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Deepwater Program: Lophelia II, continuing ecological research on deep-sea corals and deep-reef habitats in the Gulf of Mexico

The deep sea is a rich environment composed of diverse habitat types. While deep-sea coral habitats have been discovered within each ocean basin, knowledge about the ecology of these habitats and associated inhabitants continues to grow. This report presents information and results from the Lophelia II project that examined deep-sea coral habitats in the Gulf of Mexico. The Lophelia II project focused on Lophelia pertusa habitats along the continental slope, at depths ranging from 300 to 1,000 meters. The chapters are authored by several scientists from the U.S. Geological Survey, National Oceanic and Atmospheric Administration, University of North Carolina Wilmington, and Florida State University who examined the community ecology (from microbes to fishes), deep-sea coral age, growth, and reproduction, and population connectivity of deep-sea corals and inhabitants. Data from these studies are presented in the chapters and appendixes of the report as well as in journal publications. This study was conducted by the Ecosystems Mission Area of the U.S. Geological Survey to meet information needs identified by the Bureau of Ocean Energy Management.

Gulf of Mexico

Climate change, coral loss, and the curious case of the parrotfish paradigm: Why don't marine protected areas improve reef resilience?

Scientists have advocated for local interventions, such as creating marine protected areas and implementing fishery restrictions, as ways to mitigate local stressors to limit the effects of climate change on reef-building corals. However, in a literature review, we find little empirical support for the notion of managed resilience. We outline some reasons for why marine protected areas and the protection of herbivorous fish (especially parrotfish) have had little effect on coral resilience. One key explanation is that the impacts of local stressors (e.g., pollution and fishing) are often swamped by the much greater effect of ocean warming on corals. Another is the sheer complexity (including numerous context dependencies) of the five cascading links assumed by the managed-resilience hypothesis. If reefs cannot be saved by local actions alone, then it is time to face reef degradation head-on, by directly addressing anthropogenic climate change—the root cause of global coral decline.

Annual Review of Marine Science

Holocene core logs and site statistics for modern patch-reef cores: Biscayne National Park, Florida

The bedrock in Biscayne National Park (BNP), a 1,730-square kilometer (km2) region off southeast Florida, consists of Pleistocene (1.8 million years ago (Ma) to 10,000 years ago (ka)) and Holocene (10 ka to present) carbonate rocks (Enos and Perkins, 1977; Halley and others, 1997; Multer and others, 2002). Most of the surficial limestone in BNP, including the islands of the Florida Keys, was formed at ~125 ka during the highstand of marine oxygen-isotope substage 5e, when sea level was approximately 6 meters (m) higher than today (Chappell and Shackleton, 1986; Multer and others, 2002; Lidz and others, 2003; Siddall and others, 2003; Balsillie and Donoghue, 2004). During the substage-5e regression, the entire Florida Platform became exposed. Subaerial exposure lasted for approximately 115,000 years (kyr), which resulted in erosion and enhancement of karst-like features (Lidz and others, 2006). As the Holocene transgression began to flood the Florida shelf ~7 to 6 ka, the bedrock depression under Biscayne Bay began to flood, and Holocene coral and reef debris laid the foundation for the present reef system (Enos and Perkins, 1977; Lighty and others, 1982; Toscano and Macintyre, 2003; Lidz and others, 2006). More than 3,000 patch reefs exist within the BNP boundary. Most contain hermatypic corals of various species such as those belonging to Montastrea, Diploria, Siderastrea, Porites, Acropora, and Agaricia. Patch reefs within BNP have two morphologies: pinnacle and flat top. Experimental Advanced Airborne Research Lidar (EAARL) data collected along the offshore BNP coral reef tract show that these two morphologies are clearly defined both in the high-resolution bathymetry maps produced by the Lidar data and by statistical analyses of the Lidar dataset (Brock and others, 2008). Brock and others (2008) also show that the pinnacle patch reefs are deeper than the more shallow, broad, and flat patch reefs. The control for these two patch-reef morphologies is unclear; however, their shapes may be due to a slightly lowered sea level or a stillstand in the middle-Holocene around 4 ka that caused erosion of the shallower reefs and allowed the deeper reefs to remain unaffected. Lidz and others (2006) have suggested a stillstand around 4 ka that carved a 2.5-kilometer (km)-wide nearshore rock ledge into the seaward side of every island in the Florida Keys. The objectives of this study were to sample living corals to understand the more recent (<200 years) changes in climate and environmental conditions of the area and to investigate the Holocene (in this case, <8,000 years in the Florida Keys) depositional history at progressively deeper patch-reef sites. This report provides statistics for the cores and core sites and a basic lithologic description of these Holocene cores.

