redlip blenny genus Ophioblennius.

An updated molecular phylogeny of the blenny genus Ophioblennius, with a focus on two geographically disjunct morphotypes observed in Brazil, is presented. The analyses showed that specimens from the north-eastern Brazilian coast are the endemic redlip blenny Ophioblennius trinitatis, but specimens from the southern Brazilian coast are conspecific to an undescribed east Atlantic Ocean (Gulf of Guinea) species, previously unknown in Brazil. Possible explanations for this geographical pattern include: natural larval dispersal and rafting across the Atlantic; an unknown ecological attribute that enabled this species to colonize southern Brazil; oil platforms as introduction vectors.

Functional over-redundancy and high functional vulnerability in global fish faunas on tropical reefs.

When tropical systems lose species, they are often assumed to be buffered against declines in functional diversity by the ability of the species-rich biota to display high functional redundancy: i.e., a high number of species performing similar functions. We tested this hypothesis using a ninefold richness gradient in global fish faunas on tropical reefs encompassing 6,316 species distributed among 646 functional entities (FEs): i.e., unique combinations of functional traits. We found that the highest functional redundancy is located in the Central Indo-Pacific with a mean of 7.9 species per FE. However, this overall level of redundancy is disproportionately packed into few FEs, a pattern termed functional over-redundancy (FOR). For instance, the most speciose FE in the Central Indo-Pacific contains 222 species (out of 3,689) whereas 38% of FEs (180 out of 468) have no functional insurance with only one species. Surprisingly, the level of FOR is consistent across the six fish faunas, meaning that, whatever the richness, over a third of the species may still be in overrepresented FEs whereas more than one third of the FEs are left without insurance, these levels all being significantly higher than expected by chance. Thus, our study shows that, even in high-diversity systems, such as tropical reefs, functional diversity remains highly vulnerable to species loss. Although further investigations are needed to specifically address the influence of redundant vs. vulnerable FEs on ecosystem functioning, our results suggest that the promised benefits from tropical biodiversity may not be as strong as previously thought.

Adult and larval traits as determinants of geographic range size among tropical reef fishes.

Most marine organisms disperse via ocean currents as larvae, so it is often assumed that larval-stage duration is the primary determinant of geographic range size. However, empirical tests of this relationship have yielded mixed results, and alternative hypotheses have rarely been considered. Here we assess the relative influence of adult and larval-traits on geographic range size using a global dataset encompassing 590 species of tropical reef fishes in 47 families, the largest compilation of such data to date for any marine group. We analyze this database using linear mixed-effect models to control for phylogeny and geographical limits on range size. Our analysis indicates that three adult traits likely to affect the capacity of new colonizers to survive and establish reproductive populations (body size, schooling behavior, and nocturnal activity) are equal or better predictors of geographic range size than pelagic larval duration. We conclude that adult life-history traits that affect the postdispersal persistence of new populations are primary determinants of successful range extension and, consequently, of geographic range size among tropical reef fishes.

Global mismatch between species richness and vulnerability of reef fish assemblages.

The impact of anthropogenic activity on ecosystems has highlighted the need to move beyond the biogeographical delineation of species richness patterns to understanding the vulnerability of species assemblages, including the functional components that are linked to the processes they support. We developed a decision theory framework to quantitatively assess the global taxonomic and functional vulnerability of fish assemblages on tropical reefs using a combination of sensitivity to species loss, exposure to threats and extent of protection. Fish assemblages with high taxonomic and functional sensitivity are often exposed to threats but are largely missed by the global network of marine protected areas. We found that areas of high species richness spatially mismatch areas of high taxonomic and functional vulnerability. Nevertheless, there is strong spatial match between taxonomic and functional vulnerabilities suggesting a potential win-win conservation-ecosystem service strategy if more protection is set in these locations.

Thermal tolerance as a driver of reef fish community structure at the isolated tropical Mid-Atlantic Ridge Islands.

