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REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
38 | P a g e VOLUME 34 NUMBER 1 December 2019
Spread of the new coral disease “SCTLD” into the Caribbean:
implications for Puerto Rico
E Weil1, EA Hernández-Delgado2,3,4, M Gonzalez5,6, S Williams7, S Suleimán-
Ramos4, M Figuerola1,5 and T Metz-Estrella5,7
1Department of Marine Sciences, University of Puerto Rico, Road 304, La Parguera, PR 00667; 2Department of Environmental
Sciences, University of Puerto Rico, 17 Ave Universidad STE 1701, San Juan 00925-2537, Puerto Rico; 3Center for Applied Tropical
Ecology and Conservation, University of Puerto Rico, 17 Ave Universidad STE 1701, San Juan 00925-2537, Puerto Rico; 4Sociedad
Ambiente Marino, PO Box 22158, San Juan 00931-2158, Puerto Rico; 5Department of Natural and Environmental Resources, PO
BOX 366147, San Juan Puerto Rico, 00936; 6Nova Southeastern University, 3301 College Avenue, Fort Lauderdale-Davie, Florida
33314; 7Institute for Socio-Ecological Research, Inc, PO Box 3151, Lajas, Puerto Rico 00667
The ongoing deterioration and significant decline in live coral cover and diversity in coral reef communities
worldwide is strongly associated with increasing water temperatures linked to Global Climate Change, aided by
anthropogenic activities (Harvell et al. 2004, 2007, 2009; Weil and Rogers 2011; Maynard et al. 2016; Woodley et al.
2016). In the Wider Caribbean, major community structure and function decline was marked by two region-wide,
concurrent, highly virulent disease epizootics in the early 1980’s. These events almost wiped out two foundational
scleractinian species (Acropora palmata and A. cervicornis), and the keystone sea urchin Diadema antillarum. White
band disease (WBD) affected the acroporids and was caused by a complex of vibrio bacteria (Gil-Agudelo et al. 2006).
The Diadema mass mortality had all the trademark characteristics of a virulent, transmissible, bacterial or viral
infection, but the putative pathogen (pathogens) was never identified (Lessios 2016). Populations of both acroporids
and sea urchins suffered over 95% mortalities throughout the wider Caribbean (Gladfelter 1982; Lessios et al. 1984a,b;
Aronson and Precht 2001; Lessios 2016), followed by a cascade of ecological consequences (significant loss of live
coral cover, primary productivity, spatial complexity, biodiversity and fecundity; loss of ecological functions, increase
in algal cover and biomass, etc.), ending in a shift from coral- to algal-dominated communities and the loss of
ecological services to other tropical marine communities and to human beings (Aronson and Precht 2001; Weil and
Rogers 2011). Several other disease-induced mass mortalities of other cnidarians, as well as of massive, plate and
nodular reef-building genera, have in the last 30 years resulted in additional loss of biomass, diversity and live coral
cover on many Caribbean reefs (Miller et al. 2009; Weil et al. 2009a; Weil and Rogers 2011; Bastidas et al. 2011; Weil
et al. 2017).
More recently, a presumed new “white-plague type” disease, killing large numbers of corals in a short time, was
reported from southeastern Florida in 2014 (Precht et al. 2016; Walton et al. 2018). It followed dredging operations
(2013-2015) in the Port of Miami channel, that resulted in high sedimentation and turbidity near “ground zero” (Miller
et al. 2016), and came after Summer-Fall high thermal anomalies that led to extensive bleaching across the Florida
Reef Tract (Manzello 2015; Walton et al. 2018). Therefore, it is possible that the pathogen(s) could have been released
from sediment disturbance, or that pathogen virulence and/or host susceptibility were affected by the high
temperatures, or both. Often new disease outbreaks occur following a change in host-parasite biological or ecological
relationship, the introduction of a novel pathogen(s) in susceptible host populations, the emergence of newly evolved
pathogens and/or changes in environmental conditions that alter the microbiome/host physiological equilibrium,
fostering increased pathogen virulence, transmissibility and coral mortality (Daszak et al. 2000, 2001; Harvell et al.