Florida

U-series ages of solitary corals from the California coast by mass spectrometry

The purpose of this study is to evaluate the feasibility of dating fossil solitary corals from Pleistocene marine strandlines outside tropical latitudes using the recently developed high sensitivity, high-precision U-series technique based on thermal-ionization mass-spectrometry (TIMS). The TIMS technique is much more efficient than conventional a spectrometry and, as a result, multiple samples of an individual coral skeleton, or different specimens from the same bed can be analyzed. Detached and well-rounded fossil specimens of the solitary coral Balanophyllia elegans were collected from relict littoral deposits on emergent marine terraces along the California coast at Cayucos terrace (elevation 8 m, previously dated at 124 and 117 Ky by α counting), Shell Beach terrace (elevation about 25 m, previously undated), Nestor terrace, San Diego (elevation 23 m, previously dated at 131 to 109 Ky ), Bird Rock terrace, San Diego ( elevation 8 m, previously dated at 81 Ky ). Attached living specimens were collected from the intertidal zone on the modern terrace at Moss Beach. Concentrations of 232 Th in both living and fossil specimens are much higher than in reef-building corals (12 to 624 pmol/g vs. 0.1 to 1.6 pmol/g, respectively). However, because 230 Th/ 232 Th in Balanophyllia elegans are very low (2.22 × 10 −3 to 4.33 × 10 −4 ), the high 232 Th concentrations have negligible effect on the 230 Th- 234 U dates. The high 232 Th concentration in the living specimen (33.1 pmol/g) indicates that a significant amount of 232 Th is incorporated in the aragonitic skeleton during growth, or attached to clay-sized silicates trapped in the skeletal material. The calculated initial 234 U activities in the fossil specimens of Balanophyllia elegans are higher than the 234 U activity in modern seawater or in the modern specimen. The higher initial activities could possibly reflect the influx of 234 U-enriched continental water into Pleistocene coastal waters, or it could reflect minor diagenetic alteration, a persistent and fundamental problem in dating all corals. Samples from a compound specimen from the Cayucos terrace were subjected to different preparation procedures. Samples prepared by a standard acid washing procedure yielded 230 Th- 234 U ages of 125, 123, and 122 Ky, whereas samples prepared by an abbreviated procedure without acid washing yield significantly lower ages of 113 and 112 Ky. Two other specimens from the same bed yielded 230 Th- 234 U ages of 118 and 115 Ky. Also, two specimens from a stratigraphically higher bed yielded ages of 120 and 117 Ky, and three specimens from a lower bed yield ages of 115, 113, and 101 Ky. Nine of the twelve ages of the treated samples from the Cayucos terrace range from 125 to 113 Ky. However, the ages do not follow the stratigraphie order. Two possible interpretations are ( 1 ) the age of the terrace deposit is 125 Ky and all younger ages reflect variable diagenetic alteration or (2) the age of the terrace is 125 to 113 Ky and the ages reflect sediment reworking over a period of 12 Ky. Three specimens from a single bed on the Shell Beach terrace yield ages of 126, 122, and 121 Ky, similar to the older ages from Cayucos. The ages of solitary corals from the Cayucos and Shell Beach terraces are similar to ages of reef-building corals from terraces at numerous tropical localities. These are correlated with the last interglacial sea-level highstand, which probably stood 2 to 10 m above present sea level. The youngest ages and present elevations of the Cayucos and Shell Beach terraces yield tectonic uplift rates of 0.01 and 0.15 m/Ky, respectively, assuming the original elevation of each terrace was 7 m. Four specimens from the basal gravel on the Nestor terrace yielded ages of 145, 143, 137, and 133 Ky. The three oldest ages, however, are older than that associated with the last interglacial. The possible explanations for these older ages are ( 1 ) diagenic alteration or ( 2 ) the Nestor terrace deposits reflect in some way a poorly documented early phase of the last interglacial sea level high stand.

