Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

Active restoration efforts in the Central Mexican Pacific

as a strategy for coral reef recovery

Violeta Martínez-Castillo1; https://orcid.org/0000-0003-3932-4646

Alma Paola Rodríguez-Troncoso1*; https://orcid.org/0000-0001-6243-7679

José de Jesús Adolfo Tortolero-Langarica2,3; https://orcid.org/0000-0001-8857-5789

Amílcar Leví Cupul-Magaña1; https://orcid.org/0000-0002-6455-1253

1. Laboratorio de Ecología Marina, Centro Universitario de la Costa, Universidad de Guadalajara, Puerto Vallarta,

Jalisco, México, 48280; viomarcast@gmail.com, pao.rodriguezt@gmail.com (*Correspondence),

amilcar.cupul@gmail.com

2. Laboratorio de Esclerocronología de Corales Pétreos, Unidad Académica de Sistemas Arrecifales, Instituto de

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

77580; adolfo.tl@bahia.tecnm.mx

3. Tecnológico Nacional de México / IT de Bahía de Banderas, Bahía de Banderas, Bahía de Banderas, Nayarit, México,

63734.

Received 03-X-2022. Corrected 06-II-2023. Accepted 07-II-2023.

ABSTRACT

Introduction: The 1997-98 El Niño event caused massive coral bleaching and mortality in the Central Mexican

Pacific (CMP). Punta de Mita alone used to harbor more than 30 % of the coral coverage in this region, with a

mono-specific Pocillopora coverage. The 1997-1998 ENSO event caused massive coral mortality reducing live

coral coverage to < 5 %. Despite being considered a coral region unlikely to recover, recent restoration efforts

have been implemented to rehabilitate the coral community.

Objective: To assess coral recovery by analyzing the coral growth and survival rates of branching Pocillopora

species at Punta de Mita.

Methods: Healthy coral fragments of opportunity were re-attached to the natural substrata using zip ties and

measured considering their growth in terms of maximum length and width (cm) to determine their annual exten-

sion rates.

Results: After 50 weeks, corals duplicated their size, with a mean growth of ~ 4 cm year-1. After 100 weeks (2

years), corals triplicated their size, increasing on average 8–9 cm in each diameter.

Conclusions: Successful coral reef restoration activities in the Central Mexican Pacific are the result of

Pocillopora’s physiological processes, such as fast growth rates, and recent life-history traits, like the ability to

cope with thermal anomalies, which enable them to thrive in a dynamic region severely affected by natural and

anthropogenic perturbations. Indeed, a region considered unlikely to recover has regained its live coral cover

from < 5 % in 1998 up to 15 % in 20 years. This demonstrates the importance of assisting natural coral recovery

with restoration efforts, especially in coral locations that, despite environmental perturbations, have proven to be

resilient and may become coral refugia areas under the current climate change scenario.

Key words: Scleractinia; Pocillopora; ecosystem restoration; coral growth; active coral restoration.

https://doi.org/10.15517/rev.biol.trop..v71iS1.54795

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

INTRODUCTION

Interest in coral restoration has increased

in the last two decades due to the rapid global

decline of coral reefs (Hoegh-Guldberg et al.,

2007; Hughes et al., 2018). More recently,

a special interest in the use of coral species

or genotypes more tolerant to fluctuations in

environmental conditions has emerged, as ther-

mal anomalies associated with ENSO events

are expected to increase in both frequency

and intensity (Hughes et al., 2018; Rinkevich,

2014), reducing the time that these ecosys-

tems have to recover from such disturbances

(Boström-Einarsson et al., 2020). Therefore, it

is important to identify those coral ecosystems

that harbor scleractinian species capable of

withstanding future disturbances from climate

change and human activities, as they represent

future coral refugia and may become sources of

coral fragments for coral restoration programs.

Identifying resilient communities requires eco-

logical, physiological, and molecular studies

to ensure that restoration strategies maintain

and improve current restored areas and at

least maintain coral diversity at ecological and

genetic scales (Boström-Einarsoon et al., 2020;

Montoya-Maya et al., 2016).

