Two decades of skeletal density decline in Pocillopora spp. corals in the Mexican Pacific Ocean: Insight into a tropical eastern Pacific acidification scenario?

Andrés López-Pérez; Omar Jiménez-Gutiérrez; Héctor Reyes-Bonilla; Rafael A. Cabral-Tena; Gerardo Leyte-Morales; Francisco Medellín-Maldonado; Orión Norzagaray-López; Cecilia Chapa-Balcorta

doi:10.1371/journal.pone.0342741

Abstract

Corals demonstrate vulnerability to environmental changes, exhibiting the capacity to substantially modify coral calcification. In this study, we estimated declines in the density of Pocillopora coral species in the Mexican Pacific. The samples utilized in this study encompass both recently collected corals and those stored in Mexican repositories collected in the northeastern and southern Mexican Pacific regions. Density estimates indicate a 28.6% decline in coral density over the past 23 years (−0.0227 g CaCO₃ cm^-3 y^-1) in the southern Mexican Pacific, while at the entrance to the Gulf of California, density has decreased by 15.4% over the past 20 years (−0.017 g CaCO₃ cm^-3 y^-1). A comprehensive evaluation of environmental data reveals that the observed decline in Pocillopora skeletal density in Mexican Pacific reefs is concomitant with decreases in Ω_ar and pH, and an increase in ocean temperature on a substantial regional scale. When considered in conjunction with the previously documented reductions in coral growth of Pocillopora spp. skeletons in the eastern Tropical Pacific, our findings indicate a potential decline in CaCO₃ production within the region's reef systems. The results of this study underscore the significance of generating long-term series of coral growth parameters for relevant reef-building species and the carbonate system in key and representative coastal areas, particularly those that are already challenging for coral survival and reef maintenance.

Citation: López-Pérez A, Jiménez-Gutiérrez O, Reyes-Bonilla H, Cabral-Tena RA, Leyte-Morales G, Medellín-Maldonado F, et al. (2026) Two decades of skeletal density decline in Pocillopora spp. corals in the Mexican Pacific Ocean: Insight into a tropical eastern Pacific acidification scenario? PLoS One 21(2): e0342741. https://doi.org/10.1371/journal.pone.0342741

Editor: Anderson B. Mayfield, Living Oceans Foundation, TAIWAN

Received: December 16, 2024; Accepted: January 28, 2026; Published: February 26, 2026

Copyright: © 2026 López-Pérez et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Data Availability: All relevant data are within the paper and its Supporting Information files.

Funding: This study was supported by the Consejo Nacional de Humanidades, Ciencias y Tecnologías (CONAHCyT, grant number: 866397 to RACT; grant number: 278637 to CCB) and the Universidad Autónoma Metropolitana (UAM, grant number: 14705007 to ALP).

Competing interests: The authors have declared that no competing interests exist.

Introduction

Coral reefs are intricate three-dimensional structures that serve as a vital foundation for maintaining elevated levels of biodiversity and providing a multitude of goods and ecosystem services [1]. The three-dimensional structure of reefs is contingent upon the accumulation of CaCO₃ by calcifying organisms, predominantly corals [2]. However, corals are vulnerable to environmental and anthropogenic changes that have the potential to significantly alter coral calcification and, in extreme cases, threaten the maintenance of reef systems at different spatiotemporal scales [3–5]. These threats pose physiological challenges to corals, requiring modification of metabolism to adapt and acclimate. A number of parameters, including linear extension rate (cm yr^-1), skeletal density (g cm^-3), and calcification rate (g cm^-2 yr^-1), have been identified as useful in evaluating the growth rates of coral over time [3]. Consequently, growth parameters are instrumental in elucidating the impact of environmental changes on the present decline of coral populations.

Several studies have recently reported changes in coral calcification rates in response to environmental variability in various coastal regions [4,6,7]. Coral reefs are among the earliest warm-water ecosystems to be impacted by climate change. The consequences of this phenomenon are multifaceted, chiefly concerning the response of corals and other reef organisms to rising temperatures and ocean acidification [8,9]. For instance, in the eastern Tropical Pacific (ETP), coral reefs are predominantly formed by near monogeneric communities of branching Pocillopora type 1 (> 90% live coral cover). A coral-growth model for the equatorial ETP predicts a gradual reduction in the linear extension of Pocillopora, estimated at 0.9% per year, due to increased acidification [10,11]. In contrast, a decline in skeletal density has been documented in Orbicella faveolata, Siderastrea siderea and Pseudodiploria strigosa, while linear extension rates have remained constant. This phenomenon can be attributed to changes in the calcium carbonate saturation state, specifically the aragonite (Ω_ar) component, within the Florida Reef Tract and the Colombian Caribbean [12,13]. Manzello et al. [14] reported a decline in skeleton density of Porites lobata in the southern Galapagos, while extension showed no significant trend. The authors posit that elevated nutrient concentrations in upwelled waters and/or increased heterotrophy from high water-column productivity may stimulate extension and calcification, but impair skeletal density. Concurrently, Mollica et al. [5] demonstrated through modeling that skeletal density, but not extension rate, exhibited sensitivity to ocean acidification; the model, which has been validated with data from disparate geographical regions, predicts that ocean acidification will result in a 12 ± 6% reduction in skeletal density of Porites by the end of the 21^st century [5]. In summary, although the response of corals to environmental stressors, particularly climate change, exhibits variation depending on the species and geographical region, ocean acidification and the increase in sea-surface temperature have been identified as significant stressors of coral calcification.

The majority of studies that have assessed the major drivers of variation and the temporal variability of coral calcification have focused on species that form growth bands, from which skeletal extent and density can be determined [15]. Conversely, there is a paucity of data for species with branching morphologies, including Acropora and Pocillopora species. This complicates the monitoring of changes in growth parameters for these significant reef-building species in the Caribbean and eastern Pacific, which do not form growth bands [16,17]. Pocillopora corals are the primary reef-building species in the ETP since the late Pleistocene epoch, and the CaCO₃ production and reef-structural complexity of this region are closely linked to the abundance and calcification rate of these corals [17,18]. Consequently, any reduction in Pocillopora corals, whether attributable to natural or human-induced perturbations or calcification, could have an immediate or long-term detrimental impact on the functionality, stability, and biodiversity of the coral ecosystems in question [19]. The scarcity of data concerning the growth parameters of coral species relevant to reef system functionality is particularly disconcerting in the ETP, a region that is known to possess the most severe natural acidification conditions for reef development in the world [20,21]. The region’s environmental conditions are conducive to minimal reef accretion due to the interaction of upwelled subsurface water with low pH (7.8 as the lower threshold) and a near-unity subsaturation state of aragonite in seawater. It is anticipated that these conditions will become even more marginal for accretion in future ocean acidification scenarios [11,21–24].

This study presents empirical data on the decline in the density of Pocillopora type-1 skeletons over the last two decades in the Mexican Pacific. The findings are then contextualized within the progressive acidification scenario of the eastern Pacific and the potential risk to the persistence and functioning of reef systems in the region.

Materials and methods

Study area

Sampling was conducted at multiple locations in the eastern tropical Pacific, with a particular focus on the northwestern region (entrance of the Gulf of California) and the southern region (coast of Oaxaca) of the Mexican Pacific (Fig 1). A distinguishing characteristic of both regions is their elevated degree of oceanographic dynamism, which is influenced by a variety of seasonal processes. At the entrance to the Gulf of California, anticyclonic eddies with cold-water filaments of approximately 22°C are generated by the collision of the California Current with the continent. This collision subsequently leads to the former's expansion, with a portion of it entering the Gulf [25,26]. The Gulf of California Water, a water mass of local origin, exhibits the highest salinity values in the area (≥ 34.9) and demonstrates a notable variability in temperature (> 12°C annual range). In contrast, less saline (< 34.9) and comparatively warmer Tropical Surface Water, which originates from the Pacific Ocean, enters the system [25].

Download:

Fig 1. Study area and sampling sites in the Mexican Pacific (a).

(b) Green = entrance of the Gulf of California: Isla San José, San Gabriel, Swami, Bonanza, Pichilingue, Calerita, Punta Arenas, Punta Perico, Bahía Chileno, Cabo Pulmo. (c) Blue = coast of Oaxaca: Mazunte, Panteón, Estacahuite, La Tijera, Salchi, Riscalillo, La Prima, La India, Maguey, La Entrega, Dársena, Isla Montosa, Guerrilla.

https://doi.org/10.1371/journal.pone.0342741.g001

The limited reports from the Gulf of California region pertaining to carbon chemistry, encompassing dissolved inorganic carbon (DIC) concentration, Ω_ar, and pH, suggests a dependence of conditions on the predominant water masses at the seasonal scale. From spring to fall, tropical surface water is predominant, characterized by DIC values of 2008 ± 48 µmol kg^-1, Ω_ar = 3.2 ± 0.3 units, and pH = 8.01 ± 0.03. In contrast, during the winter months, the Gulf of California water predominates, characterized by higher DIC values (2079 ± 27 µmol kg^-1), the lowest Ω_ar (2.8 ± 0.2 units), and a pH of 7.93 ± 0.02. The mean pH, when both seasons are considered is 7.99 ± 0.05, with a range of 0.14 units (7.92–8.06) [26–29].

The coral communities situated along the coast of Oaxaca are shaped by the variable seasonal conditions that prevail in this region. Intense coastal upwelling, promoted by seasonal strong winds known as “Tehuanos”, is observed in the western region of the Gulf of Tehuantepec (the same mechanism is observed in Costa Rica [20,30]). This phenomenon occurs during the spring months of March and April. The mechanism in question gives rise to the vertical transportation of subsurface water, which is characterized by elevated concentrations of DIC (~ 2200 µmol kg^-1) that extend along the coastline. The upwelled subsurface water is distinguished by its relatively low pH (~7.7) and Ω_ar values (~1.4 units). It is subsequently mixed with surface water by mesoscale structures, such as dipole eddies interacting with coastal waters [31]. Throughout the remaining months of the year, the water column situated above 100 m in depth is composed of tropical surface water, with pH values ranging from 8.2 to 8.3 and Ω_ar values from 2 to 2.4 [32].

