The Limits of Coral Thermal Tolerance in Tropical South Atlantic Coastal Reefs after the Fourth Global Mass Bleaching Event

The Limits of Coral Thermal Tolerance in Tropical South Atlantic Coastal Reefs after the Fourth Global Mass Bleaching Event | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article The Limits of Coral Thermal Tolerance in Tropical South Atlantic Coastal Reefs after the Fourth Global Mass Bleaching Event Thales Jean Vidal, Ágatha Naiara Ninow, Ana Lídia Gaspar, Beatriz Fernandes, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9408247/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract The increase in the frequency and intensity of thermal anomalies, combined with local pressures, has reshaped our understanding of the thermal tolerance potential of coral species on a global scale. In this study, through detailed monitoring, we tracked the health trajectories of 290 colonies from eight reef-building species along Brazil’s northeastern coast before, during, and after the fourth global mass bleaching event. Our results reveal striking differences in species vulnerability, fine-scale spatial variability in mortality patterns, no significant variation in the response of colonies of different sizes, and marked heterogeneity in algal colonization on dead skeletons, with filamentous algae predominating nine months after the onset of bleaching. In addition, by considering heatwaves events with different levels of Degree Heating Weeks, we assessed the responses of five species based on more than 3,000 colony records collected throughout 2016 and 2024 along fixed transects and tracked colonies. The findings indicate that intensifying thermal stress has pushed some species close to their survival thresholds, reinforcing that reefs are undergoing biodiversity loss and will likely become dominated by thermotolerant species in the near future. Overall, our results underscore the vulnerability of South Atlantic coastal reefs to climate change and highlight species-specific differences in thermal tolerance, with important implications for the future composition and resilience of these ecosystems. Bleaching Climate change Coastal coral reefs Thresholds Thermal Tolerance Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction Coastal coral reefs are globally significant socioecological systems, supporting a wide array of ecosystem goods and services, including fisheries and tourism (Giglio et al., 2024 ; Silveira & Ferreira, 2024 ). Yet, they are severely impacted by accelerating climate change and compounding local stressors (Hoegh-Guldberg, 2011 ; Wolff et al., 2018 ). Historically, agricultural runoff and sedimentation have been predominant drivers of degradation in nearshore systems (Fabricius, 2005 ; Lapointe et al., 2019 ; Lewis et al., 2021 ), whereas tourism and urban development represent more recent but rapidly escalating pressures (Martinez et al., 2022 ). In parallel, global drivers such as climate change have intensified the frequency and severity of bleaching events (Hughes et al., 2018 ). As pressure accumulates and recovery opportunities narrow (Hughes et al., 2017 and 2018 ; Ortiz et al., 2018 ; Robinson et al., 2019 ), a refined understanding of coral species’ stress-tolerance remains critical. Species diversity in reef ecosystems is primarily shaped by environmental conditions (e.g., temperature, salinity, light, turbidity) (Francini-Filho et al., 2013 ), physical and spatial patterns (Hughes et al., 2018 ; Cardoso et al., 2025 ). Consequently, each species develops life-history strategies suited to the environmental context in which it occurs (e.g. temperature fluctuations, light exposure) (Anthony and Fabricius, 2000 ; Oliver & Palumbi, 2011 ; Darling et al., 2012 ). During and after heatwaves, certain intrinsic mechanisms may confer advantages, such as association with thermotolerant symbiodiniaceae (Fabricius et al., 2004 ; Abrego et al., 2008 ; Hoadley et al., 2019 ), reproduction strategies (Darling et al., 2012 ) and heterotrophic capacity (Grottoli et al., 2006 ; Borell et al., 2008 ). Additionally, subtle variations in light exposure, depth, and hydrodynamic flow can buffer temperature fluctuations and favor coral survival (Morgan et al., 2017 ; Sully & Van Woesik, 2020 ; Yadav et al., 2023 ; Vidal et al., 2025 ). In general, branching species tend to be less thermotolerant than massive corals (Loya et al., 2001 ; Van Woesik et al., 2011 ; Wooldridge et al., 2014). However, few studies have comprehensively assessed local coral diversity and described species-specific thermal tolerance, hindering the evaluation of vulnerability under current and future climate scenarios (Darling et al., 2012 ; Swain et al., 2016 , Edmunds et al., 2021 ). In the Brazilian coastal reefs, the high endemism and predominance of massive corals have led to the hypothesis that they are less vulnerable (Mies et al., 2020 ). Nevertheless, important shallow reefs exhibit high cover of hydrocorals, particularly Millepora alcicornis , the only major branching reef-building species in the Western South Atlantic (WSA), which can occupy extensive areas and provide structural complexity and refugia in coastal systems (Laborel, 1970 ; Ferreira & Maida, 2006 ; Da Silveira et al., 2021 ). Moreover, mass mortality events affecting key reef-building species have become increasingly frequent as climate change progresses (Duarte et al., 2020 ; Bauman et al., 2022; Corazza et al., 2024 ; Vidal et al., 2025 ; Mies et al., 2025 ), including thermotolerant massive corals and endemic species of the genus Mussismilia (Cardoso et al., 2025 ). Along Brazil’s northeastern coast, the frequency and intensity of bleaching events have markedly increased over the last decade (Destri et al., 2025 ). Since 1998, five major events have been recorded, four of them global (1998, 2010, 2016/2017, and 2024) and one local (2019/2020) (Vidal et al., 2025 ). Event severity is largely driven by the persistence, intensity, and frequency of heatwaves (Rodrigues et al., 2025 ). In 2024, record thermal stress values were observed (21.37°C-weeks) (NOAA, 2025), reflecting a global trend of increasing event severity. Therefore, elucidating species-specific stress tolerance thresholds under extreme thermal regimes is essential to guide adaptive management strategies and strengthen the resilience of these vulnerable reef ecosystems. In this study, we closely monitored the health of eight major reef-building species in a Brazilian coastal coral reefs complex during the largest global mass bleaching event on record (2023/2024). By following individual colonies, we tracked bleaching, mortality, and recovery before, during, and after the event. We also used predictive models to evaluate the influence of morphological and spatial variables on fine-scale mortality patterns and to identify species-specific thresholds under events of varying intensity. Finally, we assessed how corals are being replaced by quantifying the organisms colonizing the dead skeletons of the different species studied. 2. Materials and methods 2.1 Study area The Tamandaré Reef Complex (8.73°S, 35.06°W), in southern Pernambuco, northeastern Brazil, is part of a mosaic of protection regimes, including three sustainable-use MPAs and the No-Take Zone Tamandaré Fortress Marine Park (NTZ) (Fig. 1 a/b). It comprises fringing-like reef lines located within 1 km of the coast, at depths between 3–20 m (Ferreira & Maida, 2006 ; Laborel, 1970 ). Reef building is dominated by crustose coralline algae and Millepora alcicornis , followed by scleractinians such as Montastraea cavernosa and Mussismilia harttii (Laborel, 1970 ; Ferreira & Maida, 2006 ; Da Silveira et al., 2021 ). The region is mesotidal (0–2.5 m) with wave-driven circulation, a humid tropical climate (27°C; 2050 mm/year), and seasonal peaks in rainfall, wind, and wave energy from June to August (Maida & Ferreira, 1997 ; Ramos et al., 2009; Cavalcante de Macêdo, 2009; Domingues et al., 2017; Schettini et al., 2017). Hard coral cover has suffered a decline in the last half century (Ferreira & Maida, 2006 ; Ferreira et al., 2021 ), a loss believed to be associated to past overexploitation and increased sedimentation due to agricultural runoff (Maida and Ferreira, 1997 ) and prompted the creation of MPAs in the late 90's as those reefs are ecologically and socioeconomically important to local economy, supporting fishing and tourism (Ferreira et al., 2001; Da Silveira & Ferreira, 2024 ). 2.2 Fourth global mass bleaching event The thermal anomaly observed in 2024 was the most intense and prolonged reported for the region in recent decades. Seawater temperature reached 30°C, with a maximum Degree Heating Weeks (DHW) of 21.37°C-weeks (NOAA, 2025). To assess the impact of this event, we individually monitored the health of 290 colonies from eight species (Table S1 ): Millepora alcicornis (branching); Mussismilia harttii (phacelloid); Agaricia humilis (weedy); Scolymia wellsi (weedy); Porites astreoides (weedy); Montastraea cavernosa (massive); Mussismilia hispida (massive) and Siderastrea sp. (massive). Each colony was surveyed and tagged by attaching a plastic cable tie to rock crevices near its base, and georeferenced between November 2023 and February 2024. The first bleaching records were observed in early March 2024. Continuous monitoring was carried out throughout the year, totaling eight campaigns until December 2024 (Fig. 3 ). Some colonies could not be sampled every month due to logistical constraints. However, all colonies included in the results were monitored at least once before, during, and in the final survey. For six species ( M. cavernosa, M. alcicornis, M. hispida, M. harttii, A. humilis , and S. wellsi ), sampling was conducted using video and photographic surveys covering the full colony area at each sampling event. For Siderastrea sp. and P. astreoides , given the higher abundance of smaller colonies (i.e. < 20 cm), we used fixed 80 m² video transects positioned at the same locations based on in situ reference points, marking colonies along the transects to efficiently quantify community health. Based on image analyses, the health status of each colony (%) was determined using three visually distinct categories: healthy, bleached, and dead. Portions of the colony with Coral Health Chart scores ≤ 3 were classified as bleached, encompassing both initial paling and severe bleaching stages. Mortality was assigned when colony tissue was absent and the exposed skeleton became overgrown by other organisms (Fig. 2 ). The organisms that colonized the dead skeletons were identified for the species M. cavernosa , M. alcicornis , M. hispida , M. harttii , A. humilis , and S. wellsi during the last sampling (Dec/2024), conducted nine months after the onset of bleaching. These organisms were classified into six main groups: filamentous algae; crustose coralline algae (CCA); turf algae (Turf), composed of multispecific assemblages of early successional algae < 2 cm in height (Connell et al., 2014 ); calcareous turf (Taca), composed of a multispecific morphofunctional group, with calcareous articulated algae ( Jania/Amphiroa ) as dominant, followed by corticated and filamentous algae (Ninow et al. submitted ); fleshy macroalgae, and sponges. For P. astreoides and Siderastrea sp., such descriptions were not possible because the video transect method did not provide the level of detail required to identify algal groups. We generated 3D models from M. cavernosa , using Agisoft Metashape Professional software (Burns et al., 2025), to validate health assessments obtained from image analyses. We analyzed 138 models from 47 M. cavernosa colonies distributed across seven sampled sites, covering pre- ( T0 ), during- ( T1 ), and post-event ( T2 ) scenarios (Fig. 3 ). Models were constructed from overlapping photos taken from multiple angles thus covering the entire colony (House et al., 2018 ). ColorChecker® was used as a scale bar (Cardoso et al., 2025 ). For each model, we delineated the total colony area and subsequently classified subareas into the three categories (healthy, bleached, and dead). The areas (m²) of each category were converted into percentages for comparison with results from the initial image analyses. Due to the clear visual distinction of the health categories, a strong correlation was observed between the two methodologies, confirming the reliability of the direct image analyses (r² = 0.97, p < 0.01) (Fig. S1 , Supplementary Material). Therefore, health data derived from image analyses, for all species, were used for subsequent results. 