Geochimica et Cosmochimica Acta

Physicochemical controls on zones of higher coral stress where Black Band Disease occurs at Mākua Reef, Kauaʻi, Hawaiʻi

Pervasive and sustained coral diseases contribute to the systemic degradation of reef ecosystems, however, to date an understanding of the physicochemical controls on a coral disease event is still largely lacking. Water circulation and residence times and submarine groundwater discharge all determine the degree to which reef organisms are exposed to the variable chemistry of overlying waters; understanding these physical controls is thus necessary to interpret spatial patterns in coral health. The recent discovery of coral Black Band Disease at Mākua Reef on Kauaʻi, Hawaiʻi prompted an investigation into the physicochemical drivers and geomorphic controls of reef water circulation, and the temporally variable nutrient fluxes derived from submarine groundwater discharge. Results reveal localized stagnant water parcels at Mākua Reef where groundwater-derived high nutrient loading and low salinities act in concert as stressors to coralline health – and where Black Band Disease was uniquely identified. The observed high nutrient levels during low tide conditions are likely associated with nearby upstream cesspools and drain fields. Information obtained using such a multidisciplinary approach has direct value for successful management of coastal aquifers and the health and sustainability of adjacent nearshore coral reef ecosystems.

Hawaii

Sea‐level rise will drive divergent sediment transport patterns on fore reefs and reef flats, potentially causing erosion on atoll islands

Atoll reef islands primarily consist of unconsolidated sediment, and their ocean‐facing shorelines are maintained by sediment produced and transported across their reefs. Changes in incident waves can alter cross‐shore sediment exchange and, thus, affect the sediment budget and morphology of atoll reef islands. Here we investigate the influence of sea level rise and projected wave climate change on wave characteristics and cross‐shore sediment transport across an atoll reef at Kwajalein Island, Republic of the Marshall Islands. Using a phase‐resolving model, we quantify the influence on sediment transport of quantities not well captured by wave‐averaged models, namely, wave asymmetry and skewness and flow acceleration. Model results suggest that for current reef geometry, sea level, and wave climate, potential bedload transport is directed onshore, decreases from the fore reef to the beach, and is sensitive to the influence of flow acceleration. We find that a projected 12% decrease in annual wave energy by 2100 CE has negligible influence on reef flat hydrodynamics. However, 0.5–2.0 m of sea level rise increases wave heights, skewness, and shear stress on the reef flat and decreases wave skewness and shear stress on the fore reef. These hydrodynamic changes decrease potential sediment inputs onshore from the fore reef where coral production is greatest but increase potential cross‐reef sediment transport from the outer reef flat to the beach. Assuming sediment production on the fore reef remains constant or decreases due to increasing ocean temperatures and acidification, these processes have the potential to decrease net sediment delivery to atoll islands, causing erosion.

Journal of Geophysical Research – Earth Surface

Assessment of antibiotic resistance genes in Caribbean corals, including those treated with amoxicillin

The decimation of reefs from stony coral tissue loss disease prompted the use of a topical amoxicillin treatment to prevent coral mortality. Application of this treatment led to concerns about unintentional impacts such as potential alteration of the coral microbiome and possible spread of antibiotic resistance. We used three different methodologies—microbial RNA sequencing, 16S rRNA amplicon surveys, and microbial qPCR array—to assess these concerns and to establish a baseline of antibiotic resistance genes (ARGs) in untreated coral microbes. We conducted microbial RNA sequencing on wild Montastraea cavernosa coral mucus samples collected before and 24 h after amoxicillin application. While diverse antibiotic resistance genes (ARGs) were expressed, no differences in ARG expression were detected after amoxicillin treatment. Additionally, there were no notable changes in the microbial communities between the before and after samples. In a separate experiment, a microbial qPCR array was used to assess differences in ARGs over longer timescales using cores from wild Colpophyllia natans , comparing never-treated corals with ones treated a single time seven months prior and with those treated multiple times seven months and more prior. No clinically relevant ARGs were detected across any samples. A small number of above-detection reads (4 in the never-treated corals, 2 in the once-treated corals, and 0 in the multi-treated corals) may indicate weak amplification of similar environmental (non-anthropogenic) ARGs in the corals. Results indicate that the localized topical application of amoxicillin to prevent mortality of SCTLD-affected corals does not: (1) significantly disrupt microbiomes, (2) increase ARG expression in adjacent tissues of these species within 24 h, nor (3) increase abundance of clinically relevant ARGs over a 7 month time period.