2007, 2009; Weil and Rogers 2011; Woodley et al. 2016; Aeby et al. 2019).
This apparently new disease has been called “Stony Coral Tissue Loss Disease” (SCTLD). It is waterborne, highly
transmissible and highly virulent (rapidly kills coral tissues at a rate of 3-4 cm/day), affecting at least 22 foundational,
scleractinian species (generalist), both usually characteristic traits of a novel pathogen (Weil and Rogers 2011).
Furthermore, in Florida and St. Thomas, most diseased coral lesions treated with an antibiotic (amoxicillin) showed
REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
VOLUME 34 NUMBER 1 December 2019 39 | P a g e
signs that disease progression had slowed or even stopped (SCTLD, Florida Keys National Marine Sanctuary 2018; M.
Brandt pers. comm.). These characteristics indicate that a bacterium or a complex of bacteria might be responsible,
although the putative agent(s) have yet to be identified (Meyer et al. 2018; Aeby et al. 2019). Similarities of SCTLD
signs with those of white plague disease type II (WPD-II) (Richardson et al. 1998; Weil and Rogers 2011; Woodley et
al. 2006; SCTLD-Report-Florida Keys National Marine Sanctuary) has produced uncertainty when attempting to
identify the disease in the field. Some differences with WPD include: acute multifocal infections on single colonies,
tissue and mucus sloughing, and more rapid tissue mortality. These symptoms may not, however, be observed in all
colonies, or in a single survey, but only in repetitive surveys of the same colonies.
Both of the above diseases are “generalists”, affecting multiple species, although species susceptibility seems to
vary. Of the 22 species reported with SCTLD signs, most of them are important reef-building species in the Caribbean.
Six species seem to be highly susceptible in Florida (prevalence > 85%) and other localities: Meandrina meandrites,
Colpophyllia natans, Dichocoenia stokesii, Pseudodiploria strigosa, P. clivosa and Dendrogyra cylindrus. Five species
showed prevalence values higher than 45% in Florida (Eusmilia fastigiata, Diploria labyrinthiformis, Montastraea
cavernosa, Stephanocoenia intersepta and Orbicella faveolata) (Meyer et al. 2018; Aeby et al 2019; SCTLD-Report-
Florida Keys National Marine Sanctuary-2018). Four susceptible species (D. cylindrus, O. faveolata, O. franksi and O.
annularis) are listed as threatened under the United States Endangered Species Act. The Orbicella spp. complex,
Montastraea cavernosa and Siderastrea siderea are listed under intermediately susceptible to SCTLD, however,
species susceptibility could vary geographically and temporally as the disease moves through the Caribbean. Corals,
like many other modular cnidarians, do not show many different visible structural/ physiological responses to
diseases, specially, within the white band-white plague syndromes. These syndromes produce a white, clean skeletal
band after the die-off of the tissue, between the normal-looking tissue and the colonizing turf algae. The band width
Table 1. Chronological dispersion of SCTLD in Florida and the northern Caribbean, and
appearance of localized disease outbreak in Puerto Rico
2014 - First reported off the coast of Miami-Dade County, Florida
2015 - Expanded to Biscayne National Park and north to Broward County in Florida
2016 - Continued spreading in Florida, south to the Upper Keys and north to Palm Beach County
2017 - Moved south into the Middle Keys and to the northern latitudinal edge of the Florida Reef Tract
2017 - First reports from the north coast of Jamaica in July 2017
2018 - Reached the Lower Keys in Florida, more reports made from Jamaica, new reports from Mexico,
Belize and St. Maarten
Spring 2018: New observations from the northwest coast of Jamaica
July 2018: First reports from Quintana Roo, Mexico
October 2018: First report in the eastern Caribbean, St. Maarten
2019 - Moved to the southwest end of the Lower Keys, not into the Dry Tortugas National Park. AGRRA
created a map to report and track SCTLD throughout the Caribbean:
January 2019: First reports at Flat Cay, St. Thomas, U.S. Virgin Islands
March 2019: Reported at more locations in St. Maarten; First report in northwest coast of the
Dominican Republic
June 2019: First report from Belize
August 2019: First report from Saint Eustatius
2019 - First reports of a localized disease outbreak off the eastern coast of Puerto Rico
November 2019: Reports of highly virulent tissue loss disease affecting several corals in
Tamarindo Chico reef, Culebra, Puerto Rico. Many colonies with signs similar to those reported
for SCTLD.