2.4 Spatial and Oceanographic Parameters All colonies were georeferenced using a GPS (Garmin Etrex 10 2.2) towed by the diver, synchronized with the diver’s watch (Da Silveira et al., 2021 ). Colony depths were recorded in situ using a dive computer (Cressi Leonardo 2.0), and depth values were standardized to a 0.0 tidal condition. Colony size (m²) for M. cavernosa was determined from 3D models by delineating the area of living tissue of each colony prior to bleaching (Cardoso et al., 2025 ). For M. alcicornis , pilot 3D models yielded unsatisfactory results due to the shallow habitat and higher structural complexity of the colonies, therefore, colony size was measured in situ using measuring tapes (length and width). The distance from each colony to the coastline was extracted in software QGIS using their georeferenced positions. Degree Heating Weeks (DHW) values were obtained from the National Oceanic and Atmospheric Administration database (NOAA, 2025). 2.5 Analysis All statistical analyses were conducted in R version 4.4.2 (R Core Team, 2025 ). To identify similarities in the temporal trajectories of mortality among species during the 2024 event, we applied a Principal Component Analysis (PCA) (Jolliffe & Cadima, 2016 ). The PCA was performed on the standardized matrix of mean mortality per species across months, allowing dimensionality reduction and visualization of the dominant patterns in temporal variation. Subsequently, we applied a hierarchical cluster analysis using Euclidean distance and Ward’s linkage (Ward.D2 method) to group species according to the similarity of their mortality trajectories (Borcard et al., 2018 ). The number of clusters was defined by inspecting the dendrogram and the fusion height values. To evaluate how morphological and spatial characteristics (depth and distance from the coast) influenced the progression of health categories during the 2024 event, we fitted Cumulative Link Mixed Models (CLMMs) using the ordinal package in R (Christensen, 2023). The analysis included the two species with the largest sample sizes ( M. alcicornis and M. cavernosa ). Fixed effects comprised colony size, depth, DHW, and distance from the coast, along with their interactions with species. We also included the interaction between depth and distance from the coast to capture joint spatial effects. Random effects accounted for variability among sites and among individual colonies (ID). Health categories were defined as: Healthy; Bleached; 0–25% dead; 26–50% dead; 51–95% dead; and 96–100% dead. Differences in mortality rates among sites were assessed using the nonparametric Kruskal–Wallis test, due to the lack of normality in the data. Considering that the response to increasing DHW differed among the two species tested (p < 0.001), we developed two complementary CLMM modeling approaches to identify DHW thresholds. The first examined the health trajectories of all species monitored during the 2024 event. The second evaluated the responses of M. alcicornis , M. hispida , A. humilis , Siderastrea sp., and P. astreoides to past thermal anomalies based on historical data. For the first set of models, the structure was adjusted according to sample size, data distribution, and health progression observed for each species to ensure robust analyses. For M. alcicornis, M. cavernosa, A. humilis , and Siderastrea sp., random effects incorporated site- and colony-level variation. For M. hispida , colony ID could not be included because many colonies died early and remained in the “96–100% dead” category thereafter; in this case, the model accounted only for variability among sites. For S. wellsi , recorded at a single site, only colony ID was used. For M. harttii and P. astreoides , low sample sizes required CLMs without random effects, testing only the influence of DHW. In the second modeling approach, we evaluated species responses across different levels of accumulated thermal stress (maximum DHW) recorded in 2016 (2.62°C-weeks), 2017 (4.64°C-weeks), 2020 (12.07°C-weeks), and 2024 (21.37°C-weeks). Coral health for the 2016–2020 events was reconstructed from historical monitoring in the region (Ferreira et al., 2020, Coxey et al., 2024). In all assessed years, four 20 m transects (spaced one meter apart) were surveyed along the reef crest at sites P1, P4, P5, P6, and P8. The belt-transect method used for corals quantified and classified the health of all colonies within a 50-cm belt along each transect (Ferreira et al., 2018 ). In total, we compiled information on the health of 3,258 colonies across the studied species collected throughout 2016 and 2024. These data were modeled using a CLMM, with health categories (Healthy; 0–25% dead; 26–50% dead; 51–95% dead; and 96–100% dead) as the ordinal response variable, DHW as a fixed effect, and sites and years as random effects. The “bleached” category was not included because these surveys represented post-event conditions. 3. Results During the temperature anomaly period, we observed that all species, at some point, reached nearly 100% bleaching. M. alcicornis was the first species to bleach, with almost its entire population bleached in less than a month. S. wellsi was the only species that remained with high bleaching rates until December. In addition, small fragments of M. alcicornis were also observed bleached until the end of the year. The three most affected species were M. alcicornis , A. humilis , and M. harttii , with mortality rates reaching 99%, 99%, and 93%, respectively, by the end of 2024. M. hispida and S. wellsi experienced mortality rates of 78% and 70%, respectively, followed by P. astreoides with 48%. The most resistant species were M. cavernosa and Siderastrea sp., with mortality rates of 10% and 14%, respectively (Fig. 4 ). Degree heating weeks was a significant factor for all species during the 2024 event (p < 0,001); however, responses to thermal stress varied substantially among taxa (Fig. 4 ; Table S2). Multivariate analyses (PCA and hierarchical clustering) consistently revealed three distinct trajectories of colony health throughout the year, with the first two PCA axes explaining 72.7% and 20.5% of the variation, respectively. These groups captured the main response patterns observed in the field. The first group showed high vulnerability, with a sharp decline until Jun–Jul, including abrupt early mortality in M. alcicornis , M. harttii , and M. hispida . In some cases, M. harttii and M. hispida exhibited partial mortality even before full-colony bleaching, followed by nearly complete recovery from bleaching from August onward. The second group showed high to medium vulnerability, characterized by slow recovery from bleaching and high mortality occurring later in the event (Jun–Jul) in S. wellsi , A. humilis , and P. astreoides . The third group showed low vulnerability, with relatively low mortality in M. cavernosa and Siderastrea sp., and signs of tissue recovery beginning in Jun–Jul (Fig. 4 ). Through the models (CLMM), we identified that depth (p = 0.06), distance from the coast (p = 0.93), and colony size (p = 0.4) did not significantly influence the health categories of M. alcicornis and M. cavernosa (Table S3). Random effects indicated substantial variability between colonies (SD = 1.08) and sites (SD = 0.70). In contrast to the absence of spatial variation in mortality among sites for M. alcicornis (p = 0.23), likely due to the near-total mortality of monitored colonies, M. cavernosa showed a significant difference (p < 0.001). Site P7 (22%) exhibited the highest mortality rates, whereas P2 (2%) and P6 (2%) had the lowest values (Fig. S5). When exposed to events with different maximum DHW scenarios, species also showed distinct trends (Fig. 5 ). M. alcicornis and A. humilis tended to reach a population survival threshold in events with DHW exceeding 20°C-weeks. However, while A. humilis showed an increase in the probability of colonies reaching the 96–100% mortality category only above 15°C-weeks, M. alcicornis exhibited an increased probability of this category even under lower DHW values. M. hispida showed a consistent increase in mortality with rising DHW, reaching probabilities above 60% of colonies being completely dead in events with DHW of 20°C-weeks. For P. astreoides , the probability of transitioning into the most concerning categories showed a slight increasing trend, yet still indicated considerable thermal stress tolerance. Siderastrea sp. was identified as the least affected species, being the only one for which increasing DHW values did not significantly influence health categories over time (Fig. 5 ; Table S4). Overall, we found that filamentous algae were the primary colonizers of dead skeletons in December 2024, nine months after the onset of bleaching, accounting for approximately 48% of total cover across all species. Turf algae, calcareous turf (Taca), and crustose coralline algae (CCA) accounted for 25%, 15%, and 11%, respectively. However, these benthic categories varied markedly among species. For most species, filamentous algae were the predominant colonizers of dead skeletons, with the highest percentages recorded in S. wellsi (66%), M. harttii (54%), A. humilis (51%), and M. alcicornis (45%). Regarding the other benthic categories, calcareous turf was most prevalent on the dead skeletons of M. cavernosa (31%) and M. hispida (36%), reached moderate values in M. harttii (19%), and remained low in all other species. In contrast, crustose coralline algae (CCA) were markedly more abundant on the skeletons of M. alcicornis (31%) than in other taxa, with only moderate values recorded for A. humilis (15%) and M. harttii (13%). Fleshy macroalgae, primarily Dictyotales, were recorded mainly in M. harttii (2.9%), where their cover remained low. Sponges, meanwhile, colonized only a small proportion of M. hispida skeletons (0.2%) (Table 1 ). Table 1 Mean percent cover of organisms on dead skeletons for each species in December 2024. Six main benthic categories were identified: filamentous algae (Fil); crustose coralline algae (CCA); turf algae (Turf); calcareous turf (Taca); fleshy macroalgae (Fleshy); and sponges. For Siderastrea sp. and Porites astreoides , the organisms colonizing dead skeletons were not described (NA). The last column shows the mortality percentages observed for each species. Species Fil(%) CCA(%) Turf(%) Taca(%) Fleshy(%) sponges(%) Mortality(%) M. cavernosa 37.9 4.9 21.7 35.5 0 0 10 M. Hispida 32.8 0.3 30.9 35.8 0 0.2 78 M. Harttii 54.3 13.3 11 18.6 2.9 0 93 A. humilis 51.1 15 32.8 0.6 0.6 0 99 S. wellsi 66 4 30 0 0 0 70 M. alcicornis 44.6 30.7 23.5 1 0.3 0 99 Siderastrea sp. NA NA NA NA NA NA 14 P. astreoides NA NA NA NA NA NA 48 Mean 47.8 11.4 25 15.2 0.6 0.0 64 4. Discussion The intensification and increasing frequency of thermal anomalies have shaped our understanding of coral species’ resistance thresholds worldwide (Hughes et al., 2018 ; Edmunds et al., 2021 ; Marzonie et al., 2023 ). Considering that both intrinsic species traits and extrinsic factors can influence the resilience potential of reef ecosystems (Darling et al., 2012 ), understanding how these elements interact during large-scale bleaching events is essential. However, few studies focus on the response of individual organisms, which limits our understanding of the mechanisms and thermal tolerance thresholds of coral species (Edmunds et al., 2021 ). Our results indicate a clear difference in thermal stress responses among a wide range of coral species along the northeastern Brazilian coast, primarily driven by differences in life-history strategies (Darling et al., 2012 ; Swain et al., 2016 ; Hughes et al., 2018 ; Yadav et al., 2023 ). Following the trend of increased mortality previously reported for the branching hydrocoral M. alcicornis in earlier bleaching events worldwide (Lewis, 2006 ; Dubé et al., 2019 ; Teixeira et al., 2019 ; Duarte et al., 2020 ; Vidal et al., 2025 ), we observed that this species nearly reached its survival threshold during the fourth mass bleaching event on Brazilian coral reefs (Fig. 5 ). Although M. alcicornis is not a scleractinian coral, it plays a key role as a structural framework builder in shallow Brazilian reefs, occurring predominantly on reef crests and shallow submerged reef tops (Laborel, 1970 ; Leão et al., 2003 ). Therefore, future studies are needed to quantify the consequences of this significant reduction in Brazilian reef environments, particularly regarding the loss of structural complexity and the associated fish and invertebrate communities (Coni et al., 2013 ; Graham & Nash, 2013 ). Similarly, M. harttii has undergone marked population declines in recent years (Braz et al., 2022 ; Pereira et al., 2022 ) and is currently listed as “threatened” (IUCN). This phaceloid species can form large colonies and experience rapid skeletal degradation after mortality (Braz et al., 2022 ). In contrast, the massive species M. brasiliensis and M. hispida have previously been described as more thermotolerant