Florida

Do three massive coral species from the same reef record the same SST signal? A test from the Dry Tortugas, Florida Keys

Paleoclimatologists have reconstructed century-long records of sea surface temperature (SST) in the Pacific using the Sr/Ca of massive corals, whereas similar reconstructions in the Atlantic have not proceeded at the same pace. Past research in the Florida Keys has focused on Montastrea spp. , an abundant and fast-growing massive coral, thus a good candidate for climate reconstructions. However, coral records from the Florida Keys are complicated by freshwater flux, which varies the Sr/Ca in seawater, thus confounding the Sr/Ca to SST signal. In this research, we compared the monthly Sr/Ca variations in three massive corals species ( Montastraea faveolata, Diploria strigosa, and Siderastrea siderea ) from the same reef in the nearly pristine Dry Tortugas National Park (24.70N, 82.80W) at the southwestern extent of the Florida Keys. This location is ideal for a calibration study as hourly water temperature records are available and the remote reef is far from mainland freshwater influence. These corals experienced the same environmental conditions (water depth, clarity, Sr/Ca of seawater, etc.) but differ in the mean annual growth rates (0.86 &plusmn;0.10 (1&sigma;) cm/year M. faveolata ; 0.67 &plusmn;0.04 (1&sigma;) cm/year D. strigosa ; 0.44 &plusmn;0.04 (1&sigma;) cm/year S. siderea ). The mean Sr/Ca values are not the same but decrease with mean annual growth rates (9.201 &plusmn;0.091 (1&sigma;) mmol/mol M. faveolata ; 9.177 &plusmn;0.081 (1&sigma;) mmol/mol D. strigosa ; 8.964 &plusmn;0.12 (1&sigma;) mmol/mol S. siderea ), thus supporting the &ldquo;vital effect&rdquo; or biological differences during calcification between coral species. The amplitude of the seasonal cycle in Sr/Ca varies with the slower growing S. siderea having the largest mean amplitude and D. strigosa the smallest (0.340 mmol/mol S. siderea ; 0.284 mmol/mol M. faveolata ; 0.238 mmol/mol D. strigosa ). We confirmed our sampling methods by conducting several intracolony and intercolony coral Sr/Ca replication tests and found a high correlation in all tests (>0.95 S. siderea ; >0.90 D. strigosa ; >0.83 M. faveolata; p < 0.05). The weighted linear regression of monthly coral Sr/Ca to mean monthly SST revealed that S. siderea captured the seasonal and interannual variability in SST ( r = -0.97, -0.61 monthly and monthly anomalies, respectively, p < 0.05). The other corals have reduced correlation with monthly anomalies and do not capture the seasonal variability with the same fidelity as S. siderea . All three corals were sampled along the thecal wall following the same procedures; however, each coral species has a different skeletal structure, density, and micro-scale growth patterns. We hypothesize the thecal wall of S. siderea calcifies at a continuous rate along the time-growth axis whereas the wall of D. strigosa and M. faveolata reflects a more complex signal. Of the three species, the slow growing S. siderea provides a robust reconstruction of mean monthly SST for the Dry Tortugas thus suitable for longer centennial-scale reconstructions.