Reef fish communities are shaped by historical and ecological factors, including abiotic and biotic mechanisms at different spatial scales, determining species composition, abundance and biomass. The oceanic islands in the Mid-Atlantic Ridge (St. Peter and St. Paul's Archipelago - SPSPA, Ascension, and St. Helena), exhibiting differences in community structure along a 14-degree latitudinal and a 10 °C thermal gradient. We investigate the influence of sea surface temperature, area, age, isolation and phosphate on reef fish community structures. Reef fish trophic structure varies significantly across the islands, with planktivores and herbivore-detritivores showing the highest abundances in SPSPA and Ascension, while less abundant in St. Helena, aligning with the thermal gradient. Variations in reef fish community structures were predominantly influenced by thermal regimes, corroborating the expansion of species' thermal niche breadth at higher latitudes and lower temperatures. This study highlights that in addition to biogeographic factors, temperature is pivotal on shaping oceanic island reef fish community structure.

Turbidity shapes shallow Southwestern Atlantic benthic reef communities.

Southwestern Atlantic reefs (Brazilian Province) occur along a broad latitudinal range (∼5°N-27°S) and under varied environmental conditions. We combined large-scale benthic cover and environmental data into uni- and multivariate regression tree analyses to identify unique shallow (<30 m) benthic reef communities and their environmental drivers along the Brazilian Province. Turbidity was the leading environmental driver of benthic reef communities, with the occurrence of two main groups: clear-water (dominated by fleshy macroalgae) and turbid (dominated by turf algae). Seven out of 14 scleractinian coral species were more abundant in the turbid group, thus corroborating the photophobic nature of some Brazilian corals. The most abundant scleractinian in Brazil (Montastraea cavernosa), largely dominated (71-93% of total coral cover) both, the shallow turbid and deeper clear-water reefs. Because these habitat types are widely recognized as potential climate refuges, local threats (e.g. pollution, overfishing) should be averted.

TimeFISH: Long-term assessment of reef fish assemblages in a transition zone in the Southwestern Atlantic.

The TimeFISH database provides the first public time-series dataset on reef fish assemblages in the southwestern Atlantic (SWA), comprising 15 years of data (2007-2022) based on standardized Underwater Visual Censuses (UVCs). The rocky reefs covered by our dataset are influenced by pronounced seasonal cycles of ocean temperatures with warm tropical waters from the Brazil Current in the summer (~27°C) and colder waters from the La Plata River Plume discharge and upwelling from the South Atlantic Central Water in the winter (~18°C). These oceanographic conditions characterize this area as the southernmost tropical-subtropical climatic transition zone in the Atlantic Ocean. As a result, reef fish assemblages are comprised of both tropical and subtropical species. All records included in TimeFISH were collected using UVCs, a nondestructive method that allows the estimation of fish species richness, abundance, and body size distributions. UVCs were performed through 40 m2 belt transects by scuba diving in nine locations along the southern Brazilian coast (25-29°S). Four of these locations lie within the boundaries of the no-entry Arvoredo Marine Biological Reserve, where fishing and recreational activities are forbidden, and the remaining locations are unprotected from these activities. During each belt transect, a diver swam at a constant depth above and parallel to the reef, identifying fish species, counting the number of individuals, and estimating the total body length (Lt in cm) of all detected individuals. All fish individuals in the water column (up to 2 m above the substratum) and at the bottom were targeted. In total, 202,965 individuals belonging to 163 reef fish species and 53 families were recorded across 1857 UVCs. All survey campaigns were funded by either public or mixed capital (private-public) sources, including seven grants from the Brazilian federal and Santa Catarina state governments. Part of the data has already been used in multiple MS.c. and Ph.D. theses and scientific articles. TimeFISH represents an important contribution for future studies aiming to examine temporal and spatial variations of reef fish assemblages in transition zones. No copyright restrictions apply to the use of this data set, other than citing this publication.

Ecological traits influencing range expansion across large oceanic dispersal barriers: insights from tropical Atlantic reef fishes.