during bleaching events (Duarte et al., 2020 ). However, alongside the high mortality observed for M. harttii , we also recorded a substantial increase in M. hispida mortality since 2016, raising concerns about its resistance to intensifying thermal stress in the northwest coast. A similar post-bleaching mortality pattern was reported for M. brasiliensis in the Abrolhos Archipelago after the 2019/2020 event (Cardoso et al., 2025 ). Both Mussismilia species rely on a broadcast-spawning reproductive strategy (Pires et al., 1999 ; Neves and Pires, 2002 ), which increases their susceptibility to Allee effects following mass mortality, as reduced colony abundance limits cross-fertilization and ultimately decreases recruitment success (Courchamp et al., 1999 ; Knowlton, 1992 ; Darling et al., 2012 ). Regarding the so-called “weedy” species ( A. humilis, S. wellsi, and P. asteroides ), in spite of their variable susceptibility to bleaching. their brooding reproductive strategy (McGuire, 1998 ) may support faster post-stress recovery (Darling et al., 2012 ; Goodbody-Gringley & Putron, 2016 ), and opportunistic pattern, increasing in reef cover following the decline of more sensitive corals (Green et al., 2008 ; Carlos-Júnior et al., 2026 ). In our study, although these species initially showed resistance, sharp population declines occurred under high thermal stress (DHW > 15°C-weeks). A. humilis , a species commonly associated with reef walls, approached its regional survival threshold (Fig. 5 ), raising concerns about the time required for substantial population recovery. S. wellsi , was the only species that remained bleached for an extended period (eight months), a pattern that may be linked to heterotrophic plasticity documented in several coral species under thermal stress (Anthony & Fabricius, 2000 ; Grottoli et al., 2006 ; Hughes & Grottoli, 2013 ). Nevertheless, it still experienced high mortality toward the end of the event. In addition, this species has a restricted distribution range, commonly found at the base of reef formations (Francini-Filho et al., 2013 ), which may limit its recolonization following high-mortality events, as observed here. P. astreoides , although showing a slight shift toward more critical health categories since 2016 (Fig. 5 ), exhibited a mortality rate approaching 50% during the fourth global bleaching event. Historically, this species has exhibited high survival and substantial population growth following thermal stress, often dominating reefs where more sensitive taxa experienced severe losses (Green et al., 2008 ). However, despite its “weedy” life-history traits (high fecundity, relatively fast growth, and tolerance to adverse conditions) recent population declines have been reported on Caribbean reefs as climate change accelerates (Edmunds et al., 2021 ), a pattern mirrored in our observations. Additionally, the predominance of medium and small colonies further suggests a young population largely shaped by recent recruitment (Green et al., 2008 ), potentially reflecting population declines during previous events (Edmunds et al., 2021 ). A similar colony size structure was observed for Siderastrea sp. , also likely reflecting the loss of larger colonies and the predominance of a relatively young population in this species. As reported for other regions, some coral species have been classified as more resistant under global change (Van Woesik et al., 2011 ; Darling et al., 2012 ; Grottoli et al., 2014 ). Along the Brazilian coast, our results similarly indicate that Mussismilia cavernosa and Siderastrea sp. exhibit generalist strategies that favor persistence under increasing thermal stress, although through distinct mechanisms. M. cavernosa commonly occurs on reef walls, where lower light availability and higher suspended particulate matter prevail (Francini-Filho et al., 2013 ), and its high heterotrophic plasticity (Lesser et al., 2010 ) may provide a metabolic advantage during bleaching events (Hughes & Grottoli, 2013 ). In contrast, Siderastrea sp., typically found in shallow habitats with high thermal and irradiance variability, appears well adapted to extreme temperature fluctuations (Castillo et al., 2012 ). Additionally, the microbial communities associated with these species may further contribute to their thermal tolerance (Abrego et al., 2008 ; Oliver & Palumbi, 2011 ; Davies et al., 2018 ). Susceptibility to bleaching and mortality in colonies of different sizes within the same species remains a controversial topic and appears to vary among the taxa studied, potentially being positive, negative, or neutral with increasing colony size (Edmunds et al., 2021 ; Winslow et al., 2024 ; Álvarez-Noriega et al., 2025 ). For the two species tested here ( M. alcicornis and M. cavernosa ), we did not observe variability in health patterns (healthy, bleached, and dead) across colony sizes during the 2024 bleaching event. A similar lack of size-related variability was previously reported for M. alcicornis , M. cavernosa , and M. braziliensis during the 2019/2020 bleaching event along the Brazilian coast (Vidal et al., 2025 ; Cardoso et al., 2025 ). In addition to the species-specific characteristics, extrinsic factors such as water circulation, depth, and light irradiance can attenuate thermal variability and alleviate stress on corals (Jokiel et al., 2004; Loya et al., 2016 ; Yadav et al., 2023 ). In the Tamandaré reef complex, we did not detect depth-related variability in bleaching or mortality of M. alcicornis and M. cavernosa . In contrast to the patterns reported for 2019/2020 (Vidal et al., 2025 ), the lack of spatial structuring in M. alcicornis mortality suggests that the intensity of thermal stress exceeded the buffering capacity typically provided by fine-scale environmental heterogeneity for this species. However, the spatial variability observed for M. cavernosa during the 2024 event was highly similar to that previously described for M. alcicornis during the 2019/2020 event (Vidal et al., 2025 ), with sites located in the southern portion of the reef complex, such as P1, P2, and P6, showing lower mortality rates, whereas P7 exhibited higher rates. This pattern indicates the presence of distinct areas of vulnerability at a fine spatial scale in response to thermal stress events. Further investigations integrating colony-level genetic characteristics and local circulation dynamics are needed to better elucidate the environmental factors underlying the spatial patterns observed. Following mortality, rapid colonization by microorganisms (Leggat et al., 2019 ) and later by macroalgae (Diaz-Pulido & McCook, 2002 ) accelerates the structural erosion of colonies (Bleuel et al., 2021 ; Braz et al., 2022 ) and hinders the settlement of new individuals (Doropoulos et al., 2016 ). Consequently, the widespread replacement of living colonies by filamentous algae represents an obstacle to the recovery of the most affected species. In turn, the presence of crustose coralline algae covering a substantial proportion of M. alcicornis skeletons may favor the long-term persistence of colony structure and promote the settlement of new recruits on these skeletons (Doropoulos et al., 2017 ). However, the recovery potential of coral populations may be compromised by local environmental pressures, particularly declining water quality and increasing sedimentation and contamination associated with agricultural runoff, uncontrolled population growth, and urban expansion in coastal zones, as observed in the study region (Silveira & Ferreira, 2024 ; Vidal, 2024 ). These conditions may further favor the establishment of opportunistic algae, intensify competition with corals, reduce recruitment success, and hinder effective recovery (Fabricius, 2005 ; Ricardo et al., 2016 ; MacNeil et al., 2019 ). Therefore, in addition to global policy measures aimed at mitigating climate change, local management actions are essential to ensure the long-term health and resilience of coral reef ecosystems (Richmond et al., 2019 ). 5. Conclusion We observed a clear divergence in health trajectories among species during the fourth global mass bleaching event, driven primarily by their intrinsic characteristics. Our results reveal the collapse of key species responsible for maintaining the structural complexity and diversity of Brazilian reefs, such as M. alcicornis, M. harttii, and A. humilis . We also found moderate to high mortality in species previously considered thermotolerant, including M. hispida and P. astreoides , indicating that climate change is reshaping the thermal tolerance patterns once described for these taxa. In contrast, M. cavernosa and Siderastrea sp. stood out for their high tolerance to thermal stress, suggesting that they may become increasingly dominant reef builders in the future, although long term mortality may be investigated. We detected no variation in mortality among colonies of different sizes and subtle fine-scale spatial patterns, underscoring the severity and homogenizing effect of the event. Our findings also reveal variability in the composition of algal communities colonizing the dead skeletons of different species, with a predominance of filamentous algae, which may pose a substantial challenge to the recovery of species that experienced sharp declines. Collectively, our results highlight the vulnerability of South Atlantic coastal reefs to accelerating climate change, the future risk of local disappearance of species that approached their thermal tolerance limits during the fourth global mass bleaching event, and the accumulation of local and global pressures that may compromise the long-term resilience of coral populations along the northeastern coast of Brazil. Declarations Authors declare they have no competing interests that are relevant to the content of this article. Conflicts of Interest The authors declare no conflicts of interest. Author Contribution Thales J. Vidal: conceptualization, data collection and curation, formal analysis, methodology, visualization, writing – original draft. Ágatha N. Ninow: conceptualization, methodology, data collection and curation, writing – review and editing. Ana Lidia Gaspar: conceptualization, funding acquisition, methodology, project administration, resources, supervision, writing – review and editing. Beatriz Fernandes: data collection; methodology, visualization, writing – review and editing. Thiago Buchianeri: data collection; methodology, visualization, writing – review and editing. Mauro Maida: conceptualization, funding acquisition, methodology, project administration, resources, supervision, writing – review and editing. Leonardo Messias: conceptualization, funding acquisition, resources, writing – review and editing. Camila da Silveira Brasil: data collection, methodology, writing – review and editing. Beatrice P. Ferreira: conceptualization, data curation, funding acquisition, methodology, project administration, resources, supervision, writing – review and editing. Acknowledgement This is a Tamandaré Long-Term Ecological Program (PELD-TAMS, ILTER site 18) contribution. We thank the National Center for Research and Conservation of Marine Biodiversity in the Northeast (CEPENE-Tamandaré ICMBio) for invaluable field and logistic support. The Coastal Reefs Institute (IRCOS), the Tamandare Cave Project (CECAV), and WWF-Brasil (CPT 003776-2024) for their financial and logistical support during field trips and collections. We thank the Brazilian scientific council CNPQ for a grant to BPF (442139/2020-9). We also thank the Post Graduate Program in Oceanography at the Federal University of Pernambuco (PPGO/UFPE). All sampling activities were conducted under the permits issued by SISBIO for the Long-Term Biodiversity Monitoring Program (No. 45992) and Reef Check Brasil (No. 18355). Data Availability The datasets generated and/or analysed during the current study are available from the corresponding author upon reasonable request. References Abrego, D., Ulstrup, K. E., Willis, B. L., & van Oppen, M. J. (2008). 