Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

The contribution of assisted coral restoration to calcium carbonate

production in Eastern Pacific reefs

J. J. Adolfo Tortolero-Langarica1,2*; https://orcid.org/0000-0001-8857-5789

Alma P. Rodríguez-Troncoso3; https://orcid.org/0000-0001-6243-7679

Lorenzo Alvarez-Filip4; https://orcid.org/0000-0002-5726-7238

Amílcar L. Cupul-Magaña3; https://orcid.org/0000-0002-6455-1253

Juan P. Carricart-Ganivet2; https://orcid.org/0000-0001-7266-8905

1. Laboratorio de Esclerocronología de Corales Arrecifales, Unidad Académica de Sistemas Arrecifales, Instituto de

Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, Puerto Morelos, Quintana Roo, México;

adolfo.tl@bahia.tecnm.mx (*Correspondence), carricart@cmarl.unam.mx

2. Tecnológico Nacional de México / Instituto Tecnológico Bahía de Banderas, Nayarit, México;

adolfo.tl@bahia.tecnm.mx (*Correspondence).

3. Laboratorio de Ecología Marina, Centro de Investigaciones Costeras, Centro Universitario de la Costa, Universidad

de Guadalajara. Puerto Vallarta, Jalisco, México; pao.rodriguezt@gmail.com, amilcar.cupul@gmail.com

4. Laboratorio de Biodiversidad y Conservación de Arrecifes, Unidad Académica de Sistemas Arrecifales, Instituto de

Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, Puerto Morelos, Quintana. Roo, México;

lorenzo@cmarl.unam.mx

Received 29-VIII-2022. Corrected 27-I-2023. Accepted 10-II-2023.

ABSTRACT

Introduction: Hermatypic corals have the capacity to construct the physical reef-framework and maintain the

balance of coral reef functionality. However, in the past three decades, coral communities have been menaced

by natural and anthropic pressures, resulting in an abrupt coral cover decline, and slow natural recovery. To

mitigate coral reef collapse, assisted restoration techniques has been implemented and improved worldwide,

However, the long-term effects of such interventions on ecological attributes have been scarcely reported.

Objective: This study evaluated the effect of assisted coral intervention on calcium carbonate production

(kg CaCO3 m-2 yr-1) and ecological volume (cm3) yielded by branching and massive corals from the central

Mexican Pacific.

Methods: We used colony size, extension rate, and skeletal density measurements of direct outplanted

Pocillopora and Pavona coral species to calculate coral carbonate production, ecological volume, and model

their long-term potential.

Results: Coral carbonate produced after one-year of outplanting increased by 42 % (1.17 kg CaCO3 m-2 yr-1),

where Pocillopora spp. and Pavona clavus corals contribute with 0.97 and 0.20 kg CaCO3 m-2 yr-1, respectively.

The ecological volume also increased by 384 cm3 for Pocillopora and 56 cm3 for Pavona after one year period.

Furthermore, the results suggest that long-term coral restoration actions (10 years) have the potential to signifi-

cantly increase carbonate production.

Conclusions: our data indicate that coral restoration initiatives have the potential to help mitigate the current low

calcium carbonate production of Mexican Pacific reefs and may significantly contribute to the long-term main-

tenance of reef-framework based on ecological engineering tools, such initiatives represent essential functional

properties related to reef ecosystem services provision.

Keywords: direct propagation; coral fragments; eastern tropical Pacific; branching corals; massive corals.

https://doi.org/10.15517/rev.biol.trop..v71iS1.54849

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

INTRODUCTION

The construction and stability of coral reef

frameworks primarily rely on hermatypic cor-

als, which are capable of precipitating calcium

carbonate and build complex structures, which

are the base of coral reef habitats (Sheppard

et al., 2009). Yet, the combined impact of

climate-induced factors (e.g., ocean warming,

and acidification) and local human-induced

pressures (e.g., marine pollution, overfishing,

and nutrients load) has led to a rapid and relent-

less decline in coral cover and reef health. This

trend poses a significant threat to the provi-

sion of reef ecosystem services in the coming

decades (Hoegh-Guldberg et al., 2007; Hughes

et al., 2017). While passive management mea-

sures like marine protected areas (MPAs) and

marine reserve designations have been estab-

lished as key conservation strategies, they

appear to be insufficient in mitigating natural

and anthropogenic threats (Graham et al., 2008;

Selig & Bruno, 2010). As a complementary

response, active human interventions, mainly

through coral restoration approaches, based

on adapting to changing conditions and imple-

menting science-based improvements to miti-

gate coral reef degradation, have been widely

performed (reviewed in Rinkevich, 2019b).