How do biogeographically different provinces arise in response to oceanic barriers to dispersal? Here, we analyse how traits related to the pelagic dispersal and adult biology of 985 tropical reef fish species correlate with their establishing populations on both sides of two Atlantic marine barriers: the Mid-Atlantic Barrier (MAB) and the Amazon-Orinoco Plume (AOP). Generalized linear mixed-effects models indicate that predictors for successful barrier crossing are the ability to raft with flotsam for the deep-water MAB, non-reef habitat usage for the freshwater and sediment-rich AOP, and large adult-size and large latitudinal-range for both barriers. Variation in larval-development mode, often thought to be broadly related to larval-dispersal potential, is not a significant predictor in either case. Many more species of greater taxonomic diversity cross the AOP than the MAB. Rafters readily cross both barriers but represent a much smaller proportion of AOP crossers than MAB crossers. Successful establishment after crossing both barriers may be facilitated by broad environmental tolerance associated with large body size and wide latitudinal-range. These results highlight the need to look beyond larval-dispersal potential and assess adult-biology traits when assessing determinants of successful movements across marine barriers.

Low functional vulnerability of fish assemblages to coral loss in Southwestern Atlantic marginal reefs.

Marginal reefs sustain coral assemblages under conditions considered suboptimal for most corals, resulting in low coral abundance. These reefs are inhabited by numerous fishes with a generally unknown degree of association with corals that might lead to the assumption that corals play minor roles in determining fish occurrence, when corals could be actually sustaining diverse and resilient assemblages. Using site-occupancy models fitted to data of 113 reef fish species of different life stages (adults and juveniles) from 36 reefs distributed across the Southwestern Atlantic (0.87-27.6°S) we first assessed fish assemblage's response to coral and turf algal cover, and identified coral-associated fish. Then, we simulated the loss of coral-associated fishes and contrasted it with random losses, providing inferences on the resilience of fish assemblage's functional trait space to species loss. The entire fish assemblage responded more positively to coral than to turf algae, with 42 (37%) species being identified as coral-associated fish. The simulated loss of coral-associated fish reduced up to 5% the functional trait space and was not different from the random loss. These results reveal that marginal reefs of Southwestern Atlantic reefs host resilient fish assemblages that might preserve fundamental ecological functions and ecosystem services even with coral declines.

Biological trade-offs underpin coral reef ecosystem functioning.

Human impact increasingly alters global ecosystems, often reducing biodiversity and disrupting the provision of essential ecosystem services to humanity. Therefore, preserving ecosystem functioning is a critical challenge of the twenty-first century. Coral reefs are declining worldwide due to the pervasive effects of climate change and intensive fishing, and although research on coral reef ecosystem functioning has gained momentum, most studies rely on simplified proxies, such as fish biomass. This lack of quantitative assessments of multiple process-based ecosystem functions hinders local and regional conservation efforts. Here we combine global coral reef fish community surveys and bioenergetic models to quantify five key ecosystem functions mediated by coral reef fishes. We show that functions exhibit critical trade-offs driven by varying community structures, such that no community can maximize all functions. Furthermore, functions are locally dominated by few species, but the identity of dominant species substantially varies at the global scale. In fact, half of the 1,110 species in our dataset are functionally dominant in at least one location. Our results reinforce the need for a nuanced, locally tailored approach to coral reef conservation that considers multiple ecological functions beyond the effect of standing stock biomass.

Long-term monitoring projects of Brazilian marine and coastal ecosystems.