Within this context, it is important to pri-

oritize resilient coral communities impacted by

anthropogenic disturbances and climate change

to evaluate their suitability to be restored and,

eventually, rehabilitated. Along the Eastern

Tropical Pacific (ETP), local coral commu-

nities have developed rapid acclimatization

capacity and ability to remain stable over

several years, despite being under the influ-

ence of moderate and severe El Niño Southern

RESUMEN

Esfuerzos de restauración activa en el Pacífico Central Mexicano

como estrategia para la recuperación de los arrecifes de coral

Introducción: El evento El Niño de 1997-98 causó un blanqueamiento y mortalidad masiva de corales en el

Pacífico Central Mexicano (CMP). Solo Punta de Mita albergaba más del 30 % de la cobertura coralina de esta

región, con una cobertura monoespecífica de Pocillopora. El evento ENSO de 1997-1998 causó una mortalidad

masiva de corales que redujo la cobertura de corales vivos a < 5 %. A pesar de ser considerada una región de

coral con pocas probabilidades de recuperarse, se han implementado esfuerzos de restauración recientes para

rehabilitar la comunidad coralina.

Objetivo: Evaluar la recuperación de coral analizando el crecimiento coralino y las tasas de supervivencia de

especies ramificadas de Pocillopora en Punta de Mita.

Métodos: Fragmentos de oportunidad de coral sanos se volvieron a unir a los sustratos naturales usando bridas

y se midieron considerando su crecimiento en términos de longitud y ancho máximos (cm) para determinar sus

tasas de extensión anual.

Resultados: Después de 50 semanas, los corales duplicaron su tamaño, con un crecimiento promedio de ~ 4 cm

año-1. Después de 100 semanas (2 años), los corales triplicaron su tamaño, aumentando en promedio 8–9 cm en

cada diámetro.

Conclusiones: Las actividades exitosas de restauración de arrecifes de coral en el Pacífico Central Mexicano

son el resultado de los procesos fisiológicos de Pocillopora, tales como tasas de crecimiento rápido, y rasgos de

historia de vida reciente, como la capacidad de hacer frente a anomalías térmicas, que les permiten prosperar en

una región dinámica severamente afectada por perturbaciones naturales y antropogénicas. De hecho, esta región

que se consideraba poco probable que se recuperara, ha recuperado su cobertura de coral vivo de < 5 % en 1998

hasta 15 % en 20 años. Esto demuestra la importancia de ayudar a la recuperación natural de los corales con

los esfuerzos de restauración, especialmente en las ubicaciones de corales que, a pesar de las perturbaciones

ambientales, han demostrado ser resistentes y pueden convertirse en áreas de refugio de corales ante el escenario

actual de cambio climático.

Palabras clave: Scleractinia; Pocillopora; restauración de ecosistemas; crecimiento de corales; restauración de

corales.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

Oscillation (ENSO) events that have triggered

bleaching, but no mortality in recent years

(Cruz-García et al., 2020; Nava et al., 2021;

Romero-Torres et al., 2020). Within the ETP,

Punta de Mita (located in the Central Mexican

Pacific, CMP) harbors a coral community that

historically lost over 90 % of its live coral

cover during the 1997-98 El Niño massive

bleaching and mortality event (Carriquiry et

al., 2001), which caused researchers to con-

sider its recovery uncertain. However, recent

life-history traits of corals in the area have pro-

moted an increase in their ability to cope and

resist different stressful conditions: not only

has been observed thermal tolerance thresh-

old but also a natural recovery capacity as

well (Martínez-Castillo, Rodríguez-Troncoso,

Mayfield et al., 2022; Tortolero-Langarica et

al., 2017), which results in an opportunity to

implement restoration strategies that contribute

to the already observed recovery (Tortolero-

Langarica et al., 2014; Tortolero-Langarica et

al., 2017), implementing protocols that allow

it to be site-specific since coral reefs do not

thrive under the same conditions, especially

under the current climate change scenario.

Here we evaluate coral reef recovery resulting

from active restoration at Punta de Mita, a site

that has shown a natural recovery capacity from

natural and anthropogenic disturbances.

MATERIALS AND METHODS

Study site: Punta de Mita (20.7699 °N,

105.5412° W) is located within the CMP

(Fig. 1), a region considered an oceanographic

transition zone where three water masses con-

verge: the cold California Current flowing

southernly, the warm coastal Mexican current

flowing northerly, and the warm and saline

water from the Gulf of California flowing

southernly (Pantoja et al., 2012). The con-

vergence of these currents creates a dynamic

region, in addition to seasonal upwellings,

Fig. 1. A. The Central Mexican Pacific (CMP), B. Punta de Mita, North of Banderas Bay within the CMP, C. Coral

restoration site at Punta de Mita.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

nutrient pulses, and internal waves, wide daily

temperature fluctuations (± 5 °C) can be

recorded (Plata & Filonov, 2007; Portela et al.,

2016). Punta de Mita presents a well-formed

coral reef patch that used to harbor ~ 38 % of

live coral cover, which was impacted by the

1997-98 ENSO event resulting in a dramatic

loss of this ecosystem (Carriquiry & Reyes-

Bonilla, 1997; Carriquiry et al., 2001).