Florida

The influence of different deep-sea coral habitats on sediment macrofaunal community structure and function

Deep-sea corals can create a highly complex, three-dimensional structure that facilitates sediment accumulation and influences adjacent sediment environments through altered hydrodynamic regimes. Infaunal communities adjacent to different coral types, including reef-building scleractinian corals and individual colonies of octocorals, are known to exhibit higher macrofaunal densities and distinct community structure when compared to non-coral soft-sediment communities. However, the coral types have different morphologies, which may modify the adjacent sediment communities in discrete ways. Here we address: (1) how infaunal communities and their associated sediment geochemistry compare among deep-sea coral types ( Lophelia pertusa, Madrepora oculata, and octocorals) and (2) do infaunal communities adjacent to coral habitats exhibit typical regional and depth-related patterns observed in the Gulf of Mexico (GOM). Sediment push cores were collected to assess diversity, composition, numerical abundance, and functional traits of macrofauna (>300 µm) across 450 kilometers in the GOM at depths ranging from 263–1,095 m. Macrofaunal density was highest in L. pertusa habitats, but similar between M. oculata and octocorals habitats. Density overall exhibited a unimodal relationship with depth, with maximum densities between 600 and 800 m. Diversity and evenness were highest in octocoral habitats; however, there was no relationship between diversity and depth. Infaunal assemblages and functional traits differed among coral habitats, with L. pertusa habitats the most distinct from both M. oculata and octocorals. These patterns could relate to differences in sediment geochemistry as L. pertusa habitats contained high organic carbon content but low proportions of mud compared to both M. oculata and octocoral habitats. Distance-based linear modeling revealed depth, mud content, and organic carbon as the primary factors in driving coral infaunal community structure, while geographic location (longitude) was the primary factor in functional trait composition, highlighting both the location and ecological differences of L. pertusa habitats from other coral habitats. Enhanced habitat structural complexity associated with L. pertusa and differences in localized hydrodynamic flow may contribute to the dissimilarities in the communities found among the coral types. Our results suggest a decoupling for infaunal coral communities from the typical depth-related density and diversity patterns present throughout soft-sediment habitats in the GOM, highlighting the importance of deep-sea corals in structuring unique communities in the nearby benthos.

PeerJ

Pleistocene corals of the Florida keys: Architects of imposing reefs - Why?

Five asymmetrical, discontinuous, stratigraphically successive Pleistocene reef tracts rim the windward platform margin off the Florida Keys. Built of large head corals, the reefs are imposing in relief (???30 m high by 1 km wide), as measured from seismic profiles. Well dated to marine oxygen isotope substages 5c, 5b, and 5a, corals at depth are inferred to date to the Stage 6/5 transition. The size of these reefs attests to late Pleistocene conditions that repeatedly induced vigorous and sustained coral growth. In contrast, the setting today, linked to Florida Bay and the Gulf of Mexico, is generally deemed marginal for reef accretion. Incursion onto the reef tract of waters that contain seasonally inconsistent temperature, salinity, turbidity, and nutrient content impedes coral growth. Fluctuating sea level and consequent settings controlled deposition. The primary dynamic was position of eustatic zeniths relative to regional topographic elevations. Sea level during the past 150 ka reached a maximum of ???10.6 m higher than at present ???125 ka, which gave rise to an inland coral reef (Key Largo Limestone) and ooid complex (Miami Limestone) during isotope substage 5e. These formations now form the Florida Keys and a bedrock ridge beneath The Quicksands (Gulf of Mexico). High-precision radiometric ages and depths of dated corals indicate subsequent apices remained ???15 to 9 m, respectively, below present sea level. Those peaks provided accommodation space sufficient for vertical reef growth yet exposed a broad landmass landward of the reefs for >100 ka. With time, space, lack of bay waters, and protection from the Gulf of Mexico, corals thrived in clear oceanic waters of the Gulf Stream, the only waters to reach them.