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Mortality patterns and recovery challenges in Millepora alcicornis after mass bleaching event on Northeast Brazilian reefs. Marine Environmental Research, 204, 106864. https://doi.org/10.1016/j.marenvres.2024.106864 Winslow, E. M., Speare, K. E., Adam, T. C., Burkepile, D. E., Hench, J. L., & Lenihan, H. S. (2024). Corals survive severe bleaching event in refuges related to taxa, colony size, and water depth. Scientific Reports, 14(1), 9006. https://doi.org/10.1038/s41598-024-58980-1 Wooldridge, S. A. (2014). Differential thermal bleaching susceptibilities amongst coral taxa: re-posing the role of the host. Coral reefs, 33(1), 15–27. https://doi.org/10.1007/s00338-013-1111-4 Wolff, N. H., Mumby, P. J., Devlin, M., & Anthony, K. R. (2018). Vulnerability of the Great Barrier Reef to climate change and local pressures. Global change biology, 24(5), 1978–1991. https://doi.org/10.1111/gcb.14043 Yadav, S., Roach, T. N., McWilliam, M. J., Caruso, C., de Souza, M. R., Foley, C., … Madin, J. S. (2023). Fine-scale variability in coral bleaching and mortality during a marine heatwave. Frontiers in Marine Science, 10, 1108365. https://doi.org/10.3389/fmars.2023.1108365 Additional Declarations No competing interests reported. Supplementary Files Supplementarymaterial.docx Graphicalabstract.jpg Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 17 May, 2026 Reviewers agreed at journal 14 May, 2026 Reviewers agreed at journal 14 May, 2026 Reviewers agreed at journal 13 May, 2026 Reviewers agreed at journal 30 Apr, 2026 Reviewers invited by journal 28 Apr, 2026 Editor assigned by journal 17 Apr, 2026 Submission checks completed at journal 15 Apr, 2026 First submitted to journal 13 Apr, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9408247","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":633047146,"identity":"183222a1-58ce-40fc-98d3-5f8e9368f542","order_by":0,"name":"Thales Jean Vidal","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA40lEQVRIie3QsQqCQBzH8b8Euly1XhT2CgdOQdGrKA4tGYLQLATXks32FvYEKQeOzY1OTQ2ODTd0BxVNp20R9138LR+8OwCd7hfDYOTya4Jr5DXYYiK1QBjgRaBIwfmCgCAMtSHz4abKaw7e3vIrNqNk1YduWSv/MipJcaDgUXQlLKAkGsQ9P1UfzCWsG4NjyhFQ7mU5chrusqgZ55KIMaHEOzWTJWHiuWxTDkOQDJrIZRkWCcW2iW5hsTuTCLOeryRWujhWdz5F460ca/Fi24QpyTP8Xi502oDP3G+BTqfT/X8PPz5H6rXFI9gAAAAASUVORK5CYII=","orcid":"","institution":"Instituto Recifes Costeiros","correspondingAuthor":true,"prefix":"","firstName":"Thales","middleName":"Jean","lastName":"Vidal","suffix":""},{"id":633047147,"identity":"4f77d818-ad6e-456e-9f09-e7025a89cc4a","order_by":1,"name":"Ágatha Naiara Ninow","email":"","orcid":"","institution":"National Center for Research and Conservation of Marine Biodiversity in the Northeast","correspondingAuthor":false,"prefix":"","firstName":"Ágatha","middleName":"Naiara","lastName":"Ninow","suffix":""},{"id":633047148,"identity":"4e2f3346-bcc7-49b0-b8f6-c05e15b1ec73","order_by":2,"name":"Ana Lídia Gaspar","email":"","orcid":"","institution":"Instituto Recifes Costeiros","correspondingAuthor":false,"prefix":"","firstName":"Ana","middleName":"Lídia","lastName":"Gaspar","suffix":""},{"id":633047149,"identity":"e85bcd3e-cb1d-4ae8-a6e6-68825af39f69","order_by":3,"name":"Beatriz Fernandes","email":"","orcid":"","institution":"Federal University of Pernambuco","correspondingAuthor":false,"prefix":"","firstName":"Beatriz","middleName":"","lastName":"Fernandes","suffix":""},{"id":633047150,"identity":"187c316d-9df7-44d6-8cf3-c2404b4055ab","order_by":4,"name":"Thiago Buchianeri","email":"","orcid":"","institution":"University of São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Thiago","middleName":"","lastName":"Buchianeri","suffix":""},{"id":633047151,"identity":"e504c418-593b-4cb7-ae43-cc58208b753b","order_by":5,"name":"Mauro Maida","email":"","orcid":"","institution":"Federal University of Pernambuco","correspondingAuthor":false,"prefix":"","firstName":"Mauro","middleName":"","lastName":"Maida","suffix":""},{"id":633047152,"identity":"49ff32e7-84ef-4c47-b85a-b385d107ed02","order_by":6,"name":"Leonardo Messias","email":"","orcid":"","institution":"National Center for Research and Conservation of Marine Biodiversity in the Northeast","correspondingAuthor":false,"prefix":"","firstName":"Leonardo","middleName":"","lastName":"Messias","suffix":""},{"id":633047153,"identity":"1b3034f0-1f6b-4cc0-ae7f-9e8376527dc7","order_by":7,"name":"Camila Brasil da Silveira","email":"","orcid":"","institution":"Federal University of Bahia","correspondingAuthor":false,"prefix":"","firstName":"Camila","middleName":"Brasil da","lastName":"Silveira","suffix":""},{"id":633047154,"identity":"19f71662-32ed-4443-b4b4-adb521b1e608","order_by":8,"name":"Beatrice Padovani Ferreira","email":"","orcid":"","institution":"Federal University of Pernambuco","correspondingAuthor":false,"prefix":"","firstName":"Beatrice","middleName":"Padovani","lastName":"Ferreira","suffix":""}],"badges":[],"createdAt":"2026-04-13 20:39:50","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9408247/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9408247/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108609185,"identity":"7498cfe2-42c6-4318-8296-7f8fe0591fea","added_by":"auto","created_at":"2026-05-06 12:47:55","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":303992,"visible":true,"origin":"","legend":"\u003cp\u003eStudy area a) Mosaic of protected areas encompassing the Tamandaré reef complex, including three Environmental Protection Areas (APAs): Costa dos Corais (APACC), Guadalupe, and Serrambi. Each colored quadrant represents a specific protection regime. b) Distribution of the eight sampled sites. Black dots indicate sites where both methodologies were applied: tagged colonies, video transect and coral surveys transect. Sites marked with a cross represent locations where only tagged colonies were sampled. The red dashed quadrant outlines the boundaries of the No-Take Zone (NTZ). c) Zoomed-in views show the georeferenced positions of tagged colonies at each site, with different colors representing species.\u003c/p\u003e","description":"","filename":"Figure1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/1243849d81e552f2690c4036.jpg"},{"id":108804849,"identity":"ace70697-5495-489c-803d-b388a19229a2","added_by":"auto","created_at":"2026-05-08 15:23:56","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1112481,"visible":true,"origin":"","legend":"\u003cp\u003eImage illustrating health categories for each species. The corals showed are indicated with letters (a-g): \u003cem\u003eMillepora alcicornis (a); Montastraea cavernosa (b); Mussismilia hispida and Agaricia humilis (c); Mussismilia harttii (d); Porites astreoides (e); Scolymia wellsi (f); Siderastrea \u003c/em\u003esp\u003cem\u003e. (g)\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/9c7e1ad9f77c15d28d18c8b8.jpg"},{"id":108805983,"identity":"5524a667-df4e-475b-8902-5c3b3a37a08c","added_by":"auto","created_at":"2026-05-08 15:27:22","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":119645,"visible":true,"origin":"","legend":"\u003cp\u003eMonitoring of tagged \u003cem\u003eMontastraea cavernosa \u003c/em\u003ecolonies using 3D photogrammetry models. The white area within the colony indicates the absence of living tissue, and its progression across the images reflects tissue mortality. The monitored colony is shown at different periods throughout the year, transitioning through healthy (Mar/2024), bleached (Apr/2024), bleached with parcial mortality (Jun/2024), and healthy with parcial mortality (Dec/2024) conditions.\u003c/p\u003e","description":"","filename":"Figure3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/d990d611c1990040da54a01d.jpg"},{"id":108609188,"identity":"18fd78d4-263b-4d20-8e6a-0a225813da58","added_by":"auto","created_at":"2026-05-06 12:47:55","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":520144,"visible":true,"origin":"","legend":"\u003cp\u003eProgression of mortality by species. Each bar represents a sampling event, with colony status distributed among three categories: Healthy (light green), Bleached (gray), and Dead (dark green). For \u003cem\u003eSiderastrea\u003c/em\u003e sp. and \u003cem\u003ePorites astreoides\u003c/em\u003e, only three periods are shown: before (Feb/24), during (Mar/24), and after the event (Dec/24).\u003c/p\u003e","description":"","filename":"Figure4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/5492b06f59f1b0d8d96bc04c.jpg"},{"id":108609189,"identity":"24b82300-b56f-459e-b5d2-9b55e2b39233","added_by":"auto","created_at":"2026-05-06 12:47:55","extension":"jpeg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":101113,"visible":true,"origin":"","legend":"\u003cp\u003eModel predictions of the probability that colonies of five studied species (n = 3,515) exhibited different health states based on Degree Heating Weeks (°C/weeks) observed during four thermal anomaly events: 2016 (2.62 °C/weeks), 2017 (4.64 °C/weeks), 2020 (12.07 °C/weeks), and 2024 (21.37 °C/weeks). Each color represents a health category, with lighter shades indicating healthy colonies and darker shades indicating colonies with 96–100% mortality.\u003c/p\u003e","description":"","filename":"Figure5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/7e33c17cfc89be1bf09a1ca2.jpeg"},{"id":108811632,"identity":"6d073cbb-6f34-4d16-ae82-e2172a2077cc","added_by":"auto","created_at":"2026-05-08 16:06:20","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2608097,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/55c8fa20-b1df-4155-8299-aaf980c377b1.pdf"},{"id":108804718,"identity":"531f2cbe-1d8d-485f-ae0f-0577dc6692f1","added_by":"auto","created_at":"2026-05-08 15:22:56","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":993660,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/2f8b099fbc7a55b989f76f74.docx"},{"id":108609186,"identity":"eb0b19d5-9fb0-4fd1-ab33-456899b23414","added_by":"auto","created_at":"2026-05-06 12:47:55","extension":"jpg","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":125878,"visible":true,"origin":"","legend":"","description":"","filename":"Graphicalabstract.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9408247/v1/b955984ba3de489501614131.jpg"}],"financialInterests":"No competing interests reported.","formattedTitle":"The Limits of Coral Thermal Tolerance in Tropical South Atlantic Coastal Reefs after the Fourth Global Mass Bleaching Event","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eCoastal coral reefs are globally significant socioecological systems, supporting a wide array of ecosystem goods and services, including fisheries and tourism (Giglio et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Silveira \u0026amp; Ferreira, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Yet, they are severely impacted by accelerating climate change and compounding local stressors (Hoegh-Guldberg, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Wolff et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Historically, agricultural runoff and sedimentation have been predominant drivers of degradation in nearshore systems (Fabricius, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Lapointe et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Lewis et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), whereas tourism and urban development represent more recent but rapidly escalating pressures (Martinez et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In parallel, global drivers such as climate change have intensified the frequency and severity of bleaching events (Hughes et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). As pressure accumulates and recovery opportunities narrow (Hughes et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2017\u003c/span\u003e and \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Ortiz et al., \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Robinson et al., \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), a refined understanding of coral species\u0026rsquo; stress-tolerance remains critical.