Coral sampling

The experimental design was devised to ascertain trends in skeletal density of Pocillopora colonies collected during two distinct periods at both localities: (1) 1996 and 2016 at the entrance of the Gulf of California and (2) 1994 and 2017 at the coast of Oaxaca (Fig 1). The experimental design was selected because Pocillopora do not form annual bands as massive corals do. This characteristic complicates the sclerochological analysis of the genus. In 2016, 33 fragments of Pocillopora spp. colonies were collected at five locations in the Gulf of California, while 34 fragments were obtained from colonies collected in 1996 at seven localities in the same area (Fig 1a–1b). Presently, these fragments are housed in the coral collection of the Natural History Museum of the Autonomous University of Baja California Sur. Conversely, in 2017, 38 coral fragments were collected at six localities on the coast of Oaxaca, while 38 fragments were obtained from colonies collected during 1994 at ten localities in the same area, which are deposited in the reference coral collection of the Universidad del Mar (Fig 1a, 1c). Information regarding the morphotypes sampled per locality, area, and time can be consulted in supplementary material (S1 Table).

In consideration of the sampled times, the studied periods were deemed to be within the ENSO-neutral variability (https://psl.noaa.gov/data/timeseries). According to the National Oceanic and Atmospheric Administration (NOAA) (https://psl.noaa.gov/data/timeseries), the estimated levels of marine heat waves were low (<1 DHW) for each of the years considered in the study (https://coralreefwatch.noaa.gov/product/5km/index_5km_dhw.php). Therefore, it is unlikely that El Niño Southern Oscillation (ENSO)-related heat stress significantly impacted calcification during the study's time frame.

Samples were collected from colonies growing at depths ranging from 1 to 9 meters. In all cases, the fragments were obtained from 2 centimeters below the tip of the branch zone. This criterion was adopted for the purpose of obtaining density estimations from coral sections, thereby circumventing the initial rapid growth reported from colonies’ tips [33]. To remove the coral tissue remnants, the fragments were immersed in a 5% sodium hypochlorite solution for a period of 24 hours [34].

Density estimations

Due to the paucity of long-term series with which to ascertain changes in skeletal density of branching corals, we sought to obtain density data from skeletons deposited in biological reference collections. The determination of density by means of the simple ratio of weight to volume (weight/volume) of the skeleton plus the volume of the “air” represents a low-tech, robust method that allows for the accurate and reproducible determination of the density of highly porous objects, such as corals, by water freezing. Cubes measuring approximately 1 cubic centimeter were obtained from each fragment using a high-speed diamond saw (SYJ-40-LD). The density of the cubes was estimated by the freezing method, following the procedure described by Carricart-Ganivet et al. [34]. Equation (1) considers the weight of the dry fragment (W_d), the density of aragonite (d_ar, 2.93 g cm^-3), the weight of the frozen fragment (W_f), and the density of ice (d_ice, 0.915 g cm^-3). Skeletal density of the fragment was determined using the following formula (d_b):

1

The data presented herein constitutes a conservative estimate of the density of the branches of the colonies, excluding the thickening and widening of the branch tips.

Environmental setting

The objective of this study is to ascertain whether changes through time in the skeletal density have occurred among select coral species within the designated study areas. Additionally, the investigation seeks to ascertain whether these decrements may be associated with changes in environmental conditions, particularly with variables associated with ocean acidification (OA) over the past two decades. Both study areas exhibit a dynamic oceanography, characterized by numerous processes occurring at varying temporal and spatial scales. These include seasonal vertical and horizontal advection of water masses, precipitation, and interannual phenomena such as El Niño Southern Oscillation (ENSO) events, to name a few.

Time series records of OA indicators (i.e., pH, DIC, saturation state regarding aragonite [Ω_ar]) are scarce for both study areas, a common situation for many oceanic and coastal regions of the Mexican Pacific. The scenario is rendered more complex by the fact that the available data correspond to relatively short and discontinuous time periods. Moreover, the data from multiple sources exhibited considerable heterogeneity, attributable to various factors. These factors include the methodologies employed for calculation, the established best practices for data analysis, the spatial and temporal resolutions of monitoring, and additional elements contributing to the natural variability observed in both regions. The distribution of the data in an irregular pattern hinders the identification of long-term trends. A strategy employed to study long-term biogeochemical processes (i.e., OA) in areas with limited data is to utilize modeled data to complement observed data.

In this study, we have calculated the pHT (total scale pH) and the Ω_ar from atmospheric pCO₂ (pCO_2atm) and modeled total alkalinity (TA). Notwithstanding the potential oversimplification of the carbonate-system dynamics that may result from this approach, its potential application for the purposes of this study was nevertheless evaluated. To achieve this objective, monthly-averaged environmental variables (i.e., temperature, °C; silicate and phosphate concentration, μM; salinity) at annual scale for each studied site were obtained for 1° x 1° quadrats from the World Ocean Atlas 2018 (WOA18, https://www.ncei.noaa.gov/products/world-ocean-atlas). The TA (µmol kg^-1) was calculated using the equations developed by Lee et al. [35] for subtropical (Gulf of California entrance) and tropical (coast of Oaxaca) areas (± 20 µmol kg^-1). The calculation of Ω_ar and pHT was performed using the software CO2sys [36], with salinity, temperature, depth, phosphate, silicate, TA, and pCO_2atm utilized as the dependent variables. The pCO_2atm data were obtained from the CO₂ program of the Scripps Institution of Oceanography for the designated study periods (https://scrippsco2.ucsd.edu/data/atmospheric_co2/ljo.html; considering a nominal error of 20 µatm). The pHT and Ω_ar estimations presented an uncertainty of 0.01–0.03 and 0.08–0.14 units, respectively (20 µmol kg^-1 for DIC and 0.1 °C data).

To verify if calculated pHT fall within the range of measured data variability, carbon chemistry data were obtained from different sources (S2 Fig A–D). To contrast pHT and Ω_ar values between the different databases (S2 Fig A–D), the same carbonate dissociation constants were utilized [37]. In instances where TA data were not available (e.g., SOCAT database [38]), the necessary estimates were derived using the methodology outlined by Lee et al. [35]. This exercise facilitated the validation of the estimated data, which were determined to align with the range of observed natural variability. This finding suggests that the estimated data may be representative of the prevailing conditions in each region.

Furthermore, to ascertain the long-term shifts in pH within the two study areas, pHT and Ω_ar were derived from the outputs of the Community Earth System Model v2, which incorporates historical DIC, TA, temperature, and salinity data as reported by Wieners et al. [39]. The selection of this model is based on its demonstrated capacity to replicate the observed seasonal patterns and magnitudes of temperature and salinity. Despite its proclivity to underestimate average pH and smooth natural variability, this model enabled the estimation of trends in Ω_ar and pHT.

Data analysis

To assess changes in coral skeletal density between areas, times, localities, and species, the present study employed a four-way nested (area (time (locality (species))) unbalanced type III permutation-based analysis of variance (PERMANOVA). The PERMANOVA was conducted on a Euclidean distance matrix with 10,000 permutations, in accordance with the criteria set forth by Anderson et al. [40].

In addition, an analysis of covariance (ANCOVA) was employed to ascertain whether the trajectories of skeletal density (i.e., the rates of change over time) vary between regions (i.e., the Gulf of California entrance versus the coast of Oaxaca). This approach enables testing the hypothesis that the slopes of each linear regression in each area are different. In this analysis, area was utilized as a categorical variable, year as a covariate, and skeletal density as the response variable. In essence, an ANCOVA was employed to investigate a significant interaction between the categorical variable (area) and the covariate (time). Prior to conducting the aforementioned analyses, the assumptions of normality and homoscedasticity were evaluated via the Kolmogorov-Smirnov and Bartlett tests, respectively.

Due to the restricted access to Pocillopora spp. museum samples for density determinations and the inclusion of only two time periods, the analysis of environmental variables was conducted using principal component analysis. The ordination encompassed vectors for each environmental variable, with the length and direction of the vector serving as proxies for the relative importance of each variable in the ordination.

The decrements in skeletal density undergone by Pocillopora spp. within the ETP over time were examined through the application of simple linear regression analysis. The coral density data utilized in the analysis corresponded to the values determined in this contribution and to the determinations published by [11,41–43].

All statistical analyses were performed at α = 0.05. Analyses were performed using Past v4.09 [44] and PRIMER & PERMANOVA+ 6.1 [40].

Results

The findings indicated that the sole dimension of sampling across which there were statistically significant disparities in density was between time periods (Pseudo-F_(2,81)= 15.83, p = 0.0001). The analysis revealed no statistically significant differences between areas (Pseudo-F_(1,81)= 2.27, p = 0.27), between sites nested within time and area (Pseudo-F_(24,81)= 1.50, p = 0.14), or between species nested within site, time and area, were observed (Pseudo-F_(28,81)= 1.06, p = 0.4) (S4–S6 Fig).

In general, corals of the genus Pocillopora exhibited higher skeletal density, with an average of approximately 30% more calcium carbonate per unit volume, at coral reefs located at the entrance of the Gulf of California (2.04 ± 0.34 g cm^-3 CaCO₃) compared to coral reefs of Oaxaca (1.55 ± 0.39 g cm^-3 CaCO₃) (S5 Fig and Table 1). In 1996, the Gulf of California entrance site known as Punta Perico exhibited the highest average density value (2.23 ± 0.09 g CaCO₃ cm^-3), while the lowest average value was recorded at Isla San José (1.98 ± 0.04 g CaCO₃ cm^-3). In contrast, the mean skeletal density values for all sites along the coast of Oaxaca in 1994 were lower than those documented at the Gulf of California entrance in 1996, with the exception of Riscalillo (2.00 ± 0.14 g CaCO₃ cm^-3). The lowest recorded value for the coast of Oaxaca was observed at La Tijera (1.62 ± 0.76 g CaCO₃ cm^-3).

