The potential for coral reef restoration to mitigate coastal flooding as sea levels rise.

The ability of reefs to protect coastlines from storm-driven flooding hinges on their capacity to keep pace with sea-level rise. Here, we show how and whether coral restoration could achieve the often-cited goal of reversing the impacts of coral-reef degradation to preserve this essential function. We combined coral-growth measurements and carbonate-budget assessments of reef-accretion potential at Buck Island Reef, U.S. Virgin Islands, with hydrodynamic modeling to quantify future coastal flooding under various coral-restoration, sea-level rise, and storm scenarios. Our results provide guidance on how restoration of Acropora palmata, if successful, could mitigate the most extreme impacts of coastal flooding by reversing projected trajectories of reef erosion and allowing reefs to keep pace with the ~0.5 m of sea-level rise expected by 2100 with moderate carbon-emissions reductions. This highlights the potential long-term benefits of pursuing coral-reef restoration alongside climate-change mitigation to support the persistence of essential coral-reef ecosystem services.

Upwelling, climate change, and the shifting geography of coral reef development.

The eastern tropical Pacific is oceanographically unfavorable for coral-reef development. Nevertheless, reefs have persisted there for the last 7000 years. Rates of vertical accretion during the Holocene have been similar in the strong-upwelling Gulf of Panamá (GoP) and the adjacent, weak-upwelling Gulf of Chiriquí (GoC); however, seasonal upwelling in the GoP exacerbated a climate-driven hiatus in reef development in the late Holocene. The situation is now reversed and seasonal upwelling in the GoP currently buffers thermal stress, creating a refuge for coral growth. We developed carbonate budget models to project the capacity of reefs in both gulfs to keep up with future sea-level rise. On average, the GoP had significantly higher net carbonate production rates than the GoC. With an estimated contemporary reef-accretion potential (RAP) of 5.5 mm year-1, reefs in the GoP are projected to be able to keep up with sea-level rise if CO2 emissions are reduced, but not under current emissions trajectories. With an estimated RAP of just 0.3 mm year-1, reefs in the GoC are likely already unable to keep up with contemporary sea-level rise in Panamá (1.4 mm year-1). Whereas the GoP has the potential to support functional reefs in the near-term, our study indicates that their long-term persistence will depend on reduction of greenhouse gases.

Twenty years of change in benthic communities across the Belizean Barrier Reef.

Disease, storms, ocean warming, and pollution have caused the mass mortality of reef-building corals across the Caribbean over the last four decades. Subsequently, stony corals have been replaced by macroalgae, bacterial mats, and invertebrates including soft corals and sponges, causing changes to the functioning of Caribbean reef ecosystems. Here we describe changes in the absolute cover of benthic reef taxa, including corals, gorgonians, sponges, and algae, at 15 fore-reef sites (12-15m depth) across the Belizean Barrier Reef (BBR) from 1997 to 2016. We also tested whether Marine Protected Areas (MPAs), in which fishing was prohibited but likely still occurred, mitigated these changes. Additionally, we determined whether ocean-temperature anomalies (measured via satellite) or local human impacts (estimated using the Human Influence Index, HII) were related to changes in benthic community structure. We observed a reduction in the cover of reef-building corals, including the long-lived, massive corals Orbicella spp. (from 13 to 2%), and an increase in fleshy and corticated macroalgae across most sites. These and other changes to the benthic communities were unaffected by local protection. The covers of hard-coral taxa, including Acropora spp., Montastraea cavernosa, Orbicella spp., and Porites spp., were negatively related to the frequency of ocean-temperature anomalies. Only gorgonian cover was related, negatively, to our metric of the magnitude of local impacts (HII). Our results suggest that benthic communities along the BBR have experienced disturbances that are beyond the capacity of the current management structure to mitigate. We recommend that managers devote greater resources and capacity to enforcing and expanding existing marine protected areas and to mitigating local stressors, and most importantly, that government, industry, and the public act immediately to reduce global carbon emissions.

Climate and the latitudinal limits of subtropical reef development.