\u003c/p\u003e \u003cp\u003eSpecies diversity in reef ecosystems is primarily shaped by environmental conditions (e.g., temperature, salinity, light, turbidity) (Francini-Filho et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), physical and spatial patterns (Hughes et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Consequently, each species develops life-history strategies suited to the environmental context in which it occurs (e.g. temperature fluctuations, light exposure) (Anthony and Fabricius, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Oliver \u0026amp; Palumbi, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). During and after heatwaves, certain intrinsic mechanisms may confer advantages, such as association with thermotolerant symbiodiniaceae (Fabricius et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Abrego et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Hoadley et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), reproduction strategies (Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and heterotrophic capacity (Grottoli et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Borell et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Additionally, subtle variations in light exposure, depth, and hydrodynamic flow can buffer temperature fluctuations and favor coral survival (Morgan et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Sully \u0026amp; Van Woesik, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Yadav et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn general, branching species tend to be less thermotolerant than massive corals (Loya et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Van Woesik et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Wooldridge et al., 2014). However, few studies have comprehensively assessed local coral diversity and described species-specific thermal tolerance, hindering the evaluation of vulnerability under current and future climate scenarios (Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Swain et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In the Brazilian coastal reefs, the high endemism and predominance of massive corals have led to the hypothesis that they are less vulnerable (Mies et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Nevertheless, important shallow reefs exhibit high cover of hydrocorals, particularly \u003cem\u003eMillepora alcicornis\u003c/em\u003e, the only major branching reef-building species in the Western South Atlantic (WSA), which can occupy extensive areas and provide structural complexity and refugia in coastal systems (Laborel, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Ferreira \u0026amp; Maida, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Da Silveira et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Moreover, mass mortality events affecting key reef-building species have become increasingly frequent as climate change progresses (Duarte et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bauman et al., 2022; Corazza et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Mies et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), including thermotolerant massive corals and endemic species of the genus \u003cem\u003eMussismilia\u003c/em\u003e (Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlong Brazil\u0026rsquo;s northeastern coast, the frequency and intensity of bleaching events have markedly increased over the last decade (Destri et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Since 1998, five major events have been recorded, four of them global (1998, 2010, 2016/2017, and 2024) and one local (2019/2020) (Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Event severity is largely driven by the persistence, intensity, and frequency of heatwaves (Rodrigues et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In 2024, record thermal stress values were observed (21.37\u0026deg;C-weeks) (NOAA, 2025), reflecting a global trend of increasing event severity. Therefore, elucidating species-specific stress tolerance thresholds under extreme thermal regimes is essential to guide adaptive management strategies and strengthen the resilience of these vulnerable reef ecosystems.\u003c/p\u003e \u003cp\u003eIn this study, we closely monitored the health of eight major reef-building species in a Brazilian coastal coral reefs complex during the largest global mass bleaching event on record (2023/2024). By following individual colonies, we tracked bleaching, mortality, and recovery before, during, and after the event. We also used predictive models to evaluate the influence of morphological and spatial variables on fine-scale mortality patterns and to identify species-specific thresholds under events of varying intensity. Finally, we assessed how corals are being replaced by quantifying the organisms colonizing the dead skeletons of the different species studied.\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Study area\u003c/h2\u003e \u003cp\u003eThe Tamandar\u0026eacute; Reef Complex (8.73\u0026deg;S, 35.06\u0026deg;W), in southern Pernambuco, northeastern Brazil, is part of a mosaic of protection regimes, including three sustainable-use MPAs and the No-Take Zone Tamandar\u0026eacute; Fortress Marine Park (NTZ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea/b). It comprises fringing-like reef lines located within 1 km of the coast, at depths between 3\u0026ndash;20 m (Ferreira \u0026amp; Maida, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Laborel, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1970\u003c/span\u003e). Reef building is dominated by crustose coralline algae and \u003cem\u003eMillepora alcicornis\u003c/em\u003e, followed by scleractinians such as \u003cem\u003eMontastraea cavernosa\u003c/em\u003e and \u003cem\u003eMussismilia harttii\u003c/em\u003e (Laborel, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Ferreira \u0026amp; Maida, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Da Silveira et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The region is mesotidal (0\u0026ndash;2.5 m) with wave-driven circulation, a humid tropical climate (27\u0026deg;C; 2050 mm/year), and seasonal peaks in rainfall, wind, and wave energy from June to August (Maida \u0026amp; Ferreira, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Ramos et al., 2009; Cavalcante de Mac\u0026ecirc;do, 2009; Domingues et al., 2017; Schettini et al., 2017). Hard coral cover has suffered a decline in the last half century (Ferreira \u0026amp; Maida, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ferreira et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), a loss believed to be associated to past overexploitation and increased sedimentation due to agricultural runoff (Maida and Ferreira, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and prompted the creation of MPAs in the late 90's as those reefs are ecologically and socioeconomically important to local economy, supporting fishing and tourism (Ferreira et al., 2001; Da Silveira \u0026amp; Ferreira, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Fourth global mass bleaching event\u003c/h2\u003e \u003cp\u003eThe thermal anomaly observed in 2024 was the most intense and prolonged reported for the region in recent decades. Seawater temperature reached 30\u0026deg;C, with a maximum Degree Heating Weeks (DHW) of 21.37\u0026deg;C-weeks (NOAA, 2025). To assess the impact of this event, we individually monitored the health of 290 colonies from eight species (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e): \u003cem\u003eMillepora alcicornis\u003c/em\u003e (branching); \u003cem\u003eMussismilia harttii\u003c/em\u003e (phacelloid); \u003cem\u003eAgaricia humilis\u003c/em\u003e (weedy); \u003cem\u003eScolymia wellsi\u003c/em\u003e (weedy); \u003cem\u003ePorites astreoides\u003c/em\u003e (weedy); \u003cem\u003eMontastraea cavernosa\u003c/em\u003e (massive); \u003cem\u003eMussismilia hispida\u003c/em\u003e (massive) and \u003cem\u003eSiderastrea\u003c/em\u003e sp. (massive). Each colony was surveyed and tagged by attaching a plastic cable tie to rock crevices near its base, and georeferenced between November 2023 and February 2024. The first bleaching records were observed in early March 2024. Continuous monitoring was carried out throughout the year, totaling eight campaigns until December 2024 (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Some colonies could not be sampled every month due to logistical constraints. However, all colonies included in the results were monitored at least once before, during, and in the final survey.\u003c/p\u003e \u003cp\u003eFor six species (\u003cem\u003eM. cavernosa, M. alcicornis, M. hispida, M. harttii, A. humilis\u003c/em\u003e, and \u003cem\u003eS. wellsi\u003c/em\u003e), sampling was conducted using video and photographic surveys covering the full colony area at each sampling event. For \u003cem\u003eSiderastrea\u003c/em\u003e sp. and \u003cem\u003eP. astreoides\u003c/em\u003e, given the higher abundance of smaller colonies (i.e. \u0026lt; 20 cm), we used fixed 80 m\u0026sup2; video transects positioned at the same locations based on in situ reference points, marking colonies along the transects to efficiently quantify community health. Based on image analyses, the health status of each colony (%) was determined using three visually distinct categories: healthy, bleached, and dead. Portions of the colony with Coral Health Chart scores\u0026thinsp;\u0026le;\u0026thinsp;3 were classified as bleached, encompassing both initial paling and severe bleaching stages. Mortality was assigned when colony tissue was absent and the exposed skeleton became overgrown by other organisms (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe organisms that colonized the dead skeletons were identified for the species \u003cem\u003eM. cavernosa\u003c/em\u003e, \u003cem\u003eM. alcicornis\u003c/em\u003e, \u003cem\u003eM. hispida\u003c/em\u003e, \u003cem\u003eM. harttii\u003c/em\u003e, \u003cem\u003eA. humilis\u003c/em\u003e, and \u003cem\u003eS. wellsi\u003c/em\u003e during the last sampling (Dec/2024), conducted nine months after the onset of bleaching. These organisms were classified into six main groups: filamentous algae; crustose coralline algae (CCA); turf algae (Turf), composed of multispecific assemblages of early successional algae\u0026thinsp;\u0026lt;\u0026thinsp;2 cm in height (Connell et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e); calcareous turf (Taca), composed of a multispecific morphofunctional group, with calcareous articulated algae (\u003cem\u003eJania/Amphiroa\u003c/em\u003e) as dominant, followed by corticated and filamentous algae (Ninow et al. \u003cem\u003esubmitted\u003c/em\u003e); fleshy macroalgae, and sponges. For \u003cem\u003eP. astreoides\u003c/em\u003e and \u003cem\u003eSiderastrea\u003c/em\u003e sp., such descriptions were not possible because the video transect method did not provide the level of detail required to identify algal groups.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWe generated 3D models from \u003cem\u003eM. cavernosa\u003c/em\u003e, using Agisoft Metashape Professional software (Burns et al., 2025), to validate health assessments obtained from image analyses. We analyzed 138 models from 47 \u003cem\u003eM. cavernosa\u003c/em\u003e colonies distributed across seven sampled sites, covering pre- (\u003cem\u003eT0\u003c/em\u003e), during- (\u003cem\u003eT1\u003c/em\u003e), and post-event (\u003cem\u003eT2\u003c/em\u003e) scenarios (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Models were constructed from overlapping photos taken from multiple angles thus covering the entire colony (House et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). ColorChecker\u0026reg; was used as a scale bar (Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For each model, we delineated the total colony area and subsequently classified subareas into the three categories (healthy, bleached, and dead). The areas (m\u0026sup2;) of each category were converted into percentages for comparison with results from the initial image analyses. Due to the clear visual distinction of the health categories, a strong correlation was observed between the two methodologies, confirming the reliability of the direct image analyses (r\u0026sup2; = 0.97, p\u0026thinsp;\u0026lt;\u0026thinsp;0.01) (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e, Supplementary Material). Therefore, health data derived from image analyses, for all species, were used for subsequent results.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Spatial and Oceanographic Parameters\u003c/h2\u003e \u003cp\u003eAll colonies were georeferenced using a GPS (Garmin Etrex 10 2.2) towed by the diver, synchronized with the diver\u0026rsquo;s watch (Da Silveira et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Colony depths were recorded in situ using a dive computer (Cressi Leonardo 2.0), and depth values were standardized to a 0.0 tidal condition. Colony size (m\u0026sup2;) for \u003cem\u003eM. cavernosa\u003c/em\u003e was determined from 3D models by delineating the area of living tissue of each colony prior to bleaching (Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For \u003cem\u003eM. alcicornis\u003c/em\u003e, pilot 3D models yielded unsatisfactory results due to the shallow habitat and higher structural complexity of the colonies, therefore, colony size was measured in situ using measuring tapes (length and width). The distance from each colony to the coastline was extracted in software QGIS using their georeferenced positions. Degree Heating Weeks (DHW) values were obtained from the National Oceanic and Atmospheric Administration database (NOAA, 2025).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.5 Analysis\u003c/h2\u003e \u003cp\u003eAll statistical analyses were conducted in R version 4.4.2 (R Core Team, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo identify similarities in the temporal trajectories of mortality among species during the 2024 event, we applied a Principal Component Analysis (PCA) (Jolliffe \u0026amp; Cadima, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The PCA was performed on the standardized matrix of mean mortality per species across months, allowing dimensionality reduction and visualization of the dominant patterns in temporal variation. Subsequently, we applied a hierarchical cluster analysis using Euclidean distance and Ward\u0026rsquo;s linkage (Ward.D2 method) to group species according to the similarity of their mortality trajectories (Borcard et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The number of clusters was defined by inspecting the dendrogram and the fusion height values.