https://floridakeys.noaa.gov/coral-disease/disease.html; https://www.agrra.org/coral-disease-outbreak/
REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
40 | P a g e VOLUME 34 NUMBER 1 December 2019
is determined by the balance between how fast the tissue is dying (virulence) and how fast algal turf is colonizing.
Although similar signs have been observed across sites, we cannot assume that the observed signs represent the same
disease produced by the same pathogen (s), as this may not be the case all across the local and/or geographic
distribution of the disease (Sunagawa et al. 2009). Once the pathogen(s) of SCTLD has been determined, diseased
colonies in all the reported localities will have to be tested to confirm whether all cases represent the same disease.
In Florida in 2015-2017 SCTLD spread quickly (at a rate of 7–10 km/month) along the Florida Reef Tract both south
and north of “ground zero”. Since then it has expanded to several northern Caribbean localities (Table 1 -
https://www.agrra.org/coral-disease-outbreak/; https://floridakeys.noaa.gov/coral-disease/disease.html) producing
significant mortalities in populations of the susceptible species in all localities where it has been observed, raising
concerns about the overall impact it could have on the already declining coral reef communities across the region
(Lunz et al. 2017; Florida Keys National Marine Sanctuary 2018; Meyer et al. 2019; Aeby et al. 2019). The dispersion
pattern of SCTLD does not seem to follow directly the direction of local and regional currents from “ground zero”,
since it has shown up in localities in directions against the normal current, and/or at sites thousands of km apart. Since
the disease is waterborne and highly infectious, it would be expected to follow the direction and speed of ocean
currents, however it was reported from Jamaica in July 2017 with no reports from Cuba or the Cayman Islands. It was
reported in Quintana Roo, Mexico in July 2018 and in Belize in June 2019.
The disease was first observed in St. Thomas, USVI, in January 2019, before it was observed in the Dominican
Republic and then in the Turks and Caicos Islands. It quickly spread along the southwest coast of St. Thomas, producing
highly localized mortality of the susceptible species, and has been documented as spreading northeastward as well.
There are no reports of outbreaks of SCTLD from the Bahamas, Cuba or other localities intermediate to the
southernmost areas where it has been reported, leading some researchers to postulate that cruise/cargo ships or
“contaminated” dive equipment might be involved in SCTLD dispersion. There might be other
biological/oceanographic explanations for this discontinuous dispersion pattern. For example, the pathogen (s) might
be part of the normal microbiome of the holobiont, as mutualistic components, or the sediment and substrate, and
become virulent given certain changes in environmental conditions, host susceptibility or both.
Oceanographic current models projected that as a waterborne pathogen SCTLD would reach Puerto Rican waters,
close to Vieques or Culebra, by May-June 2019. It was not however until October 2019 that a few, isolated colonies
of some of the susceptible species were observed with the described SCTLD signs in Culebra. Between March and
August 2019 several colonies of S. siderea were observed with signs of what looked like acute WPD or SCTLD on many
reefs in the east (Ceiba, Humacao, Culebra and Vieques) and west (Cabo Rojo, Guanica and Mona) of Puerto Rico.