Climate plays a central role in coral-reef development, especially in marginal environments. The high-latitude reefs of southeast Florida are currently non-accreting, relict systems with low coral cover. This region also did not support the extensive Late Pleistocene reef development observed in many other locations around the world; however, there is evidence of significant reef building in southeast Florida during the Holocene. Using 146 radiometric ages from reefs extending ~ 120 km along Florida's southeast coast, we test the hypothesis that the latitudinal extent of Holocene reef development in this region was modulated by climatic variability. We demonstrate that although sea-level changes impacted rates of reef accretion and allowed reefs to backstep inshore as new habitats were flooded, sea level was not the ultimate cause of reef demise. Instead, we conclude that climate was the primary driver of the expansion and contraction of Florida's reefs during the Holocene. Reefs grew to 26.7° N in southeast Florida during the relatively warm, stable climate at the beginning of the Holocene Thermal Maximum (HTM) ~ 10,000 years ago, but subsequent cooling and increased frequency of winter cold fronts were associated with the equatorward contraction of reef building. By ~ 7800 years ago, actively accreting reefs only extended to 26.1° N. Reefs further contracted to 25.8° N after 5800 years ago, and by 3000 years ago reef development had terminated throughout southern Florida (24.5-26.7° N). Modern warming is unlikely to simply reverse this trend, however, because the climate of the Anthropocene will be fundamentally different from the HTM. By increasing the frequency and intensity of both warm and cold extreme-weather events, contemporary climate change will instead amplify conditions inimical to reef development in marginal reef environments such as southern Florida, making them more likely to continue to deteriorate than to resume accretion in the future.

Ocean acidification alters properties of the exoskeleton in adult Tanner crabs, Chionoecetes bairdi.

Ocean acidification can affect the ability of calcifying organisms to build and maintain mineralized tissue. In decapod crustaceans, the exoskeleton is a multilayered structure composed of chitin, protein and mineral, predominately magnesian calcite or amorphous calcium carbonate (ACC). We investigated the effects of acidification on the exoskeleton of mature (post-terminal-molt) female southern Tanner crabs, Chionoecetes bairdi Crabs were exposed to one of three pH levels - 8.1, 7.8 or 7.5 - for 2 years. Reduced pH led to a suite of body region-specific effects on the exoskeleton. Microhardness of the claw was 38% lower in crabs at pH 7.5 compared with those at pH 8.1, but carapace microhardness was unaffected by pH. In contrast, reduced pH altered elemental content in the carapace (reduced calcium, increased magnesium), but not the claw. Diminished structural integrity and thinning of the exoskeleton were observed at reduced pH in both body regions; internal erosion of the carapace was present in most crabs at pH 7.5, and the claws of these crabs showed substantial external erosion, with tooth-like denticles nearly or completely worn away. Using infrared spectroscopy, we observed a shift in the phase of calcium carbonate present in the carapace of pH 7.5 crabs: a mix of ACC and calcite was found in the carapace of crabs at pH 8.1, whereas the bulk of calcium carbonate had transformed to calcite in pH 7.5 crabs. With limited capacity for repair, the exoskeleton of long-lived crabs that undergo a terminal molt, such as C. bairdi, may be especially susceptible to ocean acidification.

The timing and causality of ecological shifts on Caribbean reefs.

Caribbean reefs have experienced unprecedented changes in the past four decades. Of great concern is the perceived widespread shift from coral to macroalgal dominance and the question of whether it represents a new, stable equilibrium for coral-reef communities. The primary causes of the shift-grazing pressure (top-down), nutrient loading (bottom-up) or direct coral mortality (side-in)-still remain somewhat controversial in the coral-reef literature. We have attempted to tease out the relative importance of each of these causes. Four insights emerge from our analysis of an early regional dataset of information on the benthic composition of Caribbean reefs spanning the years 1977-2001. First, although three-quarters of reef sites have experienced coral declines concomitant with macroalgal increases, fewer than 10% of the more than 200 sites studied were dominated by macroalgae in 2001, by even the most conservative definition of dominance. Using relative dominance as the threshold, a total of 49 coral-to-macroalgae shifts were detected. This total represents ~35% of all sites that were dominated by coral at the start of their monitoring periods. Four shifts (8.2%) occurred because of coral loss with no change in macroalgal cover, 15 (30.6%) occurred because of macroalgal gain without coral loss, and 30 (61.2%) occurred owing to concomitant coral decline and macroalgal increase. Second, the timing of shifts at the regional scale is most consistent with the side-in model of reef degradation, which invokes coral mortality as a precursor to macroalgal takeover, because more shifts occurred after regional coral-mortality events than expected by chance. Third, instantaneous observations taken at the start and end of the time-series for individual sites showed these reefs existed along a continuum of coral and macroalgal cover. The continuous, broadly negative relationship between coral and macroalgal cover suggests that in some cases coral-to-macroalgae phase shifts may be reversed by removing sources of perturbation or restoring critical components such as the herbivorous sea urchin Diadema antillarum to the system. The five instances in which macroalgal dominance was reversed corroborate the conclusion that macroalgal dominance is not a stable, alternative community state as has been commonly assumed. Fourth, the fact that the loss in regional coral cover and concomitant changes to the benthic community are related to punctuated, discrete events with known causes (i.e. coral disease and bleaching), lends credence to the hypothesis that coral reefs of the Caribbean have been under assault from climate-change-related maladies since the 1970s.