Coral restoration and monitoring: Coral

restoration was performed in two different

phases at a coral reef community that is at

three meters depth: 1) the first in 2013 in order

to validate the protocol, and 2) the second

phase three years later in 2016. In each phase,

a total of 50 Pocillopora fragments of oppor-

tunity, defined as fragments detached from an

adult colony through natural processes such

as water-motion, or bioerosion (Monty et al.,

2006), were collected. Each coral fragment was

individually examined to avoid signs of bleach-

ing, damage, or algae overgrowth (such as turf

and macroalgae) and then fixed to the natural

substrate using zip ties within the same reef site

(Fig. 2). All coral fragments restored during

2013 were tagged and measured for a two-year

period (from November 2013 to October 2015),

while those restored in 2016 were measured for

one year (from January 2016 to February 2017)

to record coral growth, i.e., the increase of

fragment size (cm) maximum length (ML) and

maximum width (MW) using a caliper (0.05

mm of precision). Also, survival of fragments

was recorded.

Data analysis: Overall mean values of

coral growth (± standard error) were calculated

using Statistica 8.0 software (StatSoft Inc.,

2007) and expressed as cm year-1. Coral growth

rates, defined as the average total increase in

coral length and width, were calculated using

Fig. 2. Pocillopora spp. fragment of opportunity fixed to the natural substrata in Punta de Mita, México A. initial fragment

size during the first restoration period (2013), and B. Second restoration period (2016). Final colonies size at the end of the

C. first restoration (2015) and D. second restoration period (2017).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

the original fragment size (cm ± standard error)

for each restoration period. Also, the average

and cumulative size increase in length and

width (cm) within each period was estimated.

Simple linear regressions were calculated

to evaluate the effect of changes in SST on

coral growth rates. Mean monthly sea surface

temperatures (SST) were obtained from the

GIOVANNI database of the National Aeronau-

tics and Space Administration of the United

States (Li & Hegde, 2022). Regressions were

calculated using Statistica 8.0 software (Stat-

Soft Inc., 2007).

RESULTS

The overall mean coral growth rate was

4.25 ± 0.15 cm year-1 for maximum length and

3.71 ± 0.17 cm year-1 for maximum width.

Further details of each restoration period are

given below.

First restoration period: Coral fragments

tripled their size after 100 weeks of being reat-

tached with an accumulated growth in length

and width of ∼ 8-9 cm (Fig. 3A). During the

first restoration phase, corals grew at a rate of

∼ 3.5 cm year-1 (Table 1). While coral growth

remained stable throughout the first restoration

period, the highest coral growth in both axes

occurred during the warmest season (August

2015). The increase in coral size remained at

3.79 ± 0.21 and 3.55 ± 0.40 cm year-1 for both

ML and MW (Table 1, Fig. 3B). Coral frag-

ments self-attached to the substrata one month

after being restored, and the final proportion of

surviving corals was 38 % (Fig. 4).

Fig. 3. A. Mean fragment size during the first restoration period (2013). B. Increase in coral size during the first period

between measuring times. C. Mean fragment size during the second restoration period (2016). D. Increase in coral size

during the second period between measuring times. Bars = SE. Red rectangles mark ENSO weeks during the first restoration

phase.

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Second restoration period: Coral frag-

ments almost doubled their size after 57 weeks

following their re-attachment (Fig. 3C), with a

maximum accumulated growth of ∼ 5-6 cm in

both length and width. The increase in coral

size remained at 4.49 ± 0.19 and 3.78 ± 0.16 cm

year-1 for both ML and MW (Table 1), peaking

around week 30, during the warm season of

2016 (August; Fig. 3D). Corals self-attached to

the substrate within the first two months after

their restoration, and the proportion of surviv-

ing corals after 56 weeks was 76 % (Fig. 4).