Journal of Coastal Research

Decadal-scale changes in benthic foraminiferal assemblages off Key Largo, Florida

Assemblages of foraminiferal tests in sediments sampled off Key Largo, Florida, in 1982, 1991, and 1992 were significantly different from assemblages sampled along the same traverses in 1959–1961. Larger, algal symbiont-bearing taxa, primarily Soritidae, comprised 50–80% of the specimens in samples collected in 1959–1961, whereas Miliolidae and Rotaliidae comprised 65-90% of the specimens collected in 1991 and 1992. Test abundance in 1992 samples ranged from 1.0 × 10 2 /g to 8.1 × 104/g; tests were least abundant in coarse, well-sorted sediments. The lack of test-density data for the 1959-1961 samples prevented assessment of whether densities of smaller foraminifera have increased, symbiotic foraminifera have decreased, or both. Between 1982 and 1992, densities of smaller foraminifera appear to have increased. Although the causes of these changes in foraminiferal assemblages are not known, possible factors include nutrient loading inshore, winnowing and transport of tests by storm activity, and disease. The shift in dominance from long-lived, algal symbiont-bearing taxa in 1959--1961 to small, fast-growing, heterotrophic taxa in 1992 is consistent with predictions of community response to gradually increasing nutrient flux into south Florida's coastal waters. This study indicates that published accounts of foraminiferal assemblages from sediments collected 30 or more years ago can be valuable resources in efforts to determine if biotic changes have occurred in coastal ecosystems. This study also indicates that family-level identifications may be sufficient to detect decadal-scale changes in foraminiferal assemblages in reef-tract sediments.

Florida

Methods for monitoring corals and crustose coralline algae to quantify in-situ calcification rates

The potential effect of global climate change on calcifying marine organisms, such as scleractinian (reef-building) corals, is becoming increasingly evident. Understanding the process of coral calcification and establishing baseline calcification rates are necessary to detect future changes in growth resulting from climate change or other stressors. Here we describe the methods used to establish a network of calcification-monitoring stations along the outer Florida Keys Reef Tract in 2009. In addition to detailing the initial setup and periodic monitoring of calcification stations, we discuss the utility and success of our design and offer suggestions for future deployments. Stations were designed such that whole coral colonies were securely attached to fixed apparati (n = 10 at each site) on the seafloor but also could be easily removed and reattached as needed for periodic weighing. Corals were weighed every 6 months, using the buoyant weight technique, to determine calcification rates in situ. Sites were visited in May and November to obtain winter and summer rates, respectively, and identify seasonal patterns in calcification. Calcification rates of the crustose coralline algal community also were measured by affixing commercially available plastic tiles, deployed vertically, at each station. Colonization by invertebrates and fleshy algae on the tiles was low, indicating relative specificity for the crustose coralline algal community. We also describe a new, nonlethal technique for sampling the corals, used following the completion of the monitoring period, in which two slabs were obtained from the center of each colony. Sampled corals were reattached to the seafloor, and most corals had completely recovered within 6 months. The station design and sampling methods described herein provide an effective approach to assessing coral and crustose coralline algal calcification rates across time and space, offering the ability to quantify the potential effects of ocean warming and acidification on calcification processes.

Florida

Geology and biology of the "Sticky Grounds," shelf-margin carbonate mounds, and mesophotic ecosystem in the eastern Gulf of Mexico

Shelf-margin carbonate mounds in water depths of 116–135 m in the eastern Gulf of Mexico along the central west Florida shelf were investigated using swath bathymetry, side-scan sonar, sub-bottom imaging, rock dredging, and submersible dives. These enigmatic structures, known to fisherman as the “Sticky Grounds”, trend along slope, are 5–15 m in relief with base diameters of 5–30 m, and suggest widespread potential for mesophotic reef habitat along the west Florida outer continental shelf. Possible origins are sea-level lowstand coral patch reefs, oyster reefs, or perhaps more recent post-lowstand biohermal development. Rock dredging recovered bioeroded carbonate-rock facies comprised of bored and cemented bioclastics. Rock sample components included calcified worm tubes, pelagic sediment, and oysters normally restricted to brackish nearshore areas. Several reef sites were surveyed at the Sticky Grounds during a cruise in August 2010 with the R/V Seward Johnson using the Johnson-Sea-Link II submersible to ground truth the swath-sonar maps and to quantify and characterize the benthic habitats, benthic macrofauna, fish populations, and coral/sponge cover. This study characterizes for the first time this mesophotic reef ecosystem and associated fish populations, and analyzes the interrelationships of the fish assemblages, benthic habitats and invertebrate biota. These highly eroded rock mounds provide extensive hard-bottom habitat for reef invertebrate species as well as essential fish habitat for reef fish and commercially/recreationally important fish species. The extent and significance of associated living resources with these bottom types is particularly important in light of the 2010 Deepwater Horizon oil spill in the northeastern Gulf and the proximity of the Loop Current. Mapping the distribution of these mesophotic-depth ecosystems is important for quantifying essential fish habitat and describing benthic resources. These activities can improve ecosystem management and planning of future oil and gas activities in this outer continental shelf region.