Mechanical Resistance in Decapod Claw Denticles: Contribution of Structure and Composition.

The decapod crustacean exoskeleton is a multi-layered structure composed of chitin-protein fibers embedded with calcium salts. Decapod claws display tooth-like denticles, which come into direct contact with predators and prey. They are subjected to more regular and intense mechanical stress than other parts of the exoskeleton and therefore must be especially resistant to wear and abrasion. Here, we characterized denticle properties in five decapod species. Dactyls from three brachyuran crabs (Cancer borealis, Callinectes sapidus, and Chionoecetes opilio) and two anomuran crabs (Paralomis birsteini and Paralithodes camtschaticus) were sectioned normal to the contact surface of the denticle, revealing the interior of the denticle and the bulk endocuticle in which it is embedded. Microhardness, micro- and ultrastructure, and elemental composition were assessed along a transect running the width of the cuticle using microindentation hardness testing, optical and scanning electron microscopy (SEM), and energy dispersive X-ray spectroscopy (EDS), respectively. In all species tested, hardness was dramatically higher-up to ten times-in the denticle than in the bulk endocuticle. Likewise, in all species there was an increase in packing density of mineralized chitin-protein fibers, a decrease in width of the pore canals that run through the cuticle, and a decrease in phosphorous content from endocuticle to denticle. The changes in hardness across the cuticle, and the relationship between hardness, calcium, and magnesium content, however, varied among species. Although mechanical resistance of the denticles was exceptionally high in all species, the basis for resistance appears to differ among species. STATEMENT OF SIGNIFICANCE: Understanding the diverse mechanisms by which animals attain exceptionally high mechanical resistance may enable development of novel, biologically inspired materials. Decapod crustacean claws, and particularly the tooth-like denticles that these claws display, are of interest in this regard, as they must be especially resistant to wear. We assessed mechanical, elemental, and structural properties of the claw cuticle in five decapod species. Without exception, microhardness was dramatically higher in the denticle than in the bulk endocuticle. Multivariant statistical analyses, however, showed that the relationships among microhardness, elemental content, and structural variables differed among species. Such patterns likely result from strong evolutionary pressure on feeding and defensive structures and a trade-off between mechanical properties and energetic cost of exoskeleton formation.

Upwelling buffers climate change impacts on coral reefs of the eastern tropical Pacific.

Corals of the eastern tropical Pacific live in a marginal and oceanographically dynamic environment. Along the Pacific coast of Panamá, stronger seasonal upwelling in the Gulf of Panamá in the east transitions to weaker upwelling in the Gulf of Chiriquí in the west, resulting in complex regional oceanographic conditions that drive differential coral-reef growth. Over millennial timescales, reefs in the Gulf of Chiriquí recovered more quickly from climatic disturbances compared with reefs in the Gulf of Panamá. In recent decades, corals in the Gulf of Chiriquí have also had higher growth rates than in the Gulf of Panamá. As the ocean continues to warm, however, conditions could shift to favor the growth of corals in the Gulf of Panamá, where upwelling may confer protection from high-temperature anomalies. Here we describe the recent spatial and temporal variability in surface oceanography of nearshore environments in Pacific Panamá and compare those conditions with the dynamics of contemporary coral-reef communities during and after the 2016 coral-bleaching event. Although both gulfs have warmed significantly over the last 150 yr, the annual thermal maximum in the Gulf of Chiriquí is increasing faster, and ocean temperatures there are becoming more variable than in the recent past. In contrast to historical trends, we found that coral cover, coral survival, and coral growth rates were all significantly higher in the Gulf of Panamá. Corals bleached extensively in the Gulf of Chiriquí following the 2015-2016 El Niño event, whereas upwelling in the Gulf of Panamá moderated the high temperatures caused by El Niño, allowing the corals largely to escape thermal stress. As the climate continues to warm, upwelling zones may offer a temporary and localized refuge from the thermal impacts of climate change, while reef growth in the rest of the eastern tropical Pacific continues to decline.

