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REPORT
Association of butterflyfishes and stony coral tissue loss disease
in the Florida Keys
Kara R. Noonan
1
•Michael J. Childress
1
Received: 3 January 2020 / Accepted: 23 July 2020 / Published online: 1 August 2020
ÓThe Author(s) 2020
Abstract Since 2014, stony coral tissue loss disease
(SCTLD) has rapidly spread throughout the Florida reef
tract infecting and killing dozens of coral species. Previous
studies have found that corallivorous fishes, such as but-
terflyfishes, are positively correlated with coral disease
prevalence at both local and regional scales. This study
investigates the association of SCTLD infection and but-
terflyfish abundance and behaviors on ten reefs in the
middle Florida Keys. Divers conducted video surveys of
reef fish abundance and disease prevalence in June 2017,
2018, and 2019; before, during, and after the outbreak of
SCTLD infections. SCTLD prevalence increased from
3.2% in 2017 to 36.9% in 2018 and back to 2.7% in 2019.
Butterflyfish abundances also showed a similar pattern with
a twofold increase in abundance in 2018 over abundances
in 2017 and 2019. To better understand the association of
individual species of butterflyfishes and diseased corals, 60
coral colonies (20 healthy, 20 diseased, 20 recently dead)
were tagged and monitored for butterflyfish activity using
both diver-based AGGRA fish counts and 1-h time-lapse
videophotography collected in the summers of 2018 and
2019. All reef fishes were more abundant on corals with
larger surface areas of live tissue, but only the foureye
butterflyfish preferred corals with larger surface areas of
diseased tissues. Estimates of association indicate that
foureye butterflyfish were found significantly more on
diseased corals than either healthy or recently dead corals
when compared with the other species of butterflyfishes.
Foureye butterflyfish were observed to feed directly on the
SCTLD line of infection, while other butterflyfish were not.
Furthermore, association of foureye butterflyfish with par-
ticular diseased corals decreased from 2018 to 2019 as the
SCTLD infections disappeared. Our findings suggest that
foureye butterflyfish recruit to and feed on SCTLD-infected
corals which may influence the progression and/or trans-
mission of this insidious coral disease.
Keywords Coral disease Reef fish Butterflyfish
Feeding preference
Introduction
Over the past 40 years Caribbean coral reefs have experi-
enced a dramatic decline in live coral cover attributed
largely to increases in disease (Harvell et al. 2007; Schutte
et al. 2010). Warmer sea surface temperatures along with
increased nutrient pollution have contributed to a prolif-
eration of coral diseases (Harvell et al. 2002; Rosenberg
and Ben-Haim 2002; Selig et al. 2006; Bruno et al. 2007)
and are expected to increase in the future (Maynard et al.
2015). However, evidence suggests that mortality due to
coral disease is strongly influenced by the degree of coral
stress (Lesser et al. 2007; Mueller and van Woesik 2012)
and holobiont composition (Bourne et al. 2009; Randall
et al. 2014; Mera and Bourne 2018) leaving us much to
learn about coral disease transmission.
Coral disease can be transmitted by direct contact,
water-borne, or vector-borne transmission (Shore and
Topic Editor Morgan S. Pratchett
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s00338-020-01986-8) contains sup-
plementary material, which is available to authorized users.
&Kara R. Noonan
noonan2@g.clemson.edu
1
Department of Biological Sciences, Clemson University,
Clemson, SC 26934, USA
123
Coral Reefs (2020) 39:1581–1590
https://doi.org/10.1007/s00338-020-01986-8
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
Caldwell 2019), and there is growing evidence that coral-
livorous fishes may influence coral disease (Rice et al.
2019). Butterflyfish (family Chaetodontidae) abundance
has been found to be positively correlated with coral dis-
ease prevalence at both local and regional scales (Ray-
mundo et al. 2009). However, laboratory studies have
found conflicting results regarding the impact of butter-
flyfish foraging on coral disease transmission, finding both
evidence for (Aeby and Santavy 2006) and against (Nicolet
et al. 2018) butterflyfish as disease vectors. What is clear is
that some butterflyfish are attracted to and preferentially
feed on diseased coral tissues which alters the progression
of coral disease (Aeby 2007; Cole et al. 2009; Chong-Seng
et al. 2011).
Recently, a new and previously undescribed coral dis-
ease began spreading throughout the Florida reef tract
(Precht et al. 2016). Stony coral tissue loss disease
(SCTLD), initially reported in 2014 near Virginia Key, has
spread north and south along the Florida reef tract affecting
more than 20 species of scleractinian (stony) corals (Lunz
et al. 2017; Florida Keys National Marine Sanctuary 2018).