\u003c/p\u003e \u003cp\u003eTo evaluate how morphological and spatial characteristics (depth and distance from the coast) influenced the progression of health categories during the 2024 event, we fitted Cumulative Link Mixed Models (CLMMs) using the \u003cem\u003eordinal\u003c/em\u003e package in R (Christensen, 2023). The analysis included the two species with the largest sample sizes (\u003cem\u003eM. alcicornis\u003c/em\u003e and \u003cem\u003eM. cavernosa\u003c/em\u003e). Fixed effects comprised colony size, depth, DHW, and distance from the coast, along with their interactions with species. We also included the interaction between depth and distance from the coast to capture joint spatial effects. Random effects accounted for variability among sites and among individual colonies (ID). Health categories were defined as: Healthy; Bleached; 0\u0026ndash;25% dead; 26\u0026ndash;50% dead; 51\u0026ndash;95% dead; and 96\u0026ndash;100% dead. Differences in mortality rates among sites were assessed using the nonparametric Kruskal\u0026ndash;Wallis test, due to the lack of normality in the data.\u003c/p\u003e \u003cp\u003eConsidering that the response to increasing DHW differed among the two species tested (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), we developed two complementary CLMM modeling approaches to identify DHW thresholds. The first examined the health trajectories of all species monitored during the 2024 event. The second evaluated the responses of \u003cem\u003eM. alcicornis\u003c/em\u003e, \u003cem\u003eM. hispida\u003c/em\u003e, \u003cem\u003eA. humilis\u003c/em\u003e, \u003cem\u003eSiderastrea\u003c/em\u003e sp., and \u003cem\u003eP. astreoides\u003c/em\u003e to past thermal anomalies based on historical data.\u003c/p\u003e \u003cp\u003eFor the first set of models, the structure was adjusted according to sample size, data distribution, and health progression observed for each species to ensure robust analyses. For \u003cem\u003eM. alcicornis, M. cavernosa, A. humilis\u003c/em\u003e, and \u003cem\u003eSiderastrea\u003c/em\u003e sp., random effects incorporated site- and colony-level variation. For \u003cem\u003eM. hispida\u003c/em\u003e, colony ID could not be included because many colonies died early and remained in the \u0026ldquo;96\u0026ndash;100% dead\u0026rdquo; category thereafter; in this case, the model accounted only for variability among sites. For \u003cem\u003eS. wellsi\u003c/em\u003e, recorded at a single site, only colony ID was used. For \u003cem\u003eM. harttii\u003c/em\u003e and \u003cem\u003eP. astreoides\u003c/em\u003e, low sample sizes required CLMs without random effects, testing only the influence of DHW.\u003c/p\u003e \u003cp\u003eIn the second modeling approach, we evaluated species responses across different levels of accumulated thermal stress (maximum DHW) recorded in 2016 (2.62\u0026deg;C-weeks), 2017 (4.64\u0026deg;C-weeks), 2020 (12.07\u0026deg;C-weeks), and 2024 (21.37\u0026deg;C-weeks). Coral health for the 2016\u0026ndash;2020 events was reconstructed from historical monitoring in the region (Ferreira et al., 2020, Coxey et al., 2024). In all assessed years, four 20 m transects (spaced one meter apart) were surveyed along the reef crest at sites P1, P4, P5, P6, and P8. The belt-transect method used for corals quantified and classified the health of all colonies within a 50-cm belt along each transect (Ferreira et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In total, we compiled information on the health of 3,258 colonies across the studied species collected throughout 2016 and 2024. These data were modeled using a CLMM, with health categories (Healthy; 0\u0026ndash;25% dead; 26\u0026ndash;50% dead; 51\u0026ndash;95% dead; and 96\u0026ndash;100% dead) as the ordinal response variable, DHW as a fixed effect, and sites and years as random effects. The \u0026ldquo;bleached\u0026rdquo; category was not included because these surveys represented post-event conditions.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cp\u003eDuring the temperature anomaly period, we observed that all species, at some point, reached nearly 100% bleaching. \u003cem\u003eM. alcicornis\u003c/em\u003e was the first species to bleach, with almost its entire population bleached in less than a month. \u003cem\u003eS. wellsi\u003c/em\u003e was the only species that remained with high bleaching rates until December. In addition, small fragments of \u003cem\u003eM. alcicornis\u003c/em\u003e were also observed bleached until the end of the year. The three most affected species were \u003cem\u003eM. alcicornis\u003c/em\u003e, \u003cem\u003eA. humilis\u003c/em\u003e, and \u003cem\u003eM. harttii\u003c/em\u003e, with mortality rates reaching 99%, 99%, and 93%, respectively, by the end of 2024. \u003cem\u003eM. hispida\u003c/em\u003e and \u003cem\u003eS. wellsi\u003c/em\u003e experienced mortality rates of 78% and 70%, respectively, followed by \u003cem\u003eP. astreoides\u003c/em\u003e with 48%. The most resistant species were \u003cem\u003eM. cavernosa\u003c/em\u003e and \u003cem\u003eSiderastrea\u003c/em\u003e sp., with mortality rates of 10% and 14%, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDegree heating weeks was a significant factor for all species during the 2024 event (p\u0026thinsp;\u0026lt;\u0026thinsp;0,001); however, responses to thermal stress varied substantially among taxa (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; Table S2). Multivariate analyses (PCA and hierarchical clustering) consistently revealed three distinct trajectories of colony health throughout the year, with the first two PCA axes explaining 72.7% and 20.5% of the variation, respectively. These groups captured the main response patterns observed in the field. The first group showed high vulnerability, with a sharp decline until Jun\u0026ndash;Jul, including abrupt early mortality in \u003cem\u003eM. alcicornis\u003c/em\u003e, \u003cem\u003eM. harttii\u003c/em\u003e, and \u003cem\u003eM. hispida\u003c/em\u003e. In some cases, \u003cem\u003eM. harttii\u003c/em\u003e and \u003cem\u003eM. hispida\u003c/em\u003e exhibited partial mortality even before full-colony bleaching, followed by nearly complete recovery from bleaching from August onward. The second group showed high to medium vulnerability, characterized by slow recovery from bleaching and high mortality occurring later in the event (Jun\u0026ndash;Jul) in \u003cem\u003eS. wellsi\u003c/em\u003e, \u003cem\u003eA. humilis\u003c/em\u003e, and \u003cem\u003eP. astreoides\u003c/em\u003e. The third group showed low vulnerability, with relatively low mortality in \u003cem\u003eM. cavernosa\u003c/em\u003e and \u003cem\u003eSiderastrea\u003c/em\u003e sp., and signs of tissue recovery beginning in Jun\u0026ndash;Jul (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThrough the models (CLMM), we identified that depth (p\u0026thinsp;=\u0026thinsp;0.06), distance from the coast (p\u0026thinsp;=\u0026thinsp;0.93), and colony size (p\u0026thinsp;=\u0026thinsp;0.4) did not significantly influence the health categories of \u003cem\u003eM. alcicornis\u003c/em\u003e and \u003cem\u003eM. cavernosa\u003c/em\u003e (Table S3). Random effects indicated substantial variability between colonies (SD\u0026thinsp;=\u0026thinsp;1.08) and sites (SD\u0026thinsp;=\u0026thinsp;0.70). In contrast to the absence of spatial variation in mortality among sites for \u003cem\u003eM. alcicornis\u003c/em\u003e (p\u0026thinsp;=\u0026thinsp;0.23), likely due to the near-total mortality of monitored colonies, \u003cem\u003eM. cavernosa\u003c/em\u003e showed a significant difference (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Site P7 (22%) exhibited the highest mortality rates, whereas P2 (2%) and P6 (2%) had the lowest values (Fig. S5).\u003c/p\u003e \u003cp\u003eWhen exposed to events with different maximum DHW scenarios, species also showed distinct trends (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). \u003cem\u003eM. alcicornis\u003c/em\u003e and \u003cem\u003eA. humilis\u003c/em\u003e tended to reach a population survival threshold in events with DHW exceeding 20\u0026deg;C-weeks. However, while \u003cem\u003eA. humilis\u003c/em\u003e showed an increase in the probability of colonies reaching the 96\u0026ndash;100% mortality category only above 15\u0026deg;C-weeks, \u003cem\u003eM. alcicornis\u003c/em\u003e exhibited an increased probability of this category even under lower DHW values. \u003cem\u003eM. hispida\u003c/em\u003e showed a consistent increase in mortality with rising DHW, reaching probabilities above 60% of colonies being completely dead in events with DHW of 20\u0026deg;C-weeks. For \u003cem\u003eP. astreoides\u003c/em\u003e, the probability of transitioning into the most concerning categories showed a slight increasing trend, yet still indicated considerable thermal stress tolerance. \u003cem\u003eSiderastrea\u003c/em\u003e sp. was identified as the least affected species, being the only one for which increasing DHW values did not significantly influence health categories over time (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e; Table S4).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eOverall, we found that filamentous algae were the primary colonizers of dead skeletons in December 2024, nine months after the onset of bleaching, accounting for approximately 48% of total cover across all species. Turf algae, calcareous turf (Taca), and crustose coralline algae (CCA) accounted for 25%, 15%, and 11%, respectively. However, these benthic categories varied markedly among species.\u003c/p\u003e \u003cp\u003eFor most species, filamentous algae were the predominant colonizers of dead skeletons, with the highest percentages recorded in \u003cem\u003eS. wellsi\u003c/em\u003e (66%), \u003cem\u003eM. harttii\u003c/em\u003e (54%), \u003cem\u003eA. humilis\u003c/em\u003e (51%), and \u003cem\u003eM. alcicornis\u003c/em\u003e (45%). Regarding the other benthic categories, calcareous turf was most prevalent on the dead skeletons of \u003cem\u003eM. cavernosa\u003c/em\u003e (31%) and \u003cem\u003eM. hispida\u003c/em\u003e (36%), reached moderate values in \u003cem\u003eM. harttii\u003c/em\u003e (19%), and remained low in all other species. In contrast, crustose coralline algae (CCA) were markedly more abundant on the skeletons of \u003cem\u003eM. alcicornis\u003c/em\u003e (31%) than in other taxa, with only moderate values recorded for \u003cem\u003eA. humilis\u003c/em\u003e (15%) and \u003cem\u003eM. harttii\u003c/em\u003e (13%). Fleshy macroalgae, primarily Dictyotales, were recorded mainly in \u003cem\u003eM. harttii\u003c/em\u003e (2.9%), where their cover remained low. Sponges, meanwhile, colonized only a small proportion of \u003cem\u003eM. hispida\u003c/em\u003e skeletons (0.2%) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMean percent cover of organisms on dead skeletons for each species in December 2024. Six main benthic categories were identified: filamentous algae (Fil); crustose coralline algae (CCA); turf algae (Turf); calcareous turf (Taca); fleshy macroalgae (Fleshy); and sponges. For \u003cem\u003eSiderastrea\u003c/em\u003e sp. and \u003cem\u003ePorites astreoides\u003c/em\u003e, the organisms colonizing dead skeletons were not described (NA). The last column shows the mortality percentages observed for each species.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eFil(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eCCA(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eTurf(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eTaca(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eFleshy(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003esponges(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMortality(%)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. cavernosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e37.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. Hispida\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e32.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. Harttii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e54.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e13.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e18.