Among the various methods used to restore

reef-building corals, one that has shown rapid

growth rates and resilience is coral propagation

through fragmentation or micro-fragmentation,

which can be applied to most hermatypic mor-

phospecies (i.e., branching, massive, columnar,

and foliaceous), reducing time and costs effec-

tively, and can be efficiently used in large-

scale restoration programs (Page et al., 2018;

Tortolero-Langarica et al., 2020). Small coral

fragments can be produced in large quanti-

ties, and the possibility of employing multiple

genotypes, which can be combined via fusion

RESUMEN

El aporte de la restauración coralina asistida a la producción de carbonato calcio

en arrecifes del Pacífico Oriental

Introducción: Los corales hermatípicos tienen la capacidad de construir y mantener la estructura física y

mantener el equilibrio de la funcionalidad de los arrecifes coralinos. Sin embargo, en las últimas tres décadas

las comunidades coralinas han sido amenazadas por presiones tanto naturales como antrópicas, resultando en

una disminución abrupta en la cobertura de coral y lenta recuperación natural. Para mitigar el colapso de los

arrecifes de coral, diversas técnicas de restauración asistida se han implementado y mejorado alrededor del

mundo. Sin embargo, los efectos de largo plazo de dichas intervenciones en los atributos ecológicos han sido

escasamente reportado.

Objetivo: Este estudio evaluó el efecto de la intervención asistida en la producción de carbonato (kgCaCO3 m-2

yr-1) y en el volumen ecológico (cm3) producido por corales ramificados y masivos del Pacífico central mexicano.

Métodos: Utilizamos mediciones de tamaño de colonias, la tasa de extensión y densidad esquelética de especies

de coral Pocillopora y Pavona trasplantadas directamente para calcular la producción de carbonato de coral, el

volumen ecológico y modelar su potencial a largo plazo.

Resultados: El carbonato de coral producido después de un año de la plantación aumentó un 42 % (1.17 kg

CaCO3 m-2 yr-1), donde los corales Pocillopora spp. y Pavona clavus aportaron 0.97 y 0.20 kg CaCO3 m-2 yr-1,

respectivamente. El volumen ecológico también aumentó en 384 cm3 para Pocillopora y 56 cm3 para Pavona

después de un periodo anual. Los resultados sugieren que las acciones de restauración de coral a largo-plazo (10

años) tienen el potencial de aumentar significativamente la producción de carbonato.

Conclusiones: Los datos obtenidos en este estudio indican que las iniciativas de restauración de coral tienen el

potencial de ayudar a mitigar la baja producción de carbonato de calcio actual en los arrecifes del Pacífico mexi-

cano, y pueden contribuir significativamente al mantenimiento a largo-plazo de la estructura arrecifal mediante

herramientas de ingeniería ecológica. Las cuales representan propiedades funcionales esenciales relacionadas

con la provisión de servicios ecosistémicos.

Palabras clave: propagación directa; fragmentos de coral; Pacífico tropical oriental; corales ramificados; corales

masivos.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

to create chimeras, can maximize the adaptive

potential of corals to become more resilient to

future environmental conditions (Rinkevich,

2019a). However, the biological and ecologi-

cal effects of coral restoration through micro-

fragmentation are still in the initial stages,

thus evaluation of ecological restoration appli-

cations and development of integrative pro-

tocols are needed to accelerate coral reef

resilience (Rinkevich, 2019b; Boström-Einars-

son et al., 2020).

The use of ecological engineering tools

in coral restoration actions (Rinkevich, 2020)

represents an important aspect in the re-

establishment of functional properties (e.g.,

calcification rates, carbonate production, and

structural complexity) related to coral reef

ecosystem services (i.e., habitat development,

coastal protection, sand production, and track

sea-level rise) at long-term scales (Perry &

Alvarez-Filip, 2018). Yet, most coral restora-

tion methods focus on biological attributes

(e.g., coral growth and survival), while few

interventions have been attempted to measure

ecological attributes that could determine the

influence of coral restoration approaches from

a geo-ecological perspective (Rinkevich et al.,

2019b; Rinkevich, 2020).