Continental Shelf Research

Late Quaternary sea-level history of Saipan, Commonwealth of the Northern Mariana Islands, USA: A test of tectonic uplift and glacial isostatic adjustment models

In 1979, S. Uyeda and H. Kanamori proposed a tectonic model with two end members of a subduction-boundary continuum: the “Chilean” type (shallow dip of the subducting plate, great thrust events, compression, and uplift of the overriding plate) and a “Mariana” type (steep dip of the subducting plate, no great thrust events, tension, and no uplift). This concept has been used to explain variable rates of Quaternary uplift around the Pacific Rim, yet no uplift rates have been determined for the Mariana Islands themselves, one of the end members in this model. We studied the late Quaternary Tanapag Limestone, which rims much of the eastern and southern coasts of Saipan, Northern Mariana Islands, with elevations of ∼13 m to ∼30 m. Samples from 12 well-preserved corals ( Acropora , Porites , and Goniastrea ) yielded U-series ages ranging from ca. 134 ka to ca. 126 ka. These ages correlate the emergent reef of the Tanapag Limestone with the last interglacial period, when sea level was several meters above present. Ages and measured reef elevations from the Tanapag Limestone, along with paleo–sea-level data, yield relatively low late Quaternary uplift rates of 0.002–0.19 m/k.y., consistent with the Uyeda-Kanamori model. A review of data from other localities near subduction zones around the Pacific Basin, however, indicates that many coastlines do not fit the model. Uplift rates along the Chilean coast are predicted to be relatively high, but field studies indicate they are low. On some coastlines, relatively high uplift rates are better explained by subduction of seamounts or submarine ridges rather than subduction zone geometry. Despite the low long-term uplift rate on Saipan, the island also hosts an emergent, low-elevation (+3.9–4.0 m) reef with corals in growth position below a notch (+4.2 m). The corals are dated to 3.9–3.1 ka. The occurrence of this young, emergent reef is likely not due to tectonic uplift; instead, it is interpreted to be the result of glacial isostatic adjustment processes after the end of the last glacial period. Our findings are consistent with similar observations on tectonically stable or slowly uplifting islands elsewhere in the equatorial Pacific Ocean and agree with numerical models of a higher-than-present Holocene sea level in this region due to glacial isostatic adjustment processes.

Northern Mariana Islands, Saipan

Physiological differences in bleaching response of the coral Porites astreoides along the Florida Keys reef tract during high-temperature stress

The Florida Keys reef tract (FKRT) has a unique geological history wherein Holocene sea-level rise and bathymetry interacted, resulting in a reef-building system with notable spatial differences in reef development. Overprinted on this geologic history, recent global and local stressors have led to degraded reefs dominated by fleshy algae, soft corals, and sponges. Here, we assessed how coral physiology (calcification rate, tissue thickness, reproduction, symbiosis, and bleaching) varies seasonally (winter vs. summer) and geographically using 40 colonies of the mustard hill coral Porites astreoides from four sites across 350 km along the FKRT from 2015 to 2017. The study coincided with a high-temperature event in late summer 2015 that caused heterogeneous levels of coral bleaching across sites. Bleaching severity differed by site, with bleaching response more aligned with heat stress retroactively calculated from local degree heating weeks than those predicted by satellites. Despite differences in temperature profiles and bleaching severity, all colonies hosted Symbiodiniaceae of the same genus (formerly Clade A and subtypes). Overall, P. astreoides at Dry Tortugas National Park, the consistently coolest site, had the highest calcification rates, symbiont cell densities, and reproductive potential (all colonies were reproductive, with most planula larvae per polyp). Corals at Dry Tortugas and Fowey Rocks Light demonstrated strong seasonality in net calcification (higher in summer) and did not express visual or partial-mortality responses from the bleaching event; in contrast, colonies in the middle and southern part of the upper keys, Sombrero Key and Crocker Reef, demonstrated similar reduced fitness from bleaching, but differential recovery trajectories following the heat stress. Identifying reefs, such as Dry Tortugas and possibly Fowey Rocks Light that may serve as heat-stress refugia, is important in selecting candidate sites for adaptive reef-management strategies, such as selective propagation and assisted gene flow, to increase coral-species adaptation to ocean warming.