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e93\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. humilis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e51.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e32.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e99\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eS. wellsi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e70\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. alcicornis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e44.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e30.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e23.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e99\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSiderastrea sp.\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eP. astreoides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e48\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMean\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e47.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e15.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e64\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThe intensification and increasing frequency of thermal anomalies have shaped our understanding of coral species\u0026rsquo; resistance thresholds worldwide (Hughes et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Marzonie et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Considering that both intrinsic species traits and extrinsic factors can influence the resilience potential of reef ecosystems (Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), understanding how these elements interact during large-scale bleaching events is essential. However, few studies focus on the response of individual organisms, which limits our understanding of the mechanisms and thermal tolerance thresholds of coral species (Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Our results indicate a clear difference in thermal stress responses among a wide range of coral species along the northeastern Brazilian coast, primarily driven by differences in life-history strategies (Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Swain et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hughes et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yadav et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFollowing the trend of increased mortality previously reported for the branching hydrocoral \u003cem\u003eM. alcicornis\u003c/em\u003e in earlier bleaching events worldwide (Lewis, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Dub\u0026eacute; et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Teixeira et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Duarte et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), we observed that this species nearly reached its survival threshold during the fourth mass bleaching event on Brazilian coral reefs (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Although \u003cem\u003eM. alcicornis\u003c/em\u003e is not a scleractinian coral, it plays a key role as a structural framework builder in shallow Brazilian reefs, occurring predominantly on reef crests and shallow submerged reef tops (Laborel, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Le\u0026atilde;o et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Therefore, future studies are needed to quantify the consequences of this significant reduction in Brazilian reef environments, particularly regarding the loss of structural complexity and the associated fish and invertebrate communities (Coni et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Graham \u0026amp; Nash, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSimilarly, \u003cem\u003eM. harttii\u003c/em\u003e has undergone marked population declines in recent years (Braz et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Pereira et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and is currently listed as \u0026ldquo;threatened\u0026rdquo; (IUCN). This phaceloid species can form large colonies and experience rapid skeletal degradation after mortality (Braz et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In contrast, the massive species \u003cem\u003eM. brasiliensis\u003c/em\u003e and \u003cem\u003eM. hispida\u003c/em\u003e have previously been described as more thermotolerant during bleaching events (Duarte et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, alongside the high mortality observed for \u003cem\u003eM. harttii\u003c/em\u003e, we also recorded a substantial increase in \u003cem\u003eM. hispida\u003c/em\u003e mortality since 2016, raising concerns about its resistance to intensifying thermal stress in the northwest coast. A similar post-bleaching mortality pattern was reported for \u003cem\u003eM. brasiliensis\u003c/em\u003e in the Abrolhos Archipelago after the 2019/2020 event (Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Both \u003cem\u003eMussismilia\u003c/em\u003e species rely on a broadcast-spawning reproductive strategy (Pires et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Neves and Pires, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), which increases their susceptibility to Allee effects following mass mortality, as reduced colony abundance limits cross-fertilization and ultimately decreases recruitment success (Courchamp et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Knowlton, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding the so-called \u0026ldquo;weedy\u0026rdquo; species (\u003cem\u003eA. humilis, S. wellsi, and P. asteroides\u003c/em\u003e), in spite of their variable susceptibility to bleaching. their brooding reproductive strategy (McGuire, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) may support faster post-stress recovery (Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Goodbody-Gringley \u0026amp; Putron, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and opportunistic pattern, increasing in reef cover following the decline of more sensitive corals (Green et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Carlos-J\u0026uacute;nior et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2026\u003c/span\u003e). In our study, although these species initially showed resistance, sharp population declines occurred under high thermal stress (DHW\u0026thinsp;\u0026gt;\u0026thinsp;15\u0026deg;C-weeks). \u003cem\u003eA. humilis\u003c/em\u003e, a species commonly associated with reef walls, approached its regional survival threshold (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), raising concerns about the time required for substantial population recovery. \u003cem\u003eS. wellsi\u003c/em\u003e, was the only species that remained bleached for an extended period (eight months), a pattern that may be linked to heterotrophic plasticity documented in several coral species under thermal stress (Anthony \u0026amp; Fabricius, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Grottoli et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Hughes \u0026amp; Grottoli, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Nevertheless, it still experienced high mortality toward the end of the event. In addition, this species has a restricted distribution range, commonly found at the base of reef formations (Francini-Filho et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), which may limit its recolonization following high-mortality events, as observed here.\u003c/p\u003e \u003cp\u003e \u003cem\u003eP. astreoides\u003c/em\u003e, although showing a slight shift toward more critical health categories since 2016 (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), exhibited a mortality rate approaching 50% during the fourth global bleaching event. Historically, this species has exhibited high survival and substantial population growth following thermal stress, often dominating reefs where more sensitive taxa experienced severe losses (Green et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). However, despite its \u0026ldquo;weedy\u0026rdquo; life-history traits (high fecundity, relatively fast growth, and tolerance to adverse conditions) recent population declines have been reported on Caribbean reefs as climate change accelerates (Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), a pattern mirrored in our observations. Additionally, the predominance of medium and small colonies further suggests a young population largely shaped by recent recruitment (Green et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), potentially reflecting population declines during previous events (Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). A similar colony size structure was observed for \u003cem\u003eSiderastrea sp.\u003c/em\u003e, also likely reflecting the loss of larger colonies and the predominance of a relatively young population in this species.\u003c/p\u003e \u003cp\u003eAs reported for other regions, some coral species have been classified as more resistant under global change (Van Woesik et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Darling et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Grottoli et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Along the Brazilian coast, our results similarly indicate that \u003cem\u003eMussismilia cavernosa\u003c/em\u003e and \u003cem\u003eSiderastrea\u003c/em\u003e sp. exhibit generalist strategies that favor persistence under increasing thermal stress, although through distinct mechanisms. \u003cem\u003eM. cavernosa\u003c/em\u003e commonly occurs on reef walls, where lower light availability and higher suspended particulate matter prevail (Francini-Filho et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), and its high heterotrophic plasticity (Lesser et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) may provide a metabolic advantage during bleaching events (Hughes \u0026amp; Grottoli, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). In contrast, \u003cem\u003eSiderastrea\u003c/em\u003e sp., typically found in shallow habitats with high thermal and irradiance variability, appears well adapted to extreme temperature fluctuations (Castillo et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Additionally, the microbial communities associated with these species may further contribute to their thermal tolerance (Abrego et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Oliver \u0026amp; Palumbi, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Davies et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSusceptibility to bleaching and mortality in colonies of different sizes within the same species remains a controversial topic and appears to vary among the taxa studied, potentially being positive, negative, or neutral with increasing colony size (Edmunds et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Winslow et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; \u0026Aacute;lvarez-Noriega et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For the two species tested here (\u003cem\u003eM. alcicornis\u003c/em\u003e and \u003cem\u003eM. cavernosa\u003c/em\u003e), we did not observe variability in health patterns (healthy, bleached, and dead) across colony sizes during the 2024 bleaching event. A similar lack of size-related variability was previously reported for \u003cem\u003eM. alcicornis\u003c/em\u003e, \u003cem\u003eM. cavernosa\u003c/em\u003e, and \u003cem\u003eM. braziliensis\u003c/em\u003e during the 2019/2020 bleaching event along the Brazilian coast (Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Cardoso et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn addition to the species-specific characteristics, extrinsic factors such as water circulation, depth, and light irradiance can attenuate thermal variability and alleviate stress on corals (Jokiel et al., 2004; Loya et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Yadav et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In the Tamandar\u0026eacute; reef complex, we did not detect depth-related variability in bleaching or mortality of \u003cem\u003eM. alcicornis\u003c/em\u003e and \u003cem\u003eM. cavernosa\u003c/em\u003e. In contrast to the patterns reported for 2019/2020 (Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), the lack of spatial structuring in \u003cem\u003eM. alcicornis\u003c/em\u003e mortality suggests that the intensity of thermal stress exceeded the buffering capacity typically provided by fine-scale environmental heterogeneity for this species. However, the spatial variability observed for \u003cem\u003eM. cavernosa\u003c/em\u003e during the 2024 event was highly similar to that previously described for \u003cem\u003eM. alcicornis\u003c/em\u003e during the 2019/2020 event (Vidal et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), with sites located in the southern portion of the reef complex, such as P1, P2, and P6, showing lower mortality rates, whereas P7 exhibited higher rates. This pattern indicates the presence of distinct areas of vulnerability at a fine spatial scale in response to thermal stress events. Further investigations integrating colony-level genetic characteristics and local circulation dynamics are needed to better elucidate the environmental factors underlying the spatial patterns observed.