Sea surface temperature and coral

growth: During the first restoration period,

the mean monthly sea surface temperature

ranged from 25.7 to 32.3 °C, whereas during

the second period, the mean monthly SST

ranged from 24.1 to 31.3 °C, coinciding with

an El Niño event, causing thermal anomalies

≥ 2.8 °C (Fig. 5). When assessing the effects

of sea temperature on coral growth parameters,

both regressions were not significant; hence,

coral growth was not significantly related to

sea surface temperature (R = 0.51, P = 0.10).

DISCUSSION

The capacity of any ecosystem to recover

from environmental perturbations depends on

the ability of its organisms to adapt to new con-

ditions, which is determined by their physiolog-

ical traits, including regulatory mechanisms,

performance, and environmental tolerance

(Cooke & Suski, 2008); specifically for coral

ecosystems, these physiological traits include

coral growth rates and reproductive strategies,

their acclimatization capacity, and the type of

endosymbiont they harbor, among others. The

Fig. 4. Survival of restored Pocillopora spp. coral fragments during both restoration periods in Punta de Mita, México. Red

asterisks signal ENSO weeks during the 2015-2016 El Niño event.

TABLE 1

Coral growth rates in length and width (cm year-1) in each restoration period. ML: maximum length; MW: maximum width

Restoration period ML MW

Mean Min Max Mean Min Max

First (2013) 3.79 ± 0.21 1.97 5.38 3.55 ± 0.40 0.90 7.14

Second (2016) 4.49 ± 0.19 1.94 7.33 3.78 ± 0.16 1.63 6.17

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

Central Mexican Pacific is a region considered

sub-optimal for coral development due to its

wide ranges in temperature, nutrient puls-

es, and elevated sedimentation rates (Glynn,

2017). It is also a region where in the last two

decades, at least three severe thermal anomalies

have affected coral communities such as Punta

de Mita (Hughes et al., 2018). In spite of this,

recent studies have documented that live coral

cover in this region is slowly recovering (Mar-

tínez-Castillo, Rodríguez-Troncoso, Mayfield

et al., 2022), evidencing the potential of this

particular site to be rehabilitated using active

restoration protocols.

Coral fragments in our study almost dou-

bled their size one year following their re-

attachment and increased three times their size

after two years. Pocillopora has been character-

ized as a coral genus with one of the highest

growth rates within the EP, a key physiological

trait that has allowed this genus to remain as

the most abundant species (Romero-Torres et

al., 2020; Tortolero-Langarica et al., 2017).

Mean extension rates of corals are ~ 3.5–4.5

cm y-1, which will not only increase the frag-

ment length or width but in the development

of the three-dimensional complexity that other

coral colonies have exhibited elsewhere with-

in the CMP (González-Pabón et al., 2021;

Tortolero-Langarica et al., 2014; Tortolero-

Langarica et al., 2017; Tortolero-Langarica et

al., 2020). Indeed, corals not only continued to

grow, but they also exhibited extension rates

that are among the highest recorded in the

Eastern Tropical Pacific (Martínez-Castillo,

Rodríguez-Troncoso, Mayfield et al., 2022);

therefore, corals within this region increase

coral cover and carbonate production, con-

tributing to the recovery of these reef sites

(Tortolero-Langarica et al., 2019).

Restoration, in its broad sense, aims to

assist the recovery of a damaged or degraded

ecosystem, and particularly for coral reefs,

their recovery relays on the increase of live

coral cover both in its area and in its three-

dimensional structure, which directly depends

on sustained coral growth (Rinkevich, 2014).

The fact that restored corals continued to grow

even during high thermal anomalies caused by

recent ENSO events (as seen in this study),

brings up the relevance of their physiologi-

cal traits such as the acclimatization capacity,

Fig. 5. Thermal anomalies (°C) caused by ENSO events across both restoration periods. The red frame signals positive

thermal anomalies during the most recent and severe El Niño event. Thermal anomalies correspond to the Oceanic Niño

Index (ONI) from the National Oceanic and Atmospheric Administration (NOAA, 2022).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

which have allowed corals at Punta de Mita to

recover and remain stable under stress events,

highlighting the CMP as an important EP ref-

uge, not only for corals, but also for the organ-

isms associated with them (Martínez-Castillo,

2020; Martínez-Castillo, Rodríguez-Troncoso,

Mayfield et al., 2022; Rodríguez-Troncoso et

al., 2016; Tortolero-Langarica et al., 2017).