Indirect effects of climate change altered the cannibalistic behaviour of shell-drilling gastropods in Antarctica during the Eocene.

The fossil record from Seymour Island, Antarctic Peninsula, provides a record of biotic response to the onset of global climatic cooling during the Eocene. Using drilling traces-small, round holes preserved on prey shells-we examined the effect of a cooling pulse 41 Ma on the cannibalistic behaviour of predatory naticid gastropods. We predicted that cannibalistic attacks would decline in response to the cooling climate, reflecting reduced activity levels, energy requirements and constraints on the chemically aided drilling process of the naticids. Surprisingly, however, cannibalism frequencies did not change. This counterintuitive result is best explained by a sharp reduction in durophagous (shell-crushing) predation in shallow-benthic communities in Antarctica that also occurred as the climate cooled. Reduced durophagous predation may have created a less-risky environment for foraging naticids, stimulating cannibalistic behaviour. The change in the top-down control exerted by shell-crushing predators on naticids may have counteracted the direct, negative effects of declining temperatures on the predatory performance of naticids. Our results suggest that the long-term consequences of climate change cannot be predicted solely from its direct effects on predation, because the temperature can have large indirect effects on consumer-resource interactions, especially where risk-effects dominate.

A 50-year retrospective of persistent organic pollutants in the fat and eggs of penguins of the Southern Ocean.

Persistent organic pollutants (POPs) such as dichlorodiphenyltrichloroethanes (DDTs), hexachlorobenzene (HCB), hexachlorocyclohexanes (HCHs), and polychlorinated biphenyls (PCBs) have been spreading to Antarctica for over half a century. Penguins are effective indicators of pelagic concentrations of POPs. We synthesized the literature on penguins to assess temporal trends of pelagic contamination in Antarctica, using fat and eggs to monitor changes from 1964 to 2011. DDT/DDE ratios suggest long-range atmospheric transport. Average DDT in fat (ww) increased from 44 ng g-1 in the 1960s, peaked at 171 ng g-1 in the mid-1980s, and then declined slowly to the present level of 101 ng g-1. Temporal trends in HCB contamination rose into the 1990s before declining. ∑HCHs in fat was ∼5 ng g-1 from 1960 to 1979, peaking at 33 ng g-1 during the period 1980-1989 before declining to ∼5 ng g-1 from 1990 to present. PCBs rose substantially from 1970 to 2009 in fat, varying more than DDTs and HCB in both fat and eggs. Antarctic penguins are good biological indicators of global DDT and HCB emissions, but the existing data are insufficient regarding HCHs and PCBs.

No barrier to emergence of bathyal king crabs on the Antarctic shelf.

Cold-water conditions have excluded durophagous (skeleton-breaking) predators from the Antarctic seafloor for millions of years. Rapidly warming seas off the western Antarctic Peninsula could now facilitate their return to the continental shelf, with profound consequences for the endemic fauna. Among the likely first arrivals are king crabs (Lithodidae), which were discovered recently on the adjacent continental slope. During the austral summer of 2010 ‒ 2011, we used underwater imagery to survey a slope-dwelling population of the lithodid Paralomis birsteini off Marguerite Bay, western Antarctic Peninsula for environmental or trophic impediments to shoreward expansion. The population density averaged ∼ 4.5 individuals × 1,000 m(-2) within a depth range of 1,100 ‒ 1,500 m (overall observed depth range 841-2,266 m). Images of juveniles, discarded molts, and precopulatory behavior, as well as gravid females in a trapping study, suggested a reproductively viable population on the slope. At the time of the survey, there was no thermal barrier to prevent the lithodids from expanding upward and emerging on the outer shelf (400- to 550-m depth); however, near-surface temperatures remained too cold for them to survive in inner-shelf and coastal environments (<200 m). Ambient salinity, composition of the substrate, and the depth distribution of potential predators likewise indicated no barriers to expansion of lithodids onto the outer shelf. Primary food resources for lithodids--echinoderms and mollusks--were abundant on the upper slope (550-800 m) and outer shelf. As sea temperatures continue to rise, lithodids will likely play an increasingly important role in the trophic structure of subtidal communities closer to shore.