\u003c/p\u003e \u003cp\u003eFollowing mortality, rapid colonization by microorganisms (Leggat et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and later by macroalgae (Diaz-Pulido \u0026amp; McCook, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) accelerates the structural erosion of colonies (Bleuel et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Braz et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and hinders the settlement of new individuals (Doropoulos et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Consequently, the widespread replacement of living colonies by filamentous algae represents an obstacle to the recovery of the most affected species. In turn, the presence of crustose coralline algae covering a substantial proportion of \u003cem\u003eM. alcicornis\u003c/em\u003e skeletons may favor the long-term persistence of colony structure and promote the settlement of new recruits on these skeletons (Doropoulos et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, the recovery potential of coral populations may be compromised by local environmental pressures, particularly declining water quality and increasing sedimentation and contamination associated with agricultural runoff, uncontrolled population growth, and urban expansion in coastal zones, as observed in the study region (Silveira \u0026amp; Ferreira, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Vidal, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These conditions may further favor the establishment of opportunistic algae, intensify competition with corals, reduce recruitment success, and hinder effective recovery (Fabricius, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Ricardo et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; MacNeil et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Therefore, in addition to global policy measures aimed at mitigating climate change, local management actions are essential to ensure the long-term health and resilience of coral reef ecosystems (Richmond et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eWe observed a clear divergence in health trajectories among species during the fourth global mass bleaching event, driven primarily by their intrinsic characteristics. Our results reveal the collapse of key species responsible for maintaining the structural complexity and diversity of Brazilian reefs, such as \u003cem\u003eM. alcicornis, M. harttii, and A. humilis\u003c/em\u003e. We also found moderate to high mortality in species previously considered thermotolerant, including \u003cem\u003eM. hispida\u003c/em\u003e and \u003cem\u003eP. astreoides\u003c/em\u003e, indicating that climate change is reshaping the thermal tolerance patterns once described for these taxa. In contrast, \u003cem\u003eM. cavernosa\u003c/em\u003e and \u003cem\u003eSiderastrea sp.\u003c/em\u003e stood out for their high tolerance to thermal stress, suggesting that they may become increasingly dominant reef builders in the future, although long term mortality may be investigated. We detected no variation in mortality among colonies of different sizes and subtle fine-scale spatial patterns, underscoring the severity and homogenizing effect of the event. Our findings also reveal variability in the composition of algal communities colonizing the dead skeletons of different species, with a predominance of filamentous algae, which may pose a substantial challenge to the recovery of species that experienced sharp declines. Collectively, our results highlight the vulnerability of South Atlantic coastal reefs to accelerating climate change, the future risk of local disappearance of species that approached their thermal tolerance limits during the fourth global mass bleaching event, and the accumulation of local and global pressures that may compromise the long-term resilience of coral populations along the northeastern coast of Brazil.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAuthors declare they have no competing interests that are relevant to the content of this article.\u003c/p\u003e\u003cp\u003e \u003ch2\u003eConflicts of Interest\u003c/h2\u003e \u003cp\u003eThe authors declare no conflicts of interest.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eThales J. Vidal: conceptualization, data collection and curation, formal analysis, methodology, visualization, writing \u0026ndash; original draft. \u0026Aacute;gatha N. Ninow: conceptualization, methodology, data collection and curation, writing \u0026ndash; review and editing. Ana Lidia Gaspar: conceptualization, funding acquisition, methodology, project administration, resources, supervision, writing \u0026ndash; review and editing. Beatriz Fernandes: data collection; methodology, visualization, writing \u0026ndash; review and editing. Thiago Buchianeri: data collection; methodology, visualization, writing \u0026ndash; review and editing. Mauro Maida: conceptualization, funding acquisition, methodology, project administration, resources, supervision, writing \u0026ndash; review and editing. Leonardo Messias: conceptualization, funding acquisition, resources, writing \u0026ndash; review and editing. Camila da Silveira Brasil: data collection, methodology, writing \u0026ndash; review and editing. Beatrice P. Ferreira: conceptualization, data curation, funding acquisition, methodology, project administration, resources, supervision, writing \u0026ndash; review and editing.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eThis is a Tamandar\u0026eacute; Long-Term Ecological Program (PELD-TAMS, ILTER site 18) contribution. We thank the National Center for Research and Conservation of Marine Biodiversity in the Northeast (CEPENE-Tamandar\u0026eacute; ICMBio) for invaluable field and logistic support. The Coastal Reefs Institute (IRCOS), the Tamandare Cave Project (CECAV), and WWF-Brasil (CPT 003776-2024) for their financial and logistical support during field trips and collections. We thank the Brazilian scientific council CNPQ for a grant to BPF (442139/2020-9). We also thank the Post Graduate Program in Oceanography at the Federal University of Pernambuco (PPGO/UFPE). All sampling activities were conducted under the permits issued by SISBIO for the Long-Term Biodiversity Monitoring Program (No. 45992) and Reef Check Brasil (No. 18355).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets generated and/or analysed during the current study are available from the corresponding author upon reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbrego, D., Ulstrup, K. E., Willis, B. L., \u0026amp; van Oppen, M. J. (2008). Species\u0026ndash;specific interactions between algal endosymbionts and coral hosts define their bleaching response to heat and light stress. 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J., Devlin, M., \u0026amp; Anthony, K. R. (2018). Vulnerability of the Great Barrier Reef to climate change and local pressures. Global change biology, 24(5), 1978\u0026ndash;1991. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/gcb.14043\u003c/span\u003e\u003cspan address=\"10.1111/gcb.14043\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYadav, S., Roach, T. N., McWilliam, M. J., Caruso, C., de Souza, M. R., Foley, C., \u0026hellip; Madin, J. S. (2023). Fine-scale variability in coral bleaching and mortality during a marine heatwave. Frontiers in Marine Science, 10, 1108365. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fmars.2023.1108365\u003c/span\u003e\u003cspan address=\"10.3389/fmars.2023.1108365\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"coral-reefs","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"core","sideBox":"Learn more about [Coral Reefs](http://link.springer.com/journal/338)","snPcode":"338","submissionUrl":"https://submission.nature.com/new-submission/338/3","title":"Coral Reefs","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Bleaching, Climate change, Coastal coral reefs, Thresholds, Thermal Tolerance","lastPublishedDoi":"10.21203/rs.3.rs-9408247/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9408247/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe increase in the frequency and intensity of thermal anomalies, combined with local pressures, has reshaped our understanding of the thermal tolerance potential of coral species on a global scale. In this study, through detailed monitoring, we tracked the health trajectories of 290 colonies from eight reef-building species along Brazil’s northeastern coast before, during, and after the fourth global mass bleaching event. Our results reveal striking differences in species vulnerability, fine-scale spatial variability in mortality patterns, no significant variation in the response of colonies of different sizes, and marked heterogeneity in algal colonization on dead skeletons, with filamentous algae predominating nine months after the onset of bleaching. In addition, by considering heatwaves events with different levels of Degree Heating Weeks, we assessed the responses of five species based on more than 3,000 colony records collected throughout 2016 and 2024 along fixed transects and tracked colonies. The findings indicate that intensifying thermal stress has pushed some species close to their survival thresholds, reinforcing that reefs are undergoing biodiversity loss and will likely become dominated by thermotolerant species in the near future. Overall, our results underscore the vulnerability of South Atlantic coastal reefs to climate change and highlight species-specific differences in thermal tolerance, with important implications for the future composition and resilience of these ecosystems.\u003c/p\u003e","manuscriptTitle":"The Limits of Coral Thermal Tolerance in Tropical South Atlantic Coastal Reefs after the Fourth Global Mass Bleaching Event","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-05-06 12:47:50","doi":"10.21203/rs.3.rs-9408247/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"325147960768825516168941911256505123671","date":"2026-05-17T23:02:35+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"165432120222906333385452461587319738331","date":"2026-05-14T22:22:17+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"167592238214654990145608803342769281397","date":"2026-05-14T21:59:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"299183102318839835526005717736315291782","date":"2026-05-13T11:38:13+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"317736633918560976004573635663119002516","date":"2026-04-30T21:41:23+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-28T07:31:32+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-17T06:25:52+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-15T16:11:32+00:00","index":"","fulltext":""},{"type":"submitted","content":"Coral Reefs","date":"2026-04-13T20:24:16+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"coral-reefs","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"core","sideBox":"Learn more about [Coral Reefs](http://link.springer.com/journal/338)","snPcode":"338","submissionUrl":"https://submission.nature.com/new-submission/338/3","title":"Coral Reefs","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"c642f246-607d-4db0-bd73-7992ae1e5498","owner":[],"postedDate":"May 6th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewerAgreed","content":"325147960768825516168941911256505123671","date":"2026-05-17T23:02:35+00:00","index":24,"fulltext":""},{"type":"reviewerAgreed","content":"165432120222906333385452461587319738331","date":"2026-05-14T22:22:17+00:00","index":23,"fulltext":""},{"type":"reviewerAgreed","content":"167592238214654990145608803342769281397","date":"2026-05-14T21:59:32+00:00","index":22,"fulltext":""},{"type":"reviewerAgreed","content":"299183102318839835526005717736315291782","date":"2026-05-13T11:38:13+00:00","index":20,"fulltext":""},{"type":"reviewerAgreed","content":"317736633918560976004573635663119002516","date":"2026-04-30T21:41:23+00:00","index":10,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-06T12:47:50+00:00","versionOfRecord":[],"versionCreatedAt":"2026-05-06 12:47:50","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9408247","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9408247","identity":"rs-9408247","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}