Some colonies showed mucus and tissue sloughing, and fast, multifocal, acute rapid loss of live tissue, similar to SCTLD
signs. In what seems a systemic immune response, several colonies became dark purple or just dark (Fig. 2). However,
there were no other diseased species and no signs of an outbreak, which could indicate that this is a different disease
affecting only S. siderea, possibly induced by the thermal anomaly hitting the area this year (Miller et al. 2009; Weil
and Rogers 2011). By November 2019, a minor, localized outbreak of what looked like SCTLD also affected many
colonies of several species at Punta Tamarindo Chico reef (18°18.578’N - 65°19.040’W) on Culebra Island, off the
eastern coast of Puerto Rico (Fig. 1.). Curiously, colonies with typical multifocal signs of SCTLD were not observed in
November 2019 at any of the 29 random sites surveyed in Culebra for NOAA’s National Coral Reef Monitoring Program
(NCRMP). So far, Tamarindo Chico reef is the only locality reported to have characteristic signs of SCTLD in Puerto
Rico.
Water temperatures have been above average this year for the northeastern Caribbean, reaching seven Degree
Heating Weeks (DHW) (local measurements and NOAA) by November 2019. In September 2019, this thermal anomaly
induced yet another extensive bleaching event that expanded all over Puerto Rico’s shallow and upper-mesophotic
coral communities by November. Bleached corals make it difficult to distinguish the signs of SCTLD or WPD. Bleaching
prevalence measured around Culebra during the NCRMP surveys in November 2019 varied between 50-62% (partially
to totally bleached), with the most susceptible species in this area being O. annularis, O. faveolata, P. strigosa, D.
labyrinthiformis and S. siderea. During this period hard corals in Culebra were possibly more susceptible to disease
REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
VOLUME 34 NUMBER 1 December 2019 41 | P a g e
due to thermal stress. Per anecdotal evidence, SCTLD lesion progression rates appeared to slow down or even stop
during periods of higher water temperatures and/or coral bleaching in Florida and the USVI; however the relationship
between these factors is still uncertain (Aeby et al. 2019).
Consecutive surveys in Tamarindo
Chico reef showed a significant
increase in disease prevalence in
SCTLD susceptible species, from 4% in
October, to 50% in November, and
74% in December (Hernández-
Delgado and Suleimán-Ramos,
unpublished data). Similar to the
initial stages of SCTLD in St. Thomas,
acute impacts have remained highly
localized across a single reef site,
affecting mostly meandroid species.
Disease prevalence varied across
species: >50% in D. cylindrus, 80% in P.
strigosa, P. clivosa, and D.
labyrinthiformis, and >90% in M.
meandrites and S. siderea, which, as
mentioned above, have shown signs
of disease well before this localized
outbreak. The disease indiscriminately
affected wild and nursery-restored
colonies of D. cylindrus,
Pseudodiploria spp., D.
labyrinthiformis, C. natans, M.
meandrites and S. siderea. It has also
affected colonies of D. cylindrus and E.
fastigiata on adjacent coral farms, but
with a significantly lower prevalence
(<5%). With the collaboration of Nova
Southeastern University, the NGO
Sociedad Ambiente Marino and a
provisional permit provided by the
Puerto Rico Department of Natural
and Environmental Resources (DNER),
preliminary testing with an
experimental treatment of
amoxicillin antibiotic in CoreRX
Base2B yielded promising results,
halting disease progression in 90% of
treated colonies of Pseudodiploria
spp., D. labyrinthiformis, Colpophyllia
natans, D. cylindrus, M. cavernosa and
S. siderea (N = 50 colonies).
One of the most effective responses documented thus far, from Florida and St. Thomas, has been the use of
amoxicillin; but given the characteristics of SCTLD, there is a pressing need to increase significantly the number of
colonies treated per locality (and to text other antibiotics) in order to minimize the risk of infection spreading to other
Figure 1. Photographs of diseased colonies of species susceptible to SCTLD in
Tamarindo Chico, Culebra. Several small colonies of P. strigosa with multi-focal
infected areas (A). Colony of P. clivosa with fast advancing, multi-focal infections
(B). Medium sized P. strigosa with two rapidly advancing diseased areas (C).