Lost opportunities: coral recruitment does not translate to reef recovery in the Florida Keys.

We tested the hypothesis that the poor recovery of the coral populations on reefs in the Florida Keys is related to low coral recruitment. In the summer of 2011, we deployed 240 terracotta tiles at eight study sites in a balanced design: (i) among three depths; and (ii) between fished and unfished reefs. Corals recruited to ∼ 40% of the deployed tiles, with more corals settling on tiles on unfished reefs than on fished reefs. The apparent effect of protection was not a consequence of different densities of herbivorous fishes, but was more likely related to local hydrography and the tendency of the no-take reserves to act as larval sinks, particularly in the lower Florida Keys. There was a mismatch between the coral taxa that recruited and the adult coral assemblages, suggesting that recruits were arriving but not surviving to contribute to coral recovery.

Coral reef baselines: how much macroalgae is natural?

Identifying the baseline or natural state of an ecosystem is a critical step in effective conservation and restoration. Like most marine ecosystems, coral reefs are being degraded by human activities: corals and fish have declined in abundance and seaweeds, or macroalgae, have become more prevalent. The challenge for resource managers is to reverse these trends, but by how much? Based on surveys of Caribbean reefs in the 1970s, some reef scientists believe that the average cover of seaweed was very low in the natural state: perhaps less than 3%. On the other hand, evidence from remote Pacific reefs, ecological theory, and impacts of over-harvesting in other systems all suggest that, historically, macroalgal biomass may have been higher than assumed. Uncertainties about the natural state of coral reefs illustrate the difficulty of determining the baseline condition of even well studied systems.

Is Jamaica a good model for understanding Caribbean coral reef dynamics?

Caribbean reefs have experienced unprecedented changes in the past 40 years. A major hypothesis to explain shifts in reef community composition relates to declining herbivory. This hypothesis was developed largely based on observations of Jamaican reefs from the 1980s onward, but it is widely held to be relevant regionally. We use a region-wide dataset on benthic composition to examine how well the pattern of ecological change on Jamaican reefs is mirrored by other Caribbean reefs. The extent to which macroalgal cover exceeds coral cover on Jamaican reefs is an order of magnitude more extreme than seen elsewhere. We suggest that Jamaican reefs are not representative of the degradation trajectory of Caribbean reefs and management based on the Jamaican experience may not be relevant elsewhere. However, the recovery of Jamaican reefs following the return of urchins gives us hope that Caribbean reefs are more resilient to catastrophic disturbances than previously thought.

ENSO drove 2500-year collapse of eastern Pacific coral reefs.

Cores of coral reef frameworks along an upwelling gradient in Panamá show that reef ecosystems in the tropical eastern Pacific collapsed for 2500 years, representing as much as 40% of their history, beginning about 4000 years ago. The principal cause of this millennial-scale hiatus in reef growth was increased variability of the El Niño-Southern Oscillation (ENSO) and its coupling with the Intertropical Convergence Zone. The hiatus was a Pacific-wide phenomenon with an underlying climatology similar to probable scenarios for the next century. Global climate change is probably driving eastern Pacific reefs toward another regional collapse.

Catastrophe and the life span of coral reefs.

A strong earthquake in the western Caribbean in 2009 had a catastrophic impact on uncemented, unconsolidated coral reefs in the central sector of the shelf lagoon of the Belizean barrier reef. In a set of 21 reef sites that had been observed prior to the earthquake, the benthic assemblages of 10 were eradicated, and one was partially damaged, by avalanching of their slopes. Ecological dynamics that had played out over the previous 23 years, including the mass mortalities of two sequentially dominant coral species and a large increase in the cover of an encrusting sponge, were instantaneously rendered moot in the areas of catastrophic reef-slope failure. Because these prior dynamics also determined the benthic composition and resilience of adjacent sections of reef that remained intact, the history of disturbance prior to the earthquake will strongly influence decadal-scale recovery in the failed areas. Geological analysis of the reef framework yielded a minimum return time of 2000-4000 years for this type of high-amplitude event. Anthropogenic degradation of ecosystems must be viewed against the backdrop of long-period, natural catastrophes, such as the impact of strong earthquakes on uncemented, lagoonal reefs.