Biodiversity assessment is a mandatory task for sustainable and adaptive management for the next decade, and long-term ecological monitoring programs are a cornerstone for understanding changes in ecosystems. The Brazilian Long-Term Ecological Research Program (PELD) is an integrated effort model supported by public funds that finance ecological studies at 34 locations. By interviewing and compiling data from project coordinators, we assessed monitoring efforts, targeting biological groups and scientific production from nine PELD projects encompassing coastal lagoons to mesophotic reefs and oceanic islands. Reef environments and fish groups were the most often studied within the long-term projects. PELD projects covered priority areas for conservation but missed sensitive areas close to large cities, as well as underrepresenting ecosystems on the North and Northeast Brazilian coast. Long-term monitoring projects in marine and coastal environments in Brazil are recent (<5 years), not yet integrated as a network, but scientifically productive with considerable relevance for academic and human resources training. Scientific production increased exponentially with project age, despite interruption and shortage of funding during their history. From our diagnosis, we recommend some actions to fill in observed gaps, such as: enhancing projects' collaboration and integration; focusing on priority regions for new projects; broadening the scope of monitored variables; and, maintenance of funding for existing projects.

Molecular phylogenetics and evolution of Holacanthus angelfishes (Pomacanthidae).

The angelfish genus Holacanthus includes seven species in the tropical Eastern Pacific and Atlantic. In this study we performed an analysis on all species, the closely related regal angelfish, and the Cortez angelfish, using four mitochondrial and one nuclear marker. Our results support a monophyletic Holacanthus. The Indo-Pacific regal angelfish, Pygoplytes diacanthus, was found to be the closest relative to Holacanthus. We found a split into two clades with divergences that were consistent with the rise of the Isthmus of Panama. An internally calibrated molecular clock thus placed the origin of Holacanthus to approximately 10.2-7.6 million years ago.

Restructuring of the 'Macaronesia' biogeographic unit: A marine multi-taxon biogeographical approach.

The Azores, Madeira, Selvagens, Canary Islands and Cabo Verde are commonly united under the term "Macaronesia". This study investigates the coherency and validity of Macaronesia as a biogeographic unit using six marine groups with very different dispersal abilities: coastal fishes, echinoderms, gastropod molluscs, brachyuran decapod crustaceans, polychaete annelids, and macroalgae. We found no support for the current concept of Macaronesia as a coherent marine biogeographic unit. All marine groups studied suggest the exclusion of Cabo Verde from the remaining Macaronesian archipelagos and thus, Cabo Verde should be given the status of a biogeographic subprovince within the West African Transition province. We propose to redefine the Lusitanian biogeographical province, in which we include four ecoregions: the South European Atlantic Shelf, the Saharan Upwelling, the Azores, and a new ecoregion herein named Webbnesia, which comprises the archipelagos of Madeira, Selvagens and the Canary Islands.

Molecular ecology, speciation, and evolution of the reef fish genus Anisotremus.

Grunts (family Haemulidae) in the genus Anisotremus comprise 10 described species which occur predominantly on coral reefs and subtropical rocky reefs in the Neotropics of the Tropical Eastern Pacific the Caribbean and adjacent waters. In this study, the phylogenetic relationships for all described species were examined based on one mitochondrial gene (cytochrome b) and one nuclear marker (the first intron of the ribosomal protein S7). Closely related species of the genus Haemulon and the single representative of the genus Genyatremus, G. luteus were also included in the analysis, following the suggestion, based on morphological data, that they may be closely related to, or possibly placed within, Anisotremus. Two sweetlips, genus Plectorhinchus, which belong to the recognized sister subfamily Plectorhynchinae, were used as outgroups. Two species pairs in the ingroup genus Anisotremus are geminate species, which presumably diverged after the rise of the Isthmus of Panama, thus providing an internal molecular clock calibration. This allowed for a comprehensive evolutionary scenario for the genus. Our data were consistent with two independent allopatric speciation events (geminate species), and one likely sympatric speciation event. Our data were also consistent with the placement of Genyatremus and Haemulon within the genus Anisotremus, thus breaking its monophyletic status. As proposed for other coral reef species, Anisotremus seems to have colonized coral reefs from non-coral reef habitats. In addition, ancestral species seem to have been widespread, since species that are currently found in Brazilian and Pacific waters are basal on the phylogenetic tree. Finally, this study adds to the hypothesis of a secondary evolutionary center in the New World.