Florida

Diverse coral communities in mangrove habitats suggest a novel refuge from climate change

Risk analyses indicate that more than 90% of the world's reefs will be threatened by climate change and local anthropogenic impacts by the year 2030 under "business-as-usual" climate scenarios. Increasing temperatures and solar radiation cause coral bleaching that has resulted in extensive coral mortality. Increasing carbon dioxide reduces seawater pH, slows coral growth, and may cause loss of reef structure. Management strategies include establishment of marine protected areas with environmental conditions that promote reef resiliency. However, few resilient reefs have been identified, and resiliency factors are poorly defined. Here we characterize the first natural, non-reef coral refuge from thermal stress and ocean acidification and identify resiliency factors for mangrove&ndash;coral habitats. We measured diurnal and seasonal variations in temperature, salinity, photosynthetically active radiation (PAR), and seawater chemistry; characterized substrate parameters; and examined water circulation patterns in mangrove communities where scleractinian corals are growing attached to and under mangrove prop roots in Hurricane Hole, St. John, US Virgin Islands. Additionally, we inventoried the coral species and quantified incidences of coral bleaching, mortality, and recovery for two major reef-building corals, Colpophyllia natans and Diploria labyrinthiformis , growing in mangrove-shaded and exposed (unshaded) areas. Over 30 species of scleractinian corals were growing in association with mangroves. Corals were thriving in low-light (more than 70% attenuation of incident PAR) from mangrove shading and at higher temperatures than nearby reef tract corals. A higher percentage of C. natans colonies were living shaded by mangroves, and no shaded colonies were bleached. Fewer D. labyrinthiformis colonies were shaded by mangroves, however more unshaded colonies were bleached. A combination of substrate and habitat heterogeneity, proximity of different habitat types, hydrographic conditions, and biological influences on seawater chemistry generate chemical conditions that buffer against ocean acidification. This previously undocumented refuge for corals provides evidence for adaptation of coastal organisms and ecosystem transition due to recent climate change. Identifying and protecting other natural, non-reef coral refuges is critical for sustaining corals and other reef species into the future.

Biogeosciences

Reproducibility of geochemical and climatic signals in the Atlantic coral Montastraea faveolata

Monthly resolved, 41-year-long stable isotopic and elemental ratio time series were generated from two separate heads of Montastraea faveolata from Looe Key, Florida, to assess the fidelity of using geochemical variations in Montastraea, the dominant reef-building coral of the Atlantic, to reconstruct sea surface environmental conditions at this site. The stable isotope time series of the two corals replicate well; mean values of ??18O and ??13C are indistinguishable between cores (compare 0.70??? versus 0.68??? for ??13C and -3.90??? versus - 3.94??? for ??18O). Mean values from the Sr/Ca time series differ by 0.037 mmol/mol, which is outside of analytical error and indicates that nonenvironmental factors are influencing the coral Sr/ Ca records at Looe Key. We have generated significant ?? 18O-sea surface temperature (SST) (R = -0.84) and Sr/ Ca-SST (R = -0.86) calibration equations at Looe Key; however, these equations are different from previously published equations for Montastraea. Variations in growth parameters or kinetic effects are not sufficient to explain either the observed differences in the mean offset between Sr/Ca time series or the disagreement between previous calibrations and our calculated ??18O-SST and Sr/Ca-SST relationships. Calibration differences are most likely due to variations in seawater chemistry in the continentally influenced waters at Looe Key. Additional geochemical replication studies of Montastraea are needed and should include multiple coral heads from open ocean localities complemented whenever possible by seawater chemistry determinations. Copyright 2006 by the American Geophysical Union.

Paleoceanography