Anthropogenic impacts on marine ecosystems in Antarctica.

Antarctica is the most isolated continent on Earth, but it has not escaped the negative impacts of human activity. The unique marine ecosystems of Antarctica and their endemic faunas are affected on local and regional scales by overharvesting, pollution, and the introduction of alien species. Global climate change is also having deleterious impacts: rising sea temperatures and ocean acidification already threaten benthic and pelagic food webs. The Antarctic Treaty System can address local- to regional-scale impacts, but it does not have purview over the global problems that impinge on Antarctica, such as emissions of greenhouse gases. Failure to address human impacts simultaneously at all scales will lead to the degradation of Antarctic marine ecosystems and the homogenization of their composition, structure, and processes with marine ecosystems elsewhere.

Changing patterns of microhabitat utilization by the threespot damselfish, Stegastes planifrons, on Caribbean reefs.

BACKGROUND: The threespot damselfish, Stegastes planifrons (Cuvier), is important in mediating interactions among corals, algae, and herbivores on Caribbean coral reefs. The preferred microhabitat of S. planifrons is thickets of the branching staghorn coral Acropora cervicornis. Within the past few decades, mass mortality of A. cervicornis from white-band disease and other factors has rendered this coral a minor ecological component throughout most of its range. METHODOLOGY/PRINCIPAL FINDINGS: Survey data from Jamaica (heavily fished), Florida and the Bahamas (moderately fished), the Cayman Islands (lightly to moderately fished), and Belize (lightly fished) indicate that distributional patterns of S. planifrons are positively correlated with live coral cover and topographic complexity. Our results suggest that species-specific microhabitat preferences and the availability of topographically complex microhabitats are more important than the abundance of predatory fish as proximal controls on S. planifrons distribution and abundance. CONCLUSIONS/SIGNIFICANCE: The loss of the primary microhabitat of S. planifrons-A. cervicornis-has forced a shift in the distribution and recruitment of these damselfish onto remaining high-structured corals, especially the Montastraea annularis species complex, affecting coral mortality and algal dynamics throughout the Caribbean.

Climate change and trophic response of the Antarctic bottom fauna.

BACKGROUND: As Earth warms, temperate and subpolar marine species will increasingly shift their geographic ranges poleward. The endemic shelf fauna of Antarctica is especially vulnerable to climate-mediated biological invasions because cold temperatures currently exclude the durophagous (shell-breaking) predators that structure shallow-benthic communities elsewhere. METHODOLOGY/PRINCIPAL FINDINGS: We used the Eocene fossil record from Seymour Island, Antarctic Peninsula, to project specifically how global warming will reorganize the nearshore benthos of Antarctica. A long-term cooling trend, which began with a sharp temperature drop approximately 41 Ma (million years ago), eliminated durophagous predators-teleosts (modern bony fish), decapod crustaceans (crabs and lobsters) and almost all neoselachian elasmobranchs (modern sharks and rays)-from Antarctic nearshore waters after the Eocene. Even prior to those extinctions, durophagous predators became less active as coastal sea temperatures declined from 41 Ma to the end of the Eocene, approximately 33.5 Ma. In response, dense populations of suspension-feeding ophiuroids and crinoids abruptly appeared. Dense aggregations of brachiopods transcended the cooling event with no apparent change in predation pressure, nor were there changes in the frequency of shell-drilling predation on venerid bivalves. CONCLUSIONS/SIGNIFICANCE: Rapid warming in the Southern Ocean is now removing the physiological barriers to shell-breaking predators, and crabs are returning to the Antarctic Peninsula. Over the coming decades to centuries, we predict a rapid reversal of the Eocene trends. Increasing predation will reduce or eliminate extant dense populations of suspension-feeding echinoderms from nearshore habitats along the Peninsula while brachiopods will continue to form large populations, and the intensity of shell-drilling predation on infaunal bivalves will not change appreciably. In time the ecological effects of global warming could spread to other portions of the Antarctic coast. The differential responses of faunal components will reduce the endemic character of Antarctic subtidal communities, homogenizing them with nearshore communities at lower latitudes.