Almost 100% mortality in small D. cylindrus (D). Mucus and tissue sloughing in S.
siderea (E). Rapidly advancing white band area in D. labyrinthiformis (F) and M.
cavernosa (G). Small colony of C. natans that is almost 100% dead in a short time
(H).
REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
42 | P a g e VOLUME 34 NUMBER 1 December 2019
reefs. There is also a need to reduce,
if possible, the numbers of
recreational visits (i.e., SCUBA diving,
snorkeling, kayaking) to the affected
sites and to implement strict
equipment disinfection protocols and
also initiate outreach activities to
inform and educate the stakeholders
and any visitors. The DNER has been
participating in the USVI-SCTLD
status monthly calls even before
suspicious colonies were identified in
Puerto Rico. With the support of the
DNER, SCTLD education and outreach
materials generated by other
jurisdictions have been translated
into Spanish and shared with relevant
audiences to increase awareness and
promote SCTLD prevention. In
addition, theoretical and practical
training on SCTLD was held by Sea
Grant FL, Sea Grant PR and the DNER
for coral reef experts, dive shops,
fishermen and other stakeholders.
Direct communication was also
established with tourist operators in
Vieques, where SCTLD was expected
to show-up first, following UVI
projections.
Several other Federal and local
government agencies, Institutions
and NGOs [the National Oceanic and
Atmospheric Administration (NOAA),
the University of Puerto Rico in San
Juan, Sociedad Ambiente Marino
(SAM), HJR Reefscaping, Coastal
Survey Solutions, and the
Department of Marine Sciences
(DMS) of University of Puerto Rico,
Mayaguez] have also collaborated to
educate, prevent, prepare, and
respond to the threat of SCTLD on Puerto Rico’s coral reefs. By staying in contact with key stakeholders in both Florida
and the USVI, these institutions have had the benefit of learning from other jurisdictions’ experiences to develop
response plans, protocols and “rapid response teams” (RRT) for Puerto Rico. The RRT are trained on how to identify
the disease and differentiate it from other coral diseases and bleaching, how to treat diseased colonies, and how to
decontaminate diving equipment after dives in disease-impacted areas, among other response activities. Such training
and preparedness is crucial given the threat of SCTLD to coral reefs in Puerto Rico. Researchers are exploring other
ways to control the disease.
Figure 2. Diseased colonies of S. siderea around Puerto Rico, some before the
outbreak of the possible SCTLD outbreak was reported in Tamarindo Chico,
Culebra. Two large diseased colonies with multifocal infected areas and evidence
of fast mortality in Guaniquilla, west coast of Puerto Rico (A, B). Mucus and tissue
sloughing in diseased S. siderea (C, D). Colony of S. siderea with multiple signs of
dark spots disease and multi-focal infections of a white-plague type that could be
SCTLD (E). A recently fast killed colony of M. meandrites, with skeletal structure
covered by sediment and signs of some turf colonization (F).
REEF ENCOUNTER
The News Magazine of the International Coral Reef Society
Reef Currents: SCTLD in the Caribbean – implications for Puerto Rico
VOLUME 34 NUMBER 1 December 2019 43 | P a g e
Acknowledgements
We would like to thank our colleagues Karen Neely (Nova Southeastern University), Maurizio Martinelli (Sea Grant FL), Dana Wusinich-Mendez
(NOAA) and the Florida team; Dr. Marilyn Brandt (University of the Virgin Islands), Leslie Henderson (NOAA) and the USVI team; and Dr. Judith
Lang (AGRRA) for all of their continued support, advice, and for providing us with treatment materials. Ernesto Diaz and the Puerto Rico Dept.
of Natural Resources (DNER), Sociedad Ambiente Marino, Kimberly Edwards (NOAA) and Coastal Survey Solutions (NCRMP), Dept. of Marine
Sciences, UPRM, and NSF (Grant # 2000863) provided logistical and partial financial support.
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