Coral growth is influenced by abiotic fac-

tors from which sea surface temperature is

considered the most important, and the thresh-

old is determined by the local and regional

acclimatization capacity of these organisms

(Manzello et al., 2010; Martínez-Castillo et

al., 2020). More important high temperatures

also trigger bleaching response and may cause

mortalities as observed previously in the region

(Carriquiry et al., 2001); however, there was no

significant relationship between coral growth

or survival and sea temperature even under

the influence of the most severe ENSO event

recorded to date. After almost 60 weeks from

coral re-attachment, survival rates in both res-

toration periods were above ~ 60 %, which is

considered as successful as most restoration

projects report between 60-70 % of survival

(Boström-Einarsson et al., 2020), and it was

even higher than other remote locations of

the CMP with low influence of human activ-

ity (Tortolero-Langarica et al., 2020). Unlike

those areas, Punta de Mita coral community

is a coastal site 200 m distant from the shore,

where there are luxury touristic complexes and

gulf camps, with intense marine traffic and no

regulation of touristic activities. The high mor-

tality observed at the end of the first restora-

tion period then may be due to two interacting

factors: first, even though coral growth was not

compromised by the 2015-2016 El Niño warm

temperatures, the coral community must have

suffered individual negative effects during its

first stage (which coincides with the end of the

first restoration period); second anthropogenic

activity in Punta de Mita may also have played a

role in coral mortality as this site lacks an offi-

cial protection status. In spite of this, the growth

rates of the surviving coral colonies were suf-

ficient enough to prevent the loss of coral cover

during this period, reinforcing the idea that the

CMP harbor resilient coral communities (Cruz-

García et al., 2020; Martínez-Castillo et al.,

2020; Martínez-Castillo, Rodríguez-Troncoso,

Tortolero-Langarica et al., 2022; Rodríguez-

Troncoso et al., 2016; Tortolero-Langarica et

al., 2017; Tortolero-Langarica et al., 2022). In

this sense, the coral community at Punta de

Mita may represent an important coral refuge

in the near future as corals’ in the region exhibit

a high acclimatization capacity, a key feature of

corals in the region that promotes the recovery

of CMP coral ecosystems (Martínez-Castillo et

al., 2020; Martínez-Castillo, Rodríguez-Tron-

coso, Mayfield et al., 2022; Martínez-Castillo,

Rodríguez-Troncoso, Tortolero-Langarica et

al., 2022; Rodríguez-Troncoso et al., 2016;

Tortolero-Langarica et al., 2017; this study).

While restoration efforts at Punta de Mita

have been successful so far, there is an urgent

need for management policies to ensure that

full ecosystem recovery can be achieved. Punta

de Mita is located within Bandera’s Bay, in an

area that is planned to continue high-end urban

development (Merchand-Rojas, 2012). A major

threat to coral reefs are changes in land use

caused by urban growth (Burke et al., 2011;

Carpenter et al., 2008). Therefore, if there is

no adequate management and regulation of

recreative aquatic activities in Punta de Mita,

corals will be resisting more stressors than they

can withstand, and therefore, local or official

protection of this resilient site is urgent. The

present study gives insight into the efforts of

coral restoration activities in the Central Mexi-

can Pacific. It highlights the importance of

considering ecological, biological, and physi-

ological processes when developing restoration

strategies and emphasizes the importance of

identifying coral communities with evidence of

both resistance and resilience capacity. In this

sense, Punta de Mita represents an important

site, as both natural and restored fragments

have contributed to increase and maintain coral

cover in a region unlikely to recover from ther-

mal anomalies (Carriquiry et al., 2001). More-

over, recent ENSO effects have not negatively

affected fragment growth rates, suggesting that

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71 (S1): e54795, abril 2023 (Publicado Abr. 30, 2023)

this location is essential for coral recovery and

a conservation priority as corals are more likely

to withstand future environmental perturba-

tions than other Eastern Pacific locations due to

their biological and physiological traits.

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 study was funded by two

National Geographic grants to APRT (W405-15

and NGS-55349R-19 and by the project ‘‘Res-

tauración de Arrecifes Coralinos en el PN Islas

Marietas’’ (PROCER/CCER/DROPC/09/2016)

to ALCM. The authors kindly thank the civil

association “Protección y Restauración de Islas

y Zonas Naturales” (PROZONA, A.C.) and the

Marine Ecology Laboratory (LEMAC-UdG)

for their assistance in field operations.

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