The disease first appears as lesions of sloughed tissue
which can rapidly spread and kill an entire coral colony in
a matter of weeks (Precht et al. 2016; Muller et al. 2018;
Walton et al. 2018). Characterization of the microbial
communities of lesions and response to antibiotic treat-
ments suggest that SCTLD is associated with bacterial
clades commonly found associated with other coral dis-
eases, although an exact putative agent is not yet known
(Aeby et al. 2019; Meyer et al. 2019).
In the summer of 2017, we first observed the initiation
of SCTLD infections on our ten reefs in the middle Keys
where we were conducting reef fish surveys. Divers noted
an increased frequency of butterflyfishes associated with
diseased coral heads. Here, we report our observations of
how changes in SCTLD prevalence are related to changes
in butterflyfish abundance, distribution, and behaviors
before, during, and after this coral disease outbreak. Our
hypotheses were that butterflyfish abundance would
increase with increasing SCTLD prevalence, butterflyfish
would preferentially feed on diseased corals over healthy
corals, and butterflyfish activity would influence SCTLD
disease progression.
Methods
Site selection and substrate census
Our research was conducted in the middle Keys of the
Florida Keys National Marine Sanctuary on ten reefs that
were selected from a set of 36 reefs previously studied
(Smith et al. 2018; Smith et al. 2019). These reefs, 4
inshore (10–20 m depth, \4 km from shore) and 6 off-
shore (15–25 m depth, [6 km offshore), were selected
because of their variation in structural complexity and
percent of live coral cover (Fig. 1). Each reef was censused
in the early summer (June–July) in 2017, 2018, and 2019.
Reef census area was defined by a permanent 50 m transect
line that ran parallel with the primary reef axis and four
perpendicular 30 m transects crossing the main transect at
10, 20, 30, and 40 m. Thus, the entire reef census area was
50 930 m. Substrate cover was estimated from twelve
digital photographs (50 950 cm) transect at 10-m intervals
and calculated using 25 random points per photograph
generated by Coral Point Count with Excel extensions (see
Smith et al. 2018 for descriptions of cover types).
Reef fish abundances and behaviors
To assess reef fish abundances across the entire reef, four
videos along a 30 92 m belt were taken on each of the
secondary transects. A diver holding a PVC camera frame
with two forward facing GoPro cameras attached at heights
of 30 cm and 100 cm from the substrate swam the entire
length of each transect at a pace such that each video was
3–4 min in length. The videos were then analyzed for reef
fish species identification and abundance which were
summed for all four transects. The dual cameras allowed
for identification of hard to see individuals or those moving
quickly across the field of view. For this study, the reef
fishes were classified into five functional groups, including
corallivores (five species of Chaetodon butterflyfishes),
herbivores, omnivores, grunts, and predators.
In 2018, 60 coral colonies (6 per reef) either healthy
(n= 20), actively infected with SCTLD (n= 20), or
recently dead (n= 20) were marked with numbered tags,
measured (max height, length, width) and photographed on
five sides. They included 5 common boulder coral species
(Colpophyllia natans, Montastraea cavernosa, Orbicella
faveolata, Porites astreoides, Siderastrea siderea). Each
photograph was analyzed using ImageJ software to esti-
mate the percent of live, diseased, and dead coral tissue.
Percent cover was converted into surface area by estimat-
ing total coral surface area using the surface area formula
for a half-sphere (2 pr
2
) with a radius estimated as the
average of (max height, length, width). In 2019, all 60 coral
colonies were again photographed on five sides and re-
analyzed for changes in tissue percent cover and surface
area. Percent morbidity was calculated using a percent
change equation between 2018 and 2019 tissue percent
cover calculations. Mortality was noted when there was no
longer any live or diseased tissue cover observed in 2019.
In the summers of 2018 and 2019, each coral head was
monitored for fish abundance using diver-based visual
AGRRA surveys. All fish observed during a five-minute
1582 Coral Reefs (2020) 39:1581–1590
123
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observation of a 5-m cylinder centered on each coral head
were counted and sorted into the five functional groups
described above to estimate local abundance directly
related to the status of each coral head (healthy, diseased,
recently dead). Abundance data were reported as numbers
per five minutes.
To observe fish behaviors in the absence of divers, a
single GoPro camera set for time-lapse images taken at
1-min intervals for 60 min was deployed 50 cm above the
substrate and 150 cm from the center of the coral head.