Download:

Table 1. Average environmental variability and skeletal density (± SD) of Pocillopora species across regions and time in the Mexican Pacific. For details on the methodology employed to obtain the environmental values, please refer to the text. SST = °C, Ω_ar = aragonite, skeletal density = g CaCO₃ cm^-3. *Significant differences (p < 0.05).

https://doi.org/10.1371/journal.pone.0342741.t001

A decline in skeletal density over time was observed in both areas. The density exhibited a higher value in 1994 (1.813 ± 0.08 g cm^-3 CaCO₃) compared to 2017 (1.295 ± 0.1 g cm^-3 CaCO₃) in the coastal region of Oaxaca. A similar trend was noted in the skeletal density, which exhibited a higher value in 1996 (2.21 ± 0.21 g cm^-3 CaCO₃) compared to 2016 (1.87 ± 0.37 g cm^-3) in the Gulf of California entrance (S5 Fig). The aforementioned evidence indicates that there has been a notable reduction in the density of Pocillopora spp. skeletons. The decline in density was 28.6% over the past 23 years in Oaxaca, with a rate of decrease of approximately 0.022 g CaCO₃ cm^-3 in calcification per year. In contrast, a decrease of 15.4% in density was observed over the last 20 years in the Gulf of California entrance, with a rate of decline of about 0.017 g CaCO₃ cm^-3 in calcification per year. An analysis of covariance (ANCOVA) was employed to assess the impact of area and the continuous covariate (year) on skeletal density, incorporating the interaction between year and area to compare slopes. The findings indicated that the effect of year was significant (F_(1,133)=70.62, p < 0.001), indicating that skeletal density undergoes a substantial decline over time. In addition, the effect of area proved to be significant (F_(1,133)=104.27, p < 0.001), thereby indicating disparities in mean skeletal density across distinct areas. Conversely, there was no interactive effect of year and area (F_(1,133)=2.58, p = 0.111), indicating that the rate of change in density over time is statistically comparable across regions. In the fitted model, the slope for the Gulf of California was determined to be −0.01512 g CaCO₃ cm^-3 y^-1 (p = 0.0001), while for the Oaxaca area was found to be −0.0227 g CaCO₃ cm^-3 y^-1 (p = 0.0001). In summary, density has exhibited a negative trend over the years, a pattern that is consistent across different regions.

Concurrently, the pCO2atm and TA models, as well as the CESM2 model, indicate a modest decline in pHT and Ω_ar for both regions (Table 1). This decline is evident in the data from 1995 to 2014, with a decrease of −0.001 units per year for pHT and −0.005 units per year for Ω_ar (S3 Fig). However, the magnitude of this decline appears to be slightly more pronounced in the Oaxaca coastline compared to the Gulf of California entrance.

The principal component analysis explained 99.3% of the environmental variation between areas and times (Fig 2, Table 1). Along PC1 (77.8% variance explained), the upper left part of the ordination biplot displays the environmental centroids for areas and times sampled in 1994 and 1996, while the lower right part of the ordination shows the environmental centroids of areas and times sampled in 2016 and 2017. As indicated by the aforementioned evidence, the environmental conditions have undergone modifications, particularly evident among samples obtained at different times within a given area. The analysis indicated that the most influential variable in the ordination was Ω_ar (r = 0.68 with PC1), and in accordance with the vectors’ angle, this indicates an inverse response, exhibiting lower values towards 2016 and 2017. Conversely, temperature exhibited a positive correlation with PC1 (r = −0.64 with PC1), indicating that temperature has increased in the areas towards 2016/2017. Finally, pH demonstrates a robust positive correlation (r = 0.92) with PC2. Consequently, the areas have experienced a decrease in Ω_ar and pH and an increase in ocean temperature (Fig 2, Table 1). However, according to the spatial Euclidean distance, the environmental change was more pronounced at the Gulf of California entrance (2.51) than along the coast of Oaxaca (1.80).

Download:

Fig 2. Principal component analysis (PCA) of areas and times based on environmental variables.

For further details, please refer to the text. GC denotes the Gulf of California entrance, while Oax represents the coast of Oaxaca. The green color indicates the environmental change between 1996 and 2016 within the Gulf of California entrance, while the blue color represents the environmental change between 1994 and 2017 within the coast of Oaxaca. The dark blue color denotes the environmental vectors (r = 0.8), which include surface pH, surface Ω_ar, and SST. Spatial Euclidean distance: Gulf of California entrance = 2.51, coast of Oaxaca = 1.8.

https://doi.org/10.1371/journal.pone.0342741.g002

Finally, the analysis of changes in skeletal density of Pocillopora spp. indicates a notable decline between 1994 and 2017 in the ETP (R2 = 0.23, n = 21, p < 0.03; y = −0.0217x + 45.411) at a rate of −0.0217 g CaCO₃ cm^-3 per year (Fig 3).

Download:

Fig 3. Linear trend in skeletal density in Pocillopora species in the eastern tropical Pacific Ocean over time.

The data points represent the density values for each time period and area (combining both localities’ data), while the error bars indicate the standard deviation of density. The dotted line represents the linear regression model.

https://doi.org/10.1371/journal.pone.0342741.g003

Discussion

The estimated density values for the various Pocillopora species in this study, regardless of the area and time at which they were determined, fall within the range previously reported by other studies in the ETP [11,41–43,45]. Despite the presence of minor discrepancies in skeletal density across species within sites, these differences did not reach statistical significance when considered at the site, area, and between areas. The only factor for which statistically significant differences in skeletal density were observed for corals of the genus Pocillopora was time. The permuted analysis of variance demonstrated a statistically significant reduction in skeletal density in both the Gulf of California entrance (15.4% overall, −0.01512 g CaCO₃ cm^-3 per year) and the Oaxaca area (28.6% overall, −0.0227 g CaCO₃ cm^-3 per year) over the past two decades. Despite the apparent disparities in the rate of density reduction among different areas (Oaxaca > Gulf of California entrance), analysis of covariance indicates that these differences are not statistically significant. This finding indicates that the observed decrease in skeletal density within Pocillopora is not a phenomenon confined to a specific spatial location, but rather a widespread occurrence across a considerable spatial extent. Consequently, we hypothesize that this phenomenon is evident throughout the entire Mexican tropical Pacific region, extending from the Gulf of California entrance to the coast of Oaxaca. However, the extant data on Pocillopora density suggest that this pattern is not confined to the Mexican Pacific region but rather occurs on a ETP regional scale [11,41–43,45]. According to the analysis, the skeletal density of Pocillopora in the eastern Pacific may be declining at a rate of 0.0217 g CaCO₃ cm^-3 per year, which is similar to the carbonate loss suggested for the coast of Oaxaca.

A principal component analysis reveals an increase in sea-surface temperature and a concomitant decline in Ω_ar and pH levels in both the Gulf of California entrance and the southern Mexican Pacific over the past two decades. The estimated values of Ω_ar and pH estimated are consistent with a basic model that posits that temperature and atmospheric pCO₂ are the predominant influencing factors on the carbonate system in the region. This phenomenon may be attributable to the decadal variability, where other biogeochemical processes (in addition to the CO₂ gas equilibrium) are also active and contributing to this effect [46]. The collective findings suggest a possible correlation between the observed decline in Pocillopora skeletal density in the Mexican Pacific and elevated temperatures, as well as decreases in Ω_ar and pH in the ocean waters where corals develop. However, a comprehensive record of Pocillopora growth parameters (skeletal density, extension rate, calcification rate) remains to be established. In addition, there is a paucity of long-term records of carbon chemistry to validate the modeled data. These facts hinder our capacity to elucidate the mechanism by which skeletons become less dense in response to increased temperature and acidification in both areas. We hypothesize that the cooler waters of the eastern Pacific [47] and the Pocillopora species within them demonstrate heightened rates of extension in response to elevated sea-surface temperatures. This phenomenon would result in a reduction in skeletal density, achieved by extending their skeletons to a greater extent with the same or a reduced amount of calcium carbonate. However, ex situ evidence suggests that when pocilloporids are acclimated to an elevated temperatures (i.e., 1 °C above their bleaching threshold) for extended periods (six years), skeletons exhibit an increased density but reduced skeletal extension and calcification rates. This phenomenon can be attributed to the prioritization of energy storage over skeletal growth at elevated temperatures [48]. Therefore, regardless of the other variables, environmental conditions (and stress) are the primary drivers of coral calcification. The capacity of coral to undergo changes in metabolism is contingent upon a tenuous energetic equilibrium. This balance is surrogated by energy inputs, which in the ETP are predominantly autotrophic and, to a lesser extent, heterotrophic [49]. Consequently, the constrained energy is apportioned to multiple, ordered processes (e.g., damage repair, somatic growth, calcification, reproduction, energy storage, among others). Imbalances (e.g., OA modifying internal ionic balance) have the potential to influence the organism's processes, including calcification and/or growth. This finding indicates that Pocillopora species may prioritize skeletal extension over the thickening of skeletal structures (i.e., increasing the skeletal density) as demonstrated by Carricart-Ganivet and Merino [50] and Carricart-Ganivet [51] for specific Atlantic and Caribbean coral species.

Mollica et al. [5] developed a numerical model of Porites skeletal growth. This model establishes a correlation between the skeletal density of corals and the surrounding water in which they develop, attributing this relationship to its impact on the chemistry of coral calcifying fluid. This fluid is located between the coral skeleton and its calicoblastic cell membrane [52]. In contrast to temperature, which has the potential to influence skeletal density or extension depending on the species and the area where corals develop [52–54], Mollica et al. [5] demonstrated that among the variables associated with coral calcification, skeletal density is sensitive to changes in seawater carbonate ion concentration, whereas extension is not. Despite the fact that the model was not generated for Pocillopora, the apparent strong biological control over the centers of calcification and the immediately associated fibers during calcification observed in several taxa may be a widespread phenomenon in Scleractinia [5,55–57].

Our local and regional results demonstrate a gradual decline in Pocillopora skeletal density over the past two decades, which is consistent with rising temperatures and ocean acidification. This phenomenon aligns with the documented decline in calcification observed in reef-building corals across various regions of the ocean [8]. While data regarding the extent of the affected area are currently unavailable, the coral growth model for the equatorial ETP predicts a gradual reduction in linear extension of Pocillopora, estimated at 0.9% per year, due to the effects of increased acidification [11]. The results, when considered collectively, indicate the presence of significant large-scale factors contributing to the reduction in the extent and density of Pocillopora spp. skeletons in the ETP, which in turn is leading to a decline in the CaCO₃ production in the region's reef systems. As Cabral-Tena et al. [18] have demonstrated, the potential for CaCO₃ production in the ETP reefs is maximized in monogeneric Pocillopora reefs. In light of the aforementioned points, two reasonable expectations can be proposed. The initial scenario posits a reduction in the calcification rate of Pocillopora, a genus of coral that is estimated to contribute to at least 90% of the total calcification in the region. Prior to the onset of the 2023 heat wave, carbonate balances in eastern Pacific coral ecosystems exhibited a pronounced positive trend in the eastern tropical Pacific (8.22 G) and the coast of Oaxaca (7.21 G) [18,19,58]. Consequently, a reduction in the density of the primary reef builder may have resulted in a 15–29% decline in the net calcification and reef accretion within the coral ecosystems in the eastern tropical Pacific, including the Mexican Pacific. In the absence of a skeletal extension estimate concurrent with the density estimate, an alternative hypothesis is that Pocillopora is growing more with an increase in temperature, as has been observed in other species, but the skeleton is less dense [12, and references therein]. Consequently, this phenomenon may render Pocillopora reefs more vulnerable to storm damage, wave energy, and bioerosion. Consequently, this phenomenon may potentially result in the flattening of reefs and a reduction in structural complexity observed in eastern Pacific reefs [59]. In summary, the reasonable expectation is that a decrease in calcium carbonate production will result in the possible endangerment of reefs comprising predominantly of Pocillopora in the region.

Additionally, the repercussions of ocean acidification on coral reefs, particularly with respect to coral growth, have been the focus of substantial research in recent years. Ocean acidification has been linked to reduced coral cover and diminished growth metrics in multiple coral species [5,60–63], including Pocillopora corals in Central American locations [11]. As ocean acidification intensifies, it is likely to emerge as a principal environmental stressor for corals, particularly in locations with pronounced seasonal upwelling (such as the Oaxaca area), which introduces nutrient-rich, low-pH, and low Ω_ar waters to the coastline [31,64]. In addition to the remarkable decrease in skeletal density among the primary ETP reef-building species that was observed in this study, the widespread bleaching and substantial mortality that were associated with the 2023 heatwave event have further compounded the situation. According to the data recorded in the Mexican South Pacific, all corals at all sites exhibited complete bleaching during the heatwave, with a mortality rate of 50–93% of the total coral cover [65]. The combination of declining skeletal density, as demonstrated in this study, and extensive coral mortality associated with intensifying and more frequent El Niño events [66] has the potential to disrupt the equilibrium between CaCO₃ accumulation (and the resultant structural complexity of the ecosystem) and losses through microbial, macrobiotic and chemical dissolution. This phenomenon may result in a transition from conditions conducive to elevated CaCO₃ production to those favoring reduced CaCO₃ output [67,68]. In cases where environmental pressures are particularly pronounced, a state of negative carbonate budgets, characterized by net reef erosion, could be attained [69,70]. Given that Pocillopora is the primary reef-building genus in the ETP region, our findings suggest that the long-term maintenance and reef development in the area may be compromised, potentially leading to reduced reef functionality and altered biodiversity stability, along with associated ecosystem services [19].

The resilience of reefs in the ETP is threatened by a number of factors, including rapid environmental change. Despite the changes in carbonate production and structural complexity, ETP reefs are typically highly porous and consist of uncemented accumulations of CaCO₃ and poorly cemented sediment matrix [71]. This renders them particularly susceptible to damage from storms and wave action [72]. In the event that all of the aforementioned environmental constraints to reef development in the ETP reach a critical point, the fate of reefs in the region will hinge on the resilience of the coral to adapt to a rapidly changing environment. Notwithstanding the empirical robustness of our analysis and the proposed scenarios, further studies are necessary to address the coupling between biological, chemical, and physical processes. In this regard, it is imperative to develop models that can quantify the coral response to stressors, thereby facilitating the understanding of potential positive feedbacks from calcifiers.

Potential research gaps and future work

There is a pressing need to identify trends in ocean acidification. This objective can be accomplished by maintaining monitoring efforts oriented towards generating long-term series of the carbonate system in coastal key/representative areas. Moreover, this information will facilitate the identification of the prevailing forcing and controlling factors on sea-surface temperature, pH, and aragonite saturation. This approach will also facilitate the validation of regional and local models, including the one employed in this study.

A notable aspect of this study is the utilization of historical samples for the development of a timeline of coral growth parameters for branching species. However, to obtain long-term databases, it is necessary to encourage the use of current (or custom-made) data repositories. The availability of this information and metadata will facilitate the correction of potential bias resulting from methodological approaches, the definition and adoption of quality criteria, the determination of measurement repetitiveness or precision, and the coupling with other ancillary data.

An evaluation of the effects of density and porosity is imperative. A substantial body of research has demonstrated that ocean acidification leads to an increase in porosity in specific taxa. Consequently, this results in a reduction in skeletal density and an increase in brittleness. However, it is crucial to substantiate this correlation between variables and assess its impact within the context of the ETP, which serves as a natural laboratory for ocean acidification, particularly in light of the diverse reef ecosystems that are dominated by the same Pocillopora species.

Supporting information

S1 Table. Number of fragments per morphotype sampled by locality, area, and time.

https://doi.org/10.1371/journal.pone.0342741.s001

(PDF)

S2 Fig. Surface pH (total scale; pHT) and saturation state of aragonite (Ω_ar) data compilation for both study areas.

https://doi.org/10.1371/journal.pone.0342741.s002

(PDF)

S3 Fig. Modeled pH and saturation state of aragonite (Ω_ar) for both study areas.

https://doi.org/10.1371/journal.pone.0342741.s003

(PDF)

S4 Fig. Changes in skeletal density across species, times and areas in the Mexican Pacific.

Mean (x), median (horizontal line).

https://doi.org/10.1371/journal.pone.0342741.s004

(PDF)

S5 Fig. Changes in skeletal density across times and areas in the Mexican Pacific.

Mean (x), median (horizontal line).

https://doi.org/10.1371/journal.pone.0342741.s005

(PDF)

S6 Fig. Changes in skeletal density across times in the Mexican Pacific.

Mean (x), median (horizontal line).

https://doi.org/10.1371/journal.pone.0342741.s006

(PDF)

Acknowledgments

We would like to thank all anonymous colleagues and students who diligently collected samples during field campaigns conducted at the entrance of the Gulf of California and on the coast of Oaxaca. Gerardo Barba (MHNUABCS), Gerardo Leyte (MHNUMar) provided access to museum material. OJT thanks the Department of Hydrobiology of the UAM and FMM thanks the Postgraduate Department of Marine Sciences and Limnology of the UNAM for the opportunity to study and its support.

References

1. Goldberg WM. The biology of reefs and reef organisms. Chicago: University of Chicago Press. 2013.
2. Perry CT, Spencer T, Kench PS. Carbonate budgets and reef production states: a geomorphic perspective on the ecological phase-shift concept. Coral Reefs. 2008;27(4):853–66.
- View Article
- Google Scholar
3. Lough JM, Cooper TF. New insights from coral growth band studies in an era of rapid environmental change. Earth-Science Reviews. 2011;108(3–4):170–84.
- View Article
- Google Scholar
4. Carricart-Ganivet JP, Cabanillas-Terán N, Cruz-Ortega I, Blanchon P. Sensitivity of calcification to thermal stress varies among genera of massive reef-building corals. PLoS One. 2012;7(3):e32859. pmid:22396797
- View Article
- PubMed/NCBI
- Google Scholar
5. Mollica NR, Guo W, Cohen AL, Huang K-F, Foster GL, Donald HK, et al. Ocean acidification affects coral growth by reducing skeletal density. Proc Natl Acad Sci U S A. 2018;115(8):1754–9. pmid:29378969
- View Article
- PubMed/NCBI
- Google Scholar
6. Cantin NE, Cohen AL, Karnauskas KB, Tarrant AM, McCorkle DC. Ocean warming slows coral growth in the central Red Sea. Science. 2010;329(5989):322–5. pmid:20647466
- View Article
- PubMed/NCBI
- Google Scholar
7. McCulloch MT, D’Olivo JP, Falter J, Holcomb M, Trotter JA. Coral calcification in a changing World and the interactive dynamics of pH and DIC upregulation. Nat Commun. 2017;8:15686. pmid:28555644
- View Article
- PubMed/NCBI
- Google Scholar
8. Pandolfi JM, Connolly SR, Marshall DJ, Cohen AL. Projecting coral reef futures under global warming and ocean acidification. Science. 2011;333(6041):418–22. pmid:21778392
- View Article
- PubMed/NCBI
- Google Scholar
9. Leung JYS, Zhang S, Connell SD. Is Ocean Acidification Really a Threat to Marine Calcifiers? A Systematic Review and Meta-Analysis of 980+ Studies Spanning Two Decades. Small. 2022;18(35):e2107407. pmid:35934837
- View Article
- PubMed/NCBI
- Google Scholar
10. Hoegh-Guldberg O, Mumby PJ, Hooten AJ, Steneck RS, Greenfield P, Gomez E, et al. Coral reefs under rapid climate change and ocean acidification. Science. 2007;318(5857):1737–42. pmid:18079392
- View Article
- PubMed/NCBI
- Google Scholar
11. Manzello DP. Coral growth with thermal stress and ocean acidification: lessons from the eastern tropical Pacific. Coral Reefs. 2010;29(3):749–58.
- View Article
- Google Scholar
12. Rippe JP, Baumann JH, De Leener DN, Aichelman HE, Friedlander EB, Davies SW, et al. Corals sustain growth but not skeletal density across the Florida Keys Reef Tract despite ongoing warming. Glob Chang Biol. 2018;24(11):5205–17. pmid:30102827
- View Article
- PubMed/NCBI
- Google Scholar
13. Lizcano-Sandoval LD, Marulanda-Gómez Á, López-Victoria M, Rodriguez-Ramirez A. Climate change and Atlantic Multidecadal Oscillation as drivers of recent declines in coral growth rates in the Southwestern Caribbean. Front Mar Sci. 2019;6:38.
- View Article
- Google Scholar
14. Manzello DP, Enochs IC, Bruckner A, Renaud PG, Kolodziej G, Budd DA, et al. Geophysical Research Letters. 2014;41(24):9001–8.
- View Article
- Google Scholar
15. Lough JM. Coral calcification from skeletal records revisited. Mar Ecol Prog Ser. 2008;373:257–64.
- View Article
- Google Scholar
16. Rodríguez-Martínez RE, Banaszak AT, McField MD, Beltrán-Torres AU, Álvarez-Filip L. Assessment of Acropora palmata in the Mesoamerican Reef System. PLoS One. 2014;9(4):e96140. pmid:24763319
- View Article
- PubMed/NCBI
- Google Scholar
17. Glynn PW, Alvarado JJ, Banks S, Cortés J, Feingold JS, Jiménez C, et al. Eastern Pacific coral reef provinces, coral community structure and composition: an overview. In: Glynn PW, Manzello D, Enochs I. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 107–76.
18. Cabral-Tena RA, López-Pérez A, Reyes-Bonilla H, Calderon-Aguilera LE, Norzagaray-López CO, Rodríguez-Zaragoza FA. Calcification of coral assemblages in the eastern Pacific: reshuffling calcification scenarios under climate change. Ecol Indic. 2018;95:726–34.
- View Article
- Google Scholar
19. Cabral-Tena RA, López-Pérez A, Alvarez-Filip L, González-Barrios FJ, Calderon-Aguilera LE, Aparicio-Cid C. Functional potential of coral assemblages along a typical Eastern Tropical Pacific reef tract. Ecol Indic. 2020;119:106795.
- View Article
- Google Scholar
20. Sánchez-Noguera C, Stuhldreier I, Cortés J, Jiménez C, Morales Á, Wild C, et al. Natural ocean acidification at Papagayo upwelling system (north Pacific Costa Rica): implications for reef development. Biogeosciences. 2018;15(8):2349–60.
- View Article
- Google Scholar
21. Kleypas J, Buddemeier R, Archer D, Gattuso J, Langdon C, Opdyke B. Geochemical consequences of increased atmospheric carbon dioxide on coral reefs. Science. 1999;284(5411):118–20. pmid:10102806
- View Article
- PubMed/NCBI
- Google Scholar
22. Manzello DP, Kleypas JA, Budd DA, Eakin CM, Glynn PW, Langdon C. Poorly cemented coral reefs of the eastern tropical Pacific: possible insights into reef development in a high-CO2 world. Proc Natl Acad Sci U S A. 2008;105(30):10450–5. pmid:18663220
- View Article
- PubMed/NCBI
- Google Scholar
23. Manzello DP. Ocean acidification hotspots: Spatiotemporal dynamics of the seawater CO2 system of eastern Pacific coral reefs. Limnol Oceanogr. 2010;55(1):239–48.
- View Article
- Google Scholar
24. Lavín MF, Castro R, Beier E, Godínez VM, Amador A, Guest P. SST, thermohaline structure, and circulation in the southern Gulf of California in June 2004 during the North American Monsoon Experiment. J Geophys Res. 2009;114(C2).
- View Article
- Google Scholar
25. Pantoja DA, Marinone SG, Parés-Sierra A, Gómez-Valdivia F. Modelación numérica de la hidrografía y circulación estacional y de mesoescala en el Pacífico central mexicano. Cienc Mar. 2012;38(2):363–79.
- View Article
- Google Scholar
26. Norzagaray‐López CO, Hernández‐Ayón JM, Aguilera LC, Reyes‐Bonilla H, Chapa‐Balcorta C, Ayala‐Bocos A. Aragonite saturation and pH variation in a fringing reef are strongly influenced by oceanic conditions. Limnol Oceanogr. 2017;62(6):2375–88.
- View Article
- Google Scholar
27. Chapa-Balcorta C, Sosa-Ávalos R, Hernández-Ayón JM, Espinosa-Carreón LT, Lara-Lara JR, Guerra-Mendoza RA. Almacenes y flujos en ecosistemas marinos. Estado del Ciclo del Carbono: Agenda Azul y Verde. Texcoco: Programa Mexicano del Carbono. 2019. 3–23.
28. Franco AC, Hernández-Ayón JM, Beier E, Garçon V, Maske H, Paulmier A, et al. Air-sea CO2fluxes above the stratified oxygen minimum zone in the coastal region off Mexico. J Geophys Res Oceans. 2014;119(5):2923–37.
- View Article
- Google Scholar
29. Norzagaray CO, Hernández-Ayón JM, Castro R, Calderón-Aguilera LE, Martz T, Valdivieso-Ojeda JA, et al. Seasonal controls of the carbon biogeochemistry of a fringing coral reef in the Gulf of California, Mexico. Continental Shelf Research. 2020;211:104279.
- View Article
- Google Scholar
30. Stuhldreier I, Sánchez-Noguera C, Rixen T, Cortés J, Morales A, Wild C. Effects of Seasonal Upwelling on Inorganic and Organic Matter Dynamics in the Water Column of Eastern Pacific Coral Reefs. PLoS One. 2015;10(11):e0142681. pmid:26560464
- View Article
- PubMed/NCBI
- Google Scholar
31. Chapa‐Balcorta C, Hernandez‐Ayon JM, Durazo R, Beier E, Alin SR, López‐Pérez A. Influence of post‐Tehuano oceanographic processes in the dynamics of the CO2 system in the Gulf of Tehuantepec, Mexico. JGR Oceans. 2015;120(12):7752–70.
- View Article
- Google Scholar
32. Flores-Ramírez M, Chapa-Balcorta C, López-Pérez RA, Leal-Acosta ML, García-Burciaga H. Distribución espacial del estado de saturación de aragonita y pH durante diciembre de 2020 en Isla La Blanca, Oaxaca. Estado actual del conocimiento del ciclo del carbono y sus interacciones en México: Síntesis a 2021. Monterrey: Programa Mexicano del Carbono, Instituto Tecnológico y de Estudios Superiores de Monterrey. 2021. 292–7.
33. Marting D’A, Le Tissier A. The growth and formation of branch tips of Pocillopora damicornis (Linnaeus). J Exp Mar Biol Ecol. 1988;124(2):115–31.
- View Article
- Google Scholar
34. Carricart-Ganivet JP, Beltrán-Torres AU, Merino M, Ruiz-Zárate MA. Skeletal extension, density and calcification rate of the reef building coral Montastraea annularis (Ellis and Solander) in the Mexican Caribbean. Bull Mar Sci. 2000;66(1):215–24.
- View Article
- Google Scholar
35. Lee K, Tong LT, Millero FJ, Sabine CL, Dickson AG, Goyet C. Global relationships of total alkalinity with salinity and temperature in surface waters of the world’s oceans. Geophys Res Lett. 2006;33(19).
- View Article
- Google Scholar
36. Lewis ER, Wallace DWR. Program developed for CO2 system calculations (No. cdiac: CDIAC-105). Environmental System Science Data Infrastructure for a Virtual Ecosystem (ESS-DIVE)(United States). 1988.
37. Lueker TJ, Dickson AG, Keeling CD. Ocean pCO2 calculated from dissolved inorganic carbon, alkalinity, and equations for K1 and K2: validation based on laboratory measurements of CO2 in gas and seawater at equilibrium. Marine Chemistry. 2000;70(1–3):105–19.
- View Article
- Google Scholar
38. Bakker DCE, Alin SR, Aramaki T. Surface Ocean CO2 Atlas Database Version 2025 (SOCATv2025). NOAA National Centers for Environmental Information. 2025.
- View Article
- Google Scholar
39. Wieners KH, Giorgetta M, Jungclaus J, Reick C, Esch M, Bittner M. MPI-M MPI-ESM1.2-LR model output prepared for CMIP6 CMIP historical. World Data Center for Climate (WDCC) at DKRZ. 2023. https://hdl.handle.net/21.14106/f30a8e1c2cdb373248d49bca915674687aee9421
40. Anderson M, Gorley RN, Clarke RK. Permanova for primer: Guide to software and statistical methods. Auckland: Primer-E Limited. 2008.
41. Medellín-Maldonado F, Cabral-Tena RA, López-Pérez A, Calderón-Aguilera LE, Norzagaray-López C, Chapa-Balcorta C. Calcificación de las principales especies de corales constructoras de arrecifes en la costa del Pacífico del sur de México. Cienc Mar. 2016;42(3):209–25.
- View Article
- Google Scholar
42. Norzagaray-López CO, Calderón-Aguilera LE, Alvarez-Filip L, Barranco-Servin LM, Cabral-Tena RA, Carricart-Ganivet JP. Base de datos de almacenes de carbonato de calcio en arrecifes coralinos de México. Elementos para Políticas Publicas. 2018;2:147–63.
- View Article
- Google Scholar
43. Tortolero-Langarica J de JA, Rodríguez-Troncoso AP, Cupul-Magaña AL, Carricart-Ganivet JP. Calcification and growth rate recovery of the reef-building Pocillopora species in the northeast tropical Pacific following an ENSO disturbance. PeerJ. 2017;5:e3191. pmid:28413732
- View Article
- PubMed/NCBI
- Google Scholar
44. Hammer Ø, Harper DA. Past: paleontological statistics software package for education and data analysis. John Wiley & Sons. 2001;4(1):1.
- View Article
- Google Scholar
45. González‐Pabón MA, Tortolero‐Langarica JA, Calderon‐Aguilera LE, Solana‐Arellano E, Rodríguez‐Troncoso AP, Cupul‐Magaña AL. Low calcification rate, structural complexity, and calcium carbonate production of Pocillopora corals in a biosphere reserve of the central Mexican Pacific. Mar Ecol. 2021;42(6):e12678.
- View Article
- Google Scholar
46. Coronado-Álvarez LDLA, Hernández-Ayón JM, Delgado-Contreras JA, Durazo R, Espinosa-Carreón TL, Norzagaray-López O, et al. Variability of pCO₂ and FCO₂ in the Mexican Pacific during 25 years. J Mar Syst. 2023;239:103853.
- View Article
- Google Scholar
47. Fiedler PC, Lavín MF. Oceanographic conditions of the eastern tropical Pacific. In: Glynn P, Manzello D, Enochs I. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 59–83.
48. Roik A, Wall M, Dobelmann M, Nietzer S, Brefeld D, Fiesinger A, et al. Trade-offs in a reef-building coral after six years of thermal acclimation. Sci Total Environ. 2024;949:174589. pmid:38981551
- View Article
- PubMed/NCBI
- Google Scholar
49. Houlbrèque F, Ferrier-Pagès C. Heterotrophy in tropical scleractinian corals. Biol Rev Camb Philos Soc. 2009;84(1):1–17. pmid:19046402
- View Article
- PubMed/NCBI
- Google Scholar
50. Carricart-Ganivet JP, Merino M. Growth responses of the reef-building coral Montastraea annularis along a gradient of continental influence in the southern Gulf of Mexico. Bull Mar Sci. 2001;68(1):133–46.
- View Article
- Google Scholar
51. Carricart-Ganivet JP. Sea surface temperature and the growth of the West Atlantic reef-building coral Montastraea annularis. J Exp Mar Biol Ecol. 2004;302(2):249–60.
- View Article
- Google Scholar
52. Allemand D, Tambutté É, Zoccola D, Tambutté S. Coral Calcification, Cells to Reefs. Coral Reefs: An Ecosystem in Transition. Springer Netherlands. 2010. 119–50.
- View Article
- Google Scholar
53. Erez J, Reynaud S, Silverman J, Schneider K, Allemand D. Coral Calcification Under Ocean Acidification and Global Change. Coral Reefs: An Ecosystem in Transition. Springer Netherlands. 2010. 151–76.
- View Article
- Google Scholar
54. Tambutté S, Holcomb M, Ferrier-Pagès C, Reynaud S, Tambutté É, Zoccola D, et al. Coral biomineralization: from the gene to the environment. J Exp Mar Biol Ecol. 2011;408(1–2):58–78.
- View Article
- Google Scholar
55. Cohen AL, McConnaughey TA. Geochemical perspectives on coral mineralization. Rev Mineral Geochem. 2003;54(1):151–87.
- View Article
- Google Scholar
56. Cuif JP, Dauphin Y, Doucet J, Salome M, Susini J. XANES mapping of organic sulfate in three scleractinian coral skeletons. GCA. 2003;67(1):75–83.
- View Article
- Google Scholar
57. Shirai K, Sowa K, Watanabe T, Sano Y, Nakamura T, Clode P. Visualization of sub-daily skeletal growth patterns in massive Porites corals grown in Sr-enriched seawater. J Struct Biol. 2012;180(1):47–56. pmid:22683766
- View Article
- PubMed/NCBI
- Google Scholar
58. Medellín Maldonado F. Macrobioerosion of the coral reef-building species and impact on carbonate budgets on the reefs of Huatulco, Mexico. Hidrobiológica. 2023;33(2):179–88.
- View Article
- Google Scholar
59. Alvarez-Filip L, Dulvy NK, Gill JA, Côté IM, Watkinson AR. Flattening of Caribbean coral reefs: region-wide declines in architectural complexity. Proc Biol Sci. 2009;276(1669):3019–25. pmid:19515663
- View Article
- PubMed/NCBI
- Google Scholar
60. Barkley HC, Cohen AL, Golbuu Y, Starczak VR, DeCarlo TM, Shamberger KEF. Changes in coral reef communities across a natural gradient in seawater pH. Sci Adv. 2015;1(5):e1500328. pmid:26601203
- View Article
- PubMed/NCBI
- Google Scholar
61. Enochs IC, Manzello DP, Carlton RD, Graham DM, Ruzicka R, Colella MA. Ocean acidification enhances the bioerosion of a common coral reef sponge: implications for the persistence of the Florida Reef Tract. Bull Mar Sci. 2015;91(2):271–90.
- View Article
- Google Scholar
62. Fabricius KE, Langdon C, Uthicke S, Humphrey C, Noonan S, De’ath G, et al. Losers and winners in coral reefs acclimatized to elevated carbon dioxide concentrations. Nat Clim Chang. 2011;1(3):165–9.
- View Article
- Google Scholar
63. Manzello DP, Kolodziej G, Kirkland A, Besemer N, Enochs IC. Increasing coral calcification in Orbicella faveolata and Pseudodiploria strigosa at Flower Garden Banks, Gulf of Mexico. Coral Reefs. 2021;40(4):1097–111.
- View Article
- Google Scholar
64. Pennington JT, Mahoney KL, Kuwahara VS, Kolber DD, Calienes R, Chavez FP. Primary production in the eastern tropical Pacific: A review. Prog Oceanogr. 2006;69(2–4):285–317.
- View Article
- Google Scholar
65. López-Pérez A, Granja-Fernández R, Ramírez-Chávez E, Valencia-Méndez O, Rodríguez-Zaragoza FA, González-Mendoza T, et al. Widespread Coral Bleaching and Mass Mortality of Reef-Building Corals in Southern Mexican Pacific Reefs Due to 2023 El Niño Warming. Oceans. 2024;5(2):196–209.
- View Article
- Google Scholar
66. Reimer JD, Peixoto RS, Davies SW, Traylor-Knowles N, Short ML, Cabral-Tena RA. The Fourth Global Coral Bleaching Event: Where do we go from here?. Coral Reefs. 2024;:1–5.
- View Article
- Google Scholar
67. Molina‐Hernández A, Medellín‐Maldonado F, Lange ID, Perry CT, Álvarez‐Filip L. Coral reef erosion: In situ measurement on different dead coral substrates on a Caribbean reef. Limnol Oceanogr. 2022;67(12):2734–49.
- View Article
- Google Scholar
68. Medellín-Maldonado F, Cruz-Ortega I, Pérez-Cervantes E, Norzogaray-López O, Carricart-Ganivet JP, López-Pérez A, et al. Newly deceased Caribbean reef-building corals experience rapid carbonate loss and colonization by endolithic organisms. Commun Biol. 2023;6(1):934. pmid:37699971
- View Article
- PubMed/NCBI
- Google Scholar
69. Perry CT, Alvarez-Filip L, Graham NAJ, Mumby PJ, Wilson SK, Kench PS, et al. Loss of coral reef growth capacity to track future increases in sea level. Nature. 2018;558(7710):396–400. pmid:29904103
- View Article
- PubMed/NCBI
- Google Scholar
70. Cornwall CE, Comeau S, Kornder NA, Perry CT, van Hooidonk R, DeCarlo TM, et al. Global declines in coral reef calcium carbonate production under ocean acidification and warming. Proc Natl Acad Sci U S A. 2021;118(21):e2015265118. pmid:33972407
- View Article
- PubMed/NCBI
- Google Scholar
71. Manzello DP, Eakin CM, Glynn PW. Effects of global warming and ocean acidification on carbonate budgets of Eastern Pacific coral reefs. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 517–33.
72. Lirman D, Glynn PW, Baker AC, Leyte Morales GE. Combined effects of three sequential storms on the Huatulco coral reef tract, Mexico. Bull Mar Sci. 2001;69(1):267–78.
- View Article
- Google Scholar

[ref1] 1. Goldberg WM. The biology of reefs and reef organisms. Chicago: University of Chicago Press. 2013.

[ref2] 2. Perry CT, Spencer T, Kench PS. Carbonate budgets and reef production states: a geomorphic perspective on the ecological phase-shift concept. Coral Reefs. 2008;27(4):853–66.
View Article
Google Scholar

[3] View Article

[4] Google Scholar

[ref3] 3. Lough JM, Cooper TF. New insights from coral growth band studies in an era of rapid environmental change. Earth-Science Reviews. 2011;108(3–4):170–84.
View Article
Google Scholar

[6] View Article

[7] Google Scholar

[ref4] 4. Carricart-Ganivet JP, Cabanillas-Terán N, Cruz-Ortega I, Blanchon P. Sensitivity of calcification to thermal stress varies among genera of massive reef-building corals. PLoS One. 2012;7(3):e32859. pmid:22396797
View Article
PubMed/NCBI
Google Scholar

[9] View Article

[10] PubMed/NCBI

[11] Google Scholar

[ref5] 5. Mollica NR, Guo W, Cohen AL, Huang K-F, Foster GL, Donald HK, et al. Ocean acidification affects coral growth by reducing skeletal density. Proc Natl Acad Sci U S A. 2018;115(8):1754–9. pmid:29378969
View Article
PubMed/NCBI
Google Scholar

[13] View Article

[14] PubMed/NCBI

[15] Google Scholar

[ref6] 6. Cantin NE, Cohen AL, Karnauskas KB, Tarrant AM, McCorkle DC. Ocean warming slows coral growth in the central Red Sea. Science. 2010;329(5989):322–5. pmid:20647466
View Article
PubMed/NCBI
Google Scholar

[17] View Article

[18] PubMed/NCBI

[19] Google Scholar

[ref7] 7. McCulloch MT, D’Olivo JP, Falter J, Holcomb M, Trotter JA. Coral calcification in a changing World and the interactive dynamics of pH and DIC upregulation. Nat Commun. 2017;8:15686. pmid:28555644
View Article
PubMed/NCBI
Google Scholar

[21] View Article

[22] PubMed/NCBI

[23] Google Scholar

[ref8] 8. Pandolfi JM, Connolly SR, Marshall DJ, Cohen AL. Projecting coral reef futures under global warming and ocean acidification. Science. 2011;333(6041):418–22. pmid:21778392
View Article
PubMed/NCBI
Google Scholar

[25] View Article

[26] PubMed/NCBI

[27] Google Scholar

[ref9] 9. Leung JYS, Zhang S, Connell SD. Is Ocean Acidification Really a Threat to Marine Calcifiers? A Systematic Review and Meta-Analysis of 980+ Studies Spanning Two Decades. Small. 2022;18(35):e2107407. pmid:35934837
View Article
PubMed/NCBI
Google Scholar

[29] View Article

[30] PubMed/NCBI

[31] Google Scholar

[ref10] 10. Hoegh-Guldberg O, Mumby PJ, Hooten AJ, Steneck RS, Greenfield P, Gomez E, et al. Coral reefs under rapid climate change and ocean acidification. Science. 2007;318(5857):1737–42. pmid:18079392
View Article
PubMed/NCBI
Google Scholar

[33] View Article

[34] PubMed/NCBI

[35] Google Scholar

[ref11] 11. Manzello DP. Coral growth with thermal stress and ocean acidification: lessons from the eastern tropical Pacific. Coral Reefs. 2010;29(3):749–58.
View Article
Google Scholar

[37] View Article

[38] Google Scholar

[ref12] 12. Rippe JP, Baumann JH, De Leener DN, Aichelman HE, Friedlander EB, Davies SW, et al. Corals sustain growth but not skeletal density across the Florida Keys Reef Tract despite ongoing warming. Glob Chang Biol. 2018;24(11):5205–17. pmid:30102827
View Article
PubMed/NCBI
Google Scholar

[40] View Article

[41] PubMed/NCBI

[42] Google Scholar

[ref13] 13. Lizcano-Sandoval LD, Marulanda-Gómez Á, López-Victoria M, Rodriguez-Ramirez A. Climate change and Atlantic Multidecadal Oscillation as drivers of recent declines in coral growth rates in the Southwestern Caribbean. Front Mar Sci. 2019;6:38.
View Article
Google Scholar

[44] View Article

[45] Google Scholar

[ref14] 14. Manzello DP, Enochs IC, Bruckner A, Renaud PG, Kolodziej G, Budd DA, et al. Geophysical Research Letters. 2014;41(24):9001–8.
View Article
Google Scholar

[47] View Article

[48] Google Scholar

[ref15] 15. Lough JM. Coral calcification from skeletal records revisited. Mar Ecol Prog Ser. 2008;373:257–64.
View Article
Google Scholar

[50] View Article

[51] Google Scholar

[ref16] 16. Rodríguez-Martínez RE, Banaszak AT, McField MD, Beltrán-Torres AU, Álvarez-Filip L. Assessment of Acropora palmata in the Mesoamerican Reef System. PLoS One. 2014;9(4):e96140. pmid:24763319
View Article
PubMed/NCBI
Google Scholar

[53] View Article

[54] PubMed/NCBI

[55] Google Scholar

[ref17] 17. Glynn PW, Alvarado JJ, Banks S, Cortés J, Feingold JS, Jiménez C, et al. Eastern Pacific coral reef provinces, coral community structure and composition: an overview. In: Glynn PW, Manzello D, Enochs I. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 107–76.

[ref18] 18. Cabral-Tena RA, López-Pérez A, Reyes-Bonilla H, Calderon-Aguilera LE, Norzagaray-López CO, Rodríguez-Zaragoza FA. Calcification of coral assemblages in the eastern Pacific: reshuffling calcification scenarios under climate change. Ecol Indic. 2018;95:726–34.
View Article
Google Scholar

[58] View Article

[59] Google Scholar

[ref19] 19. Cabral-Tena RA, López-Pérez A, Alvarez-Filip L, González-Barrios FJ, Calderon-Aguilera LE, Aparicio-Cid C. Functional potential of coral assemblages along a typical Eastern Tropical Pacific reef tract. Ecol Indic. 2020;119:106795.
View Article
Google Scholar

[61] View Article

[62] Google Scholar

[ref20] 20. Sánchez-Noguera C, Stuhldreier I, Cortés J, Jiménez C, Morales Á, Wild C, et al. Natural ocean acidification at Papagayo upwelling system (north Pacific Costa Rica): implications for reef development. Biogeosciences. 2018;15(8):2349–60.
View Article
Google Scholar

[64] View Article

[65] Google Scholar

[ref21] 21. Kleypas J, Buddemeier R, Archer D, Gattuso J, Langdon C, Opdyke B. Geochemical consequences of increased atmospheric carbon dioxide on coral reefs. Science. 1999;284(5411):118–20. pmid:10102806
View Article
PubMed/NCBI
Google Scholar

[67] View Article

[68] PubMed/NCBI

[69] Google Scholar

[ref22] 22. Manzello DP, Kleypas JA, Budd DA, Eakin CM, Glynn PW, Langdon C. Poorly cemented coral reefs of the eastern tropical Pacific: possible insights into reef development in a high-CO2 world. Proc Natl Acad Sci U S A. 2008;105(30):10450–5. pmid:18663220
View Article
PubMed/NCBI
Google Scholar

[71] View Article

[72] PubMed/NCBI

[73] Google Scholar

[ref23] 23. Manzello DP. Ocean acidification hotspots: Spatiotemporal dynamics of the seawater CO2 system of eastern Pacific coral reefs. Limnol Oceanogr. 2010;55(1):239–48.
View Article
Google Scholar

[75] View Article

[76] Google Scholar

[ref24] 24. Lavín MF, Castro R, Beier E, Godínez VM, Amador A, Guest P. SST, thermohaline structure, and circulation in the southern Gulf of California in June 2004 during the North American Monsoon Experiment. J Geophys Res. 2009;114(C2).
View Article
Google Scholar

[78] View Article

[79] Google Scholar

[ref25] 25. Pantoja DA, Marinone SG, Parés-Sierra A, Gómez-Valdivia F. Modelación numérica de la hidrografía y circulación estacional y de mesoescala en el Pacífico central mexicano. Cienc Mar. 2012;38(2):363–79.
View Article
Google Scholar

[81] View Article

[82] Google Scholar

[ref26] 26. Norzagaray‐López CO, Hernández‐Ayón JM, Aguilera LC, Reyes‐Bonilla H, Chapa‐Balcorta C, Ayala‐Bocos A. Aragonite saturation and pH variation in a fringing reef are strongly influenced by oceanic conditions. Limnol Oceanogr. 2017;62(6):2375–88.
View Article
Google Scholar

[84] View Article

[85] Google Scholar

[ref27] 27. Chapa-Balcorta C, Sosa-Ávalos R, Hernández-Ayón JM, Espinosa-Carreón LT, Lara-Lara JR, Guerra-Mendoza RA. Almacenes y flujos en ecosistemas marinos. Estado del Ciclo del Carbono: Agenda Azul y Verde. Texcoco: Programa Mexicano del Carbono. 2019. 3–23.

[ref28] 28. Franco AC, Hernández-Ayón JM, Beier E, Garçon V, Maske H, Paulmier A, et al. Air-sea CO2fluxes above the stratified oxygen minimum zone in the coastal region off Mexico. J Geophys Res Oceans. 2014;119(5):2923–37.
View Article
Google Scholar

[88] View Article

[89] Google Scholar

[ref29] 29. Norzagaray CO, Hernández-Ayón JM, Castro R, Calderón-Aguilera LE, Martz T, Valdivieso-Ojeda JA, et al. Seasonal controls of the carbon biogeochemistry of a fringing coral reef in the Gulf of California, Mexico. Continental Shelf Research. 2020;211:104279.
View Article
Google Scholar

[91] View Article

[92] Google Scholar

[ref30] 30. Stuhldreier I, Sánchez-Noguera C, Rixen T, Cortés J, Morales A, Wild C. Effects of Seasonal Upwelling on Inorganic and Organic Matter Dynamics in the Water Column of Eastern Pacific Coral Reefs. PLoS One. 2015;10(11):e0142681. pmid:26560464
View Article
PubMed/NCBI
Google Scholar

[94] View Article

[95] PubMed/NCBI

[96] Google Scholar

[ref31] 31. Chapa‐Balcorta C, Hernandez‐Ayon JM, Durazo R, Beier E, Alin SR, López‐Pérez A. Influence of post‐Tehuano oceanographic processes in the dynamics of the CO2 system in the Gulf of Tehuantepec, Mexico. JGR Oceans. 2015;120(12):7752–70.
View Article
Google Scholar

[98] View Article

[99] Google Scholar

[ref32] 32. Flores-Ramírez M, Chapa-Balcorta C, López-Pérez RA, Leal-Acosta ML, García-Burciaga H. Distribución espacial del estado de saturación de aragonita y pH durante diciembre de 2020 en Isla La Blanca, Oaxaca. Estado actual del conocimiento del ciclo del carbono y sus interacciones en México: Síntesis a 2021. Monterrey: Programa Mexicano del Carbono, Instituto Tecnológico y de Estudios Superiores de Monterrey. 2021. 292–7.

[ref33] 33. Marting D’A, Le Tissier A. The growth and formation of branch tips of Pocillopora damicornis (Linnaeus). J Exp Mar Biol Ecol. 1988;124(2):115–31.
View Article
Google Scholar

[102] View Article

[103] Google Scholar

[ref34] 34. Carricart-Ganivet JP, Beltrán-Torres AU, Merino M, Ruiz-Zárate MA. Skeletal extension, density and calcification rate of the reef building coral Montastraea annularis (Ellis and Solander) in the Mexican Caribbean. Bull Mar Sci. 2000;66(1):215–24.
View Article
Google Scholar

[105] View Article

[106] Google Scholar

[ref35] 35. Lee K, Tong LT, Millero FJ, Sabine CL, Dickson AG, Goyet C. Global relationships of total alkalinity with salinity and temperature in surface waters of the world’s oceans. Geophys Res Lett. 2006;33(19).
View Article
Google Scholar

[108] View Article

[109] Google Scholar

[ref36] 36. Lewis ER, Wallace DWR. Program developed for CO2 system calculations (No. cdiac: CDIAC-105). Environmental System Science Data Infrastructure for a Virtual Ecosystem (ESS-DIVE)(United States). 1988.

[ref37] 37. Lueker TJ, Dickson AG, Keeling CD. Ocean pCO2 calculated from dissolved inorganic carbon, alkalinity, and equations for K1 and K2: validation based on laboratory measurements of CO2 in gas and seawater at equilibrium. Marine Chemistry. 2000;70(1–3):105–19.
View Article
Google Scholar

[112] View Article

[113] Google Scholar

[ref38] 38. Bakker DCE, Alin SR, Aramaki T. Surface Ocean CO2 Atlas Database Version 2025 (SOCATv2025). NOAA National Centers for Environmental Information. 2025.
View Article
Google Scholar

[115] View Article

[116] Google Scholar

[ref39] 39. Wieners KH, Giorgetta M, Jungclaus J, Reick C, Esch M, Bittner M. MPI-M MPI-ESM1.2-LR model output prepared for CMIP6 CMIP historical. World Data Center for Climate (WDCC) at DKRZ. 2023. https://hdl.handle.net/21.14106/f30a8e1c2cdb373248d49bca915674687aee9421

[ref40] 40. Anderson M, Gorley RN, Clarke RK. Permanova for primer: Guide to software and statistical methods. Auckland: Primer-E Limited. 2008.

[ref41] 41. Medellín-Maldonado F, Cabral-Tena RA, López-Pérez A, Calderón-Aguilera LE, Norzagaray-López C, Chapa-Balcorta C. Calcificación de las principales especies de corales constructoras de arrecifes en la costa del Pacífico del sur de México. Cienc Mar. 2016;42(3):209–25.
View Article
Google Scholar

[120] View Article

[121] Google Scholar

[ref42] 42. Norzagaray-López CO, Calderón-Aguilera LE, Alvarez-Filip L, Barranco-Servin LM, Cabral-Tena RA, Carricart-Ganivet JP. Base de datos de almacenes de carbonato de calcio en arrecifes coralinos de México. Elementos para Políticas Publicas. 2018;2:147–63.
View Article
Google Scholar

[123] View Article

[124] Google Scholar

[ref43] 43. Tortolero-Langarica J de JA, Rodríguez-Troncoso AP, Cupul-Magaña AL, Carricart-Ganivet JP. Calcification and growth rate recovery of the reef-building Pocillopora species in the northeast tropical Pacific following an ENSO disturbance. PeerJ. 2017;5:e3191. pmid:28413732
View Article
PubMed/NCBI
Google Scholar

[126] View Article

[127] PubMed/NCBI

[128] Google Scholar

[ref44] 44. Hammer Ø, Harper DA. Past: paleontological statistics software package for education and data analysis. John Wiley & Sons. 2001;4(1):1.
View Article
Google Scholar

[130] View Article

[131] Google Scholar

[ref45] 45. González‐Pabón MA, Tortolero‐Langarica JA, Calderon‐Aguilera LE, Solana‐Arellano E, Rodríguez‐Troncoso AP, Cupul‐Magaña AL. Low calcification rate, structural complexity, and calcium carbonate production of Pocillopora corals in a biosphere reserve of the central Mexican Pacific. Mar Ecol. 2021;42(6):e12678.
View Article
Google Scholar

[133] View Article

[134] Google Scholar

[ref46] 46. Coronado-Álvarez LDLA, Hernández-Ayón JM, Delgado-Contreras JA, Durazo R, Espinosa-Carreón TL, Norzagaray-López O, et al. Variability of pCO₂ and FCO₂ in the Mexican Pacific during 25 years. J Mar Syst. 2023;239:103853.
View Article
Google Scholar

[136] View Article

[137] Google Scholar

[ref47] 47. Fiedler PC, Lavín MF. Oceanographic conditions of the eastern tropical Pacific. In: Glynn P, Manzello D, Enochs I. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 59–83.

[ref48] 48. Roik A, Wall M, Dobelmann M, Nietzer S, Brefeld D, Fiesinger A, et al. Trade-offs in a reef-building coral after six years of thermal acclimation. Sci Total Environ. 2024;949:174589. pmid:38981551
View Article
PubMed/NCBI
Google Scholar

[140] View Article

[141] PubMed/NCBI

[142] Google Scholar

[ref49] 49. Houlbrèque F, Ferrier-Pagès C. Heterotrophy in tropical scleractinian corals. Biol Rev Camb Philos Soc. 2009;84(1):1–17. pmid:19046402
View Article
PubMed/NCBI
Google Scholar

[144] View Article

[145] PubMed/NCBI

[146] Google Scholar

[ref50] 50. Carricart-Ganivet JP, Merino M. Growth responses of the reef-building coral Montastraea annularis along a gradient of continental influence in the southern Gulf of Mexico. Bull Mar Sci. 2001;68(1):133–46.
View Article
Google Scholar

[148] View Article

[149] Google Scholar

[ref51] 51. Carricart-Ganivet JP. Sea surface temperature and the growth of the West Atlantic reef-building coral Montastraea annularis. J Exp Mar Biol Ecol. 2004;302(2):249–60.
View Article
Google Scholar

[151] View Article

[152] Google Scholar

[ref52] 52. Allemand D, Tambutté É, Zoccola D, Tambutté S. Coral Calcification, Cells to Reefs. Coral Reefs: An Ecosystem in Transition. Springer Netherlands. 2010. 119–50.
View Article
Google Scholar

[154] View Article

[155] Google Scholar

[ref53] 53. Erez J, Reynaud S, Silverman J, Schneider K, Allemand D. Coral Calcification Under Ocean Acidification and Global Change. Coral Reefs: An Ecosystem in Transition. Springer Netherlands. 2010. 151–76.
View Article
Google Scholar

[157] View Article

[158] Google Scholar

[ref54] 54. Tambutté S, Holcomb M, Ferrier-Pagès C, Reynaud S, Tambutté É, Zoccola D, et al. Coral biomineralization: from the gene to the environment. J Exp Mar Biol Ecol. 2011;408(1–2):58–78.
View Article
Google Scholar

[160] View Article

[161] Google Scholar

[ref55] 55. Cohen AL, McConnaughey TA. Geochemical perspectives on coral mineralization. Rev Mineral Geochem. 2003;54(1):151–87.
View Article
Google Scholar

[163] View Article

[164] Google Scholar

[ref56] 56. Cuif JP, Dauphin Y, Doucet J, Salome M, Susini J. XANES mapping of organic sulfate in three scleractinian coral skeletons. GCA. 2003;67(1):75–83.
View Article
Google Scholar

[166] View Article

[167] Google Scholar

[ref57] 57. Shirai K, Sowa K, Watanabe T, Sano Y, Nakamura T, Clode P. Visualization of sub-daily skeletal growth patterns in massive Porites corals grown in Sr-enriched seawater. J Struct Biol. 2012;180(1):47–56. pmid:22683766
View Article
PubMed/NCBI
Google Scholar

[169] View Article

[170] PubMed/NCBI

[171] Google Scholar

[ref58] 58. Medellín Maldonado F. Macrobioerosion of the coral reef-building species and impact on carbonate budgets on the reefs of Huatulco, Mexico. Hidrobiológica. 2023;33(2):179–88.
View Article
Google Scholar

[173] View Article

[174] Google Scholar

[ref59] 59. Alvarez-Filip L, Dulvy NK, Gill JA, Côté IM, Watkinson AR. Flattening of Caribbean coral reefs: region-wide declines in architectural complexity. Proc Biol Sci. 2009;276(1669):3019–25. pmid:19515663
View Article
PubMed/NCBI
Google Scholar

[176] View Article

[177] PubMed/NCBI

[178] Google Scholar

[ref60] 60. Barkley HC, Cohen AL, Golbuu Y, Starczak VR, DeCarlo TM, Shamberger KEF. Changes in coral reef communities across a natural gradient in seawater pH. Sci Adv. 2015;1(5):e1500328. pmid:26601203
View Article
PubMed/NCBI
Google Scholar

[180] View Article

[181] PubMed/NCBI

[182] Google Scholar

[ref61] 61. Enochs IC, Manzello DP, Carlton RD, Graham DM, Ruzicka R, Colella MA. Ocean acidification enhances the bioerosion of a common coral reef sponge: implications for the persistence of the Florida Reef Tract. Bull Mar Sci. 2015;91(2):271–90.
View Article
Google Scholar

[184] View Article

[185] Google Scholar

[ref62] 62. Fabricius KE, Langdon C, Uthicke S, Humphrey C, Noonan S, De’ath G, et al. Losers and winners in coral reefs acclimatized to elevated carbon dioxide concentrations. Nat Clim Chang. 2011;1(3):165–9.
View Article
Google Scholar

[187] View Article

[188] Google Scholar

[ref63] 63. Manzello DP, Kolodziej G, Kirkland A, Besemer N, Enochs IC. Increasing coral calcification in Orbicella faveolata and Pseudodiploria strigosa at Flower Garden Banks, Gulf of Mexico. Coral Reefs. 2021;40(4):1097–111.
View Article
Google Scholar

[190] View Article

[191] Google Scholar

[ref64] 64. Pennington JT, Mahoney KL, Kuwahara VS, Kolber DD, Calienes R, Chavez FP. Primary production in the eastern tropical Pacific: A review. Prog Oceanogr. 2006;69(2–4):285–317.
View Article
Google Scholar

[193] View Article

[194] Google Scholar

[ref65] 65. López-Pérez A, Granja-Fernández R, Ramírez-Chávez E, Valencia-Méndez O, Rodríguez-Zaragoza FA, González-Mendoza T, et al. Widespread Coral Bleaching and Mass Mortality of Reef-Building Corals in Southern Mexican Pacific Reefs Due to 2023 El Niño Warming. Oceans. 2024;5(2):196–209.
View Article
Google Scholar

[196] View Article

[197] Google Scholar

[ref66] 66. Reimer JD, Peixoto RS, Davies SW, Traylor-Knowles N, Short ML, Cabral-Tena RA. The Fourth Global Coral Bleaching Event: Where do we go from here?. Coral Reefs. 2024;:1–5.
View Article
Google Scholar

[199] View Article

[200] Google Scholar

[ref67] 67. Molina‐Hernández A, Medellín‐Maldonado F, Lange ID, Perry CT, Álvarez‐Filip L. Coral reef erosion: In situ measurement on different dead coral substrates on a Caribbean reef. Limnol Oceanogr. 2022;67(12):2734–49.
View Article
Google Scholar

[202] View Article

[203] Google Scholar

[ref68] 68. Medellín-Maldonado F, Cruz-Ortega I, Pérez-Cervantes E, Norzogaray-López O, Carricart-Ganivet JP, López-Pérez A, et al. Newly deceased Caribbean reef-building corals experience rapid carbonate loss and colonization by endolithic organisms. Commun Biol. 2023;6(1):934. pmid:37699971
View Article
PubMed/NCBI
Google Scholar

[205] View Article

[206] PubMed/NCBI

[207] Google Scholar

[ref69] 69. Perry CT, Alvarez-Filip L, Graham NAJ, Mumby PJ, Wilson SK, Kench PS, et al. Loss of coral reef growth capacity to track future increases in sea level. Nature. 2018;558(7710):396–400. pmid:29904103
View Article
PubMed/NCBI
Google Scholar

[209] View Article

[210] PubMed/NCBI

[211] Google Scholar

[ref70] 70. Cornwall CE, Comeau S, Kornder NA, Perry CT, van Hooidonk R, DeCarlo TM, et al. Global declines in coral reef calcium carbonate production under ocean acidification and warming. Proc Natl Acad Sci U S A. 2021;118(21):e2015265118. pmid:33972407
View Article
PubMed/NCBI
Google Scholar

[213] View Article

[214] PubMed/NCBI

[215] Google Scholar

[ref71] 71. Manzello DP, Eakin CM, Glynn PW. Effects of global warming and ocean acidification on carbonate budgets of Eastern Pacific coral reefs. Coral Reefs of the Eastern Tropical Pacific. Dordrecht: Springer. 2017. 517–33.

[ref72] 72. Lirman D, Glynn PW, Baker AC, Leyte Morales GE. Combined effects of three sequential storms on the Huatulco coral reef tract, Mexico. Bull Mar Sci. 2001;69(1):267–78.
View Article
Google Scholar

[218] View Article

[219] Google Scholar

Figures

Abstract

Introduction

Materials and methods

Study area

Coral sampling

Density estimations

Environmental setting

Data analysis

Results

Discussion

Potential research gaps and future work

Supporting information

S1 Table. Number of fragments per morphotype sampled by locality, area, and time.

S2 Fig. Surface pH (total scale; pHT) and saturation state of aragonite (Ωar) data compilation for both study areas.

S3 Fig. Modeled pH and saturation state of aragonite (Ωar) for both study areas.

S4 Fig. Changes in skeletal density across species, times and areas in the Mexican Pacific.

S5 Fig. Changes in skeletal density across times and areas in the Mexican Pacific.

S6 Fig. Changes in skeletal density across times in the Mexican Pacific.

Acknowledgments

References

S2 Fig. Surface pH (total scale; pHT) and saturation state of aragonite (Ω_ar) data compilation for both study areas.

S3 Fig. Modeled pH and saturation state of aragonite (Ω_ar) for both study areas.