Coral reefs from the Eastern Tropical

Pacific (ETP) region have been damaged due

to repeated intensive thermal-anomalies and

bleaching events, causing a coral cover decline

of 50-98 % in the last two decades (Alvarado

et al., 2020; Carriquiry et al., 2001; Eakin,

2001; Glynn, 2000; Reyes-Bonilla et al., 2002;

Romero-Torres et al., 2020). Despite this dam-

age, coral reef species in the ETP have revealed

a high thermo-tolerance threshold, resulting in

different natural recovery paths among coral

reefs (Glynn et al., 2015; Glynn et al., 2017;

Hueerkamp et al., 2001; Romero-Torres et al.,

2020). The recovery of reef-building coral spe-

cies is essential for sustaining ecological pro-

cesses such as reef carbonate production and

the topographical complexity of coral reef eco-

systems (Lange et al., 2020; Perry & Alvarez-

Filip, 2018). However, coral calcium carbonate

and the three-dimensional contribution of coral

species are rarely considered when determining

coral restoration efficiency from an ecological

perspective (Forsman et al., 2006; Tortolero-

Langarica et al., 2020). This study aims to

evaluate the bio-geological effect on calcium

carbonate production (kgCaCO3 m-2 yr-1) and

ecological space occupied (ecological volume;

cm3) by corals resulting from an active coral

restoration, using the direct propagation of

small fragments of branching Pocillopora spp.

and massive Pavona clavus corals from the

Central Mexican Pacific. The results provide

us insights not only into the carbonate contribu-

tion of the most abundant coral species (Pocil-

loporids and Pavonids) along the ETP, but also

for their long-term potential to maintain the

coral reef framework in the region.

MATERIALS AND METHODS

Study area: This study was conducted

between June 2018 to August 2019 at Isla

María Cleofas, Islas Marías Biosphere reserve

(Fig. 1), located in the north ETP at a distance

of 130 km offshore of the coast (CONANP,

2010). Isla María Cleofas consists of isolated

coral reef patches dominated by branching cor-

als of the family Pocilloporidae in shallow reef

areas at 2-6 m, and massive corals of the family

Agariicidae and Poritidae corals at depths > 6

m (López-Pérez et al., 2015; Tortolero-Lan-

garica et al., 2022). The annual seawater tem-

perature (SWT) of the Islas Marías archipelago

is regulated by two inter-annual ocean currents,

and it varies from 18-21 °C during December

to May when the California Current is domi-

nant, and from 27-31 °C from July through

November when the coastal Costa Rica current

prevails (Pennington et al., 2006). The Islas

Marías are also affected by frequent heatwaves

associated with El Niño Southern-Oscillation

(ENSO) events, seasonal upwelling, and tropi-

cal storms (Pennington et al., 2006; Wang &

Feldler, 2006).

Coral propagation and coral measure-

ments: Hermatypic corals propagation was

implemented using 154 small coral fragments

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

(sizes 3-4 cm2) collected from corals of oppor-

tunity of Pocillopora spp. (n = 78), and Pavona

clavus (n = 78), and glued using water-resistant

silicon (MS-express, Fischer®) directly onto

the natural limestone rock (see, Tortolero-Lan-

garica et al., 2020). The coral fragments were

placed on two coral plots of 6 m2 at a depth of

5 m in the Regidor-2 reef site (Fig. 2). Colony

size (height, length, and width) was measured

using calipers (0.05 mm precision) at the

beginning and end of the experimental period.

Height growth (H) was calculated as the linear

distances from the bottom to the uppermost

growth of each coral fragment. Length growth

(L) was determined as the maximum diameter

of the coral fragment, while width growth (W)

was obtained from the contiguous diameter

perpendicular to the length growth.

The three-dimensional growth of coral

fragments was estimated based on the vol-

ume of a cylinder since this most accurately

expressed the total space occupied by the

colony (Shafir et al., 2006). Growth metrics (H,

L, and W) were used to estimate the increase

in coral ecological volume (EV) of each frag-

ment at the end of the evaluation period, EV =

πr2H, where r = radius (L+W /4) (Rinkevich

& Loya, 1983; Shafir et al., 2006). Following

the exponential growth rates (%) of outplanted

corals, growth rate constants (k) of ecologi-

cal volumes (EV) were calculated for each

Pocillopora and Pavona clavus coral using the

formula EVt = EVoekt, providing k = (ln EVt/

EVo)/t, where t = time and 0 = values at initial

stage of the experiment.

To compare the carbonate production of

outplanted corals with that of naturally occur-

ring corals, measurements of colony size (H,

L, and W) measurements were recorded at the

genus level (Pocillopora spp. and Pavona spp.)

in five shallow reef sites (3-5 m depth; Fig. 1),

using coral colonies found along belt transects

Fig. 1. Study map of the Islas Marías Biosphere Reserve in the Northeastern Tropical Pacific. The green stars indicate the

surveyed reef sites on Isla María Cleofas (IMC), a. Regidor 1, b. Regidor 2, c. Muelas, d. Cleofas 1, e. Cleofas 2.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

(n = 5) of 25 m in length. To complete the car-

bonate production assessment, information on

coral growth (extension rate and skeletal den-

sity) was obtained from published literature for

Pocillopora (2.74 cm-2 yr-1, and 2.18 cm-3) and

Pavona (0.92 cm-2 yr-1, and 1.39 cm-3) corals

from Islas María Cleofas (Tortolero-Langarica

et al. 2020).

Colony size, extension rate and skeletal

density data were used to calculate the cal-

cium carbonate production per coral colony/

transplant (kgCaCO3 m-2 yr-1), defined as the

calcium carbonate produced by corals based

on colony morphology following the method

described by Perry et al. (2018). Annual coral

carbonate production (kgCaCO3 m-2 yr-1) was

estimated using both Pocillopora and Pavona

coral genera at Cleofa’s reef sites and coral

restoration plots, by means of the census-based

method (Lange et al., 2020; Perry et al., 2018).

The proportion of calcium carbonate produced

during the coral intervention period was also

Fig. 2. Coral restoration plots were established using small coral fragments of A. Pavona clavus and B. Pocillopora spp.

from Isla María Cleofas, Mexico. Panels (C-D) show the initial size of fragments of massive Pavona clavus and branching

Pocillopora spp. respectively. Panels (E-F) Show the final coral sizes after one-year of growth. Scale bars in panels (A-B)

represent the area size, while those in panels (C-F) indicate the size of the nearest fragment.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

calculated as CP= Nt–N0 / t, where N0 and Nt

denote the carbonate production after outplant-

ing and at the end of the experiment, respec-

tively, and (t) is the time-lapse. The calcium

carbonate produced by the intervention was

added to the naturally produced annual coral

carbonate in Cleofas and extrapolated for a

ten-year period to estimate the potential of

calcium carbonate production over a decade.

After evaluating of normality (Shapiro-Wilk,

P <0.001) and homoscedasticity (Levine, P=

0.621), differences in annual carbonate produc-

tion between reef sites was assessed using a

non-parametric analysis of variance (Kruskal-

Wallis) with Sigma plot ver.11, using a confi-

dence interval of 95 % (alfa = 0.05).

RESULTS

Coral carbonate production after inter-

vention: From the 154 total fragments that

were outplanted, the survival rate was of 59 %

(92 corals survived, including 46 Pocillopora,

and n = 46 Pavona). Coral growth parameters

indicated that coral height increased threefold

(~ 3.52 cm), width doubled (~ 5.00 cm), and

length increase twofold (~ 5.52 cm) compared

with initial size of Pocillopora. For massive

Pavona clavus fragments, there was an equita-

ble increase (1.5-fold) in all growth dimensions

(height ~ 1.35 cm, width ~1 .73 cm, and length

~ 1.69 cm). At the end of the study period, the

mean colony calcification (± SD) for Pocil-

lopora corals was of 19.58 ± 4.13 kg CaCO3

m-2 yr-1, and mean ecological volume (EV)

increased more than 15-fold, accumulating to

384 cm3 (EV initial = 25 cm3, and EV final =

409 cm3). For P. clavus fragments, the mean

colony calcification was 3.97 ± 0.63 kg CaCO3

m-2 yr-1, EV increased fourfold, accumulating

to a total of 56 cm3 (EV initial = 22 cm3, and

EV final = 78 cm3) after 13-months of inter-

vention (Fig. 2). The total calcium carbonate

at the beginning of the experiment was 2.80 kg

CaCO3 m-2 yr-1 compared to 3.98 kg CaCO3 m-2

yr-1 at the end of the experiment, with Pocil-

lopora and Pavona corals contributing 3.59

and 0.38 kg CaCO3 m-2 yr-1, respectively (Fig.

2). The amount of calcium carbonate produced

by coral outplants was 0.97 kg CaCO3 m-2

yr-1 for Pocillopora spp. and 0.20 kg CaCO3

m-2 yr-1 for P. clavus coral fragments, carbon-

ate production increased by 26 % and 54 %,

respectively (Fig. 3).

The naturally carbonate production:

In 2018, Isla Cleofas’s sites showed a mean

annual coral carbonate production of 1.78 kg

CaCO3 m-2 yr-1 (ranged 0.43-6.40), with signifi-

cant differences among reef sites (H4 = 11.028,

P = 0.026). Among the sites, Cleofas 1 reef

had the highest production (6.40 kg CaCO3 m-2

yr-1), while Las Muelas had the lowest (0.48

kg CaCO3 m-2 yr-1; Fig. 4). By including the

calcium carbonate produced by coral outplants

in natural Isla Cleofas’ carbonate production,

the potential carbonate production increased by

4.2 times for 5 years and 7.5 times for 10 years,

using active coral restoration actions (Fig. 5).

DISCUSSION

The slow or absent natural coral recovery

at many reef locations necessitates the imple-

mentation of reliable ecological restoration

methods to mitigate local coral degradation and

restore reef-associated biodiversity and ecosys-

tem functionality (Boström-Einarsson et al.,

Fig. 3. The production of calcium carbonate accumulated

by branching and massive corals resulted from the

intervention. The colored bars indicate the proportion of

carbonate production at the initial stage (blue) and after one

year of growth (green).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

2020; Rinkevich et al., 2020). The results pre-

sented here provide an ecological insight into

coral carbonate production and its response

resulting from coral restoration efforts. Direct

relocation of asexual recruits (corals of oppor-

tunity) is not commonly used (Rinkevich

2019b), it has been shown to be a practical

and effective tool for restoring coral reef-

framework and improving reef biogeochemical

dynamics in terms of coral carbonate produc-

tion (Tortolero-Langarica et al., 2019). This

technique is both time- and cost-effective,

avoiding the need for fragment transportation,

over-manipulation, pre-growth, and acclimati-

zation periods required in both the farming and

nursery phases. It is, therefore, an affordable

technique for restoring remote and marginal

reefs (Tortolero-Langarica et al., 2020). While

Fig. 4. The annual coral carbonate production between Cleofas´s reef sites. The blue bars represent naturally produced

carbonate production, while the single green bar represents carbonate produced by outplanted corals at the intervened site.

Fig. 5. The potential coral carbonate production (kgCaCO3 m-2 yr-1) estimated at the beginning of the experiment and

projected over 10 years using values produced by coral intervention. The blue segmented line represents the maximum

carbonate potential for Eastern Tropical Pacific Reef (sensu Alvarado et al., 2016; Cabral-Tena et al., 2018).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

complementary long-term approaches are

required, the direct outplanting of both branch-

ing and massive coral species used in this study

facilitates the increment of ecological volume

(EV) and coral carbonate production during

active coral restoration actions.

The results showed an accumulated eco-

logical volume of 384 cm3 yr-1 for Pocillopora

spp. and 56 cm3 yr-1 for Pavona clavus, which

is similar increments observeded with homolo-

gous species from other coral reef locations

(Boch and Morse, 2012; Shaish et al., 2010;

Tortolero-Langarica et al., 2019). Pocillopora

spp. fragments increased more than ten-fold in

EV, while P clavus fragments increased three-

fold in contrast with their initial EV sizes in a

13-month period. This corroborates that direct

propagation techniques allows the develop-

ment of three-dimensional colonies and an

increase in the structural complexity of coral

reefs, which is critical for ecological restora-

tion (Tortolero-Langarica et al., 2019; Rinkev-

ich, 2020). The use of branching and massive

morphologies in restoration programs benefits

not only coral diversity and abundance, but

also avoids morpho-specific reefs, increasing

habitat types, substrata, and resource variabil-

ity that promote ecosystem properties recov-

ery (e.g., biodiversity recruitment, food webs,

landscape, and soundscape) (Lamont et al.,

2021; Rinkevich, 2020).

Live coral cover and carbonate produc-

tion from the ETP region have both suffered

a decline in the last thirty-years mainly due

to multiple ENSO events (El Niño and La

Niña) (Manzello et al., 2017; Romero-Torres

et al., 2020; Tortolero-Langarica et al., 2022).

Although coral reefs in the study area have pre-

sented slow recovery in coral accretion follow-

ing thermal-stress periods, the recovery is not

sufficient (Cabral-Tena et al., 2018; González-

Pabón et al., 2021; Tortolero-Langarica et al.,

2017); yet some Cleofas´ reefs have been resil-

ient, acting as a nursery areas and reefs of hope.

Thus, Cleofas´ coral reef has the potential to

recover coral growth and carbonate production

rates via active restoration (Tortolero-Langari-

ca et al., 2022). This is congruent with coral

carbonate production obtained in this study

(1.78 kg CaCO3 m-2 yr-1), which is three-times

higher compared the previous year (0.46 kg

CaCO3 m-2 yr-1; González-Pabón et al., 2021).

However, Cleofas´s carbonate production is

still low compared to the carbonate potential

estimated for ETP reefs (8.22-11.47 kg CaCO3

m-2 yr-1; Alvarado et al., 2016; Cabral-Tena et

al., 2018, 2020) and those net values (> 5 kg

CaCO3 m-2 yr-1) considered for healthy reef sys-

tems (Perry et al, 2012; Lange & Perry, 2019).

This production variability can be attributed to

the decrease in coral calcification during and

following coral bleaching events, where coral

growth rate recovery may take from months to

years after organisms’ functions are repaired

(Allemand et al., 2011; Tortolero-Langarica et

al., 2017). For a successful long-term restora-

tion, it is important to employ coral species

with a wide tolerance and resistance to local

stressors, which have more opportunities to

resilience to global stressors such as thermal

anomalies (Guest et al., 2011; Montero-Serra et

al., 2018; Tortolero-Langarica et al., 2019). It is

also crucial to conduct maintenance campaigns

after thermal anomalies and hurricane events to

retrieve the survival of the restoration program.

Novel restoration approaches based on

ecological engineering can help to accelerate

the recovery of calcium carbonate production in

damaged and degraded coral reefs (Rinkevich,

2019b; Rinkevich, 2020). These approaches

include the direct relocation of coral fragments,

which promotes optimal effects on coral car-

bonate production with a high potential of suc-

cess in long-term and large-scale interventions,

relatively low-cost and less-labor (Tortolero-

Langarica et al., 2019; Tortolero-Langarica et

al., 2020). This idea is consistent with the coral

carbonate production rates observed in this

study, which showed higher annual values than

those naturally produced (3.98 kg CaCO3 m-2

yr-1 vs 1.78 kg CaCO3 m-2 yr-1) over the same

period (Fig. 4, Fig. 5). This assertion can be

also corroborated with the potential forcarbon-

ate production resulting from this study, where

the specific values for Pocillopora and Pavona

values (Fig. 5) are similar to the carbonate pro-

duction modeled for future coral assemblage

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54849, abril 2023 (Publicado Abr. 30, 2023)

in the ETP (sensu Cabral-Tena et al., 2018).

Therefore, the continued effect of human inter-

vention through active coral restoration, in

conjunction with an integrative conservation

strategy (i.e., management, self-sustainability,

outreach, government, and community owner-

ship) may be a cornerstone of effective coral

community conservation and the prevention

coral degradation in the current and future

states of coral reef ecosystems (Boström-Ein-

arsson et al., 2020; Suding et al., 2015).

The direct relocation of reef-building spe-

cies, such as massive P. clavus and branching

Pocillopora spp. from the central Mexican

Pacific, has resulted in reliable and effec-

tive ecological restoration. This is due to the

positive effect on coral ecological volume

and carbonate production, which may con-

tribute significantly to coral recovery and the

persistence of the reef-framework (Tortolero-

Langarica et al., 2019; Tortolero-Langarica et

al., 2020). The use of small coral fragments is

an alternative practical ecological engineering

technique that has proved to be a long-term

intervention initiative in the Isla Marías archi-

pelago, and a promising tool to be adopted in

other ETP coral reef localities. Moreover, the

development and scale-up of active restoration

programs are imperative to both, short- and

long-term maintenance and functionality of

reef ecosystems, as remediation and a comple-

mentary strategy under the current global coral

reef crisis.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are

fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

The present work was supported by the

Consejo Nacional de Ciencia y Tecnología

(CONACYT) postdoctoral fellowship (CVU

410380) to JJATL, and the project PAPIIT

IN200420 to JPCG. We thank to the Mexican

authorities of Reserva de la Biosfera Islas

Marias (SEMARNAT/CONANP), and Secre-

taría de Gobierno (SEGOB) for the permits

and facilities provided. We especially loke to

thank to the organization Protección y Restau-

ración de Islas y Zonas Naturales (PROZONA

AC), La Punta Outdoors SA de CV, and Grupo

Cleofas “Maria Cleofas vessel” and their crew

for their accommodations and assistance during

field expeditions.

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