Each photograph of each coral head was scored for the
presence and behaviors of all species of butterflyfishes and
was categorized into five categories (absent, present but not
over coral head, present over coral head but not feeding,
present and feeding on coral head but not on diseased tis-
sue, present and feeding on diseased tissue). Foureye but-
terflyfish (Chaetodon capistratus) were the most abundant
of all the butterflyfishes and were the only species observed
to feed on diseased coral tissue. The three other species of
butterflyfish (spotfin—C. ocellatus, banded—C. striatus,
reef—C. sedentarius) were much less common and never
observed to feed on diseased tissue. We summed our
observations of these three species for the purpose of sta-
tistical comparison with the more common C. capistratus.
Percent occurrence was calculated as the number of images
within each category over the total number of images taken
for that coral head. Since occurrence may be strongly
influenced by multiple images of the same individual, we
consider it a measure of coral use rather than an estimate of
fish abundance.
Stony coral disease census
Disease prevalence in 2017, 2018, and 2019 was estimated
from a 2 950 m video transect taken along the main
transect on each of our ten reefs. A coral was noted as
‘‘diseased’’ when acute to subacute tissue loss was
observed with indistinct bands of pale tissue (diseased
Fig. 1 Map of our ten reefs surveyed in the middle Florida Keys.
Black circles indicate nearshore reefs, and gray circles indicate
offshore reefs. Inset map of the Florida Keys National Marine
Sanctuary indicates the spread of stony coral tissue loss disease
(SCTLD) from Miami in 2014 to Key West in 2019. SCTLD
infections on our reefs were just beginning in June 2017 and were
gone by June 2019
Coral Reefs (2020) 39:1581–1590 1583
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tissue) progressing to normal pigmentation (healthy tissue).
Each video was analyzed for evidence of disease corals
relative to the total number of live corals along the tran-
sect. Coral species that were surveyed include Acropora
spp., Agaricia spp., Colpophyllia spp., Diploria spp., Di-
chocoenia spp., Montastraea spp., Orbicella spp., Porites
spp., and Siderastrea spp.
Statistical analyses
Reef-wide fish counts and disease prevalence by year were
analyzed by two single-factor ANOVA with Tukey’s post
hoc comparisons. Fish abundances and fish occurrences
(square-root transformed to meet assumptions of homo-
geneity of variances) for each type of coral head were
analyzed by single-factor ANOVA with Tukey’s post hoc
comparisons. Fish abundances and coral characteristics
(coral species, colony length, width, height, surface area,
and percent live/dead tissue) were estimated using two-
factor ANOVA and Pearson’s correlations with avalues
adjusted for multiple comparisons by a sequential Bon-
ferroni correction.
Results
Across our ten sites, stony coral tissue loss disease infec-
tions in 2017 ranged from 0.0 to 8.0% with an average
prevalence of 3.2%. In 2018, SCTLD prevalence peaked
with percentages ranging from 23.1 to 45.8% with an
average prevalence of 36.9% (Table 1). Then, in 2019,
disease occurrence diminished with infections ranging
from 0.0 to 5.2% with an average prevalence of 2.7%.
Percent disease in 2018 was unrelated to region (t= 0.937,
df = 2.28, p= 0.3976) or the amount of live hard coral
cover before the outbreak (F= 0.964, df = 2. 26,
p= 0.3589). Disease prevalence was significantly higher in
2018 than either 2017 or 2019 (F = 150.9, df = 2.26,
p\0.0001).
Of the 20 diseased corals, we tagged and monitored
beginning in the summer of 2018, 19 survived and showed
no signs of active SCTLD infection in 2019 (Table S1).
Percent morbidity ranged from -0.5471% (increase in
live tissue cover) to 100% with a mean coral morbidity of
35.1% (Table S1). However, we did not see a regional
(F= 0.812, df = 1.27, p= 0.3729), site (F= 1.173, df =
9.19, p= 0.3455), or species effect on morbidity
(F= 1.322, df = 4.24, p= 0.2803).
Total reef fish abundance did not change over this three-
year period (F= 3.22, df = 2.26, p= 0.0561). However,
the abundance of corallivores (family Chaetodontidae) did
show a higher abundance in 2018 (F= 2.92, df = 2.26,
p= 0.0716) (Fig. 2) and was positively related to disease
prevalence (F= 3.767, df = 1.27, p= 0.0627), although
not significantly. Total reef fish abundance around focal
corals did not change from 2018 to 2019 and was unrelated
to coral status (Table 2). However, foureye butterflyfish
abundance was significantly higher in 2018 than in 2019,
regardless of coral status (F= 11.07, df = 1.27,
p= 0.0012) (Table 2).
The characteristics of coral heads with the largest
impact on fish abundances were height and surface area.
All reef fish were more abundant around tall coral heads
and those with the greatest surface area of live tissue
(Table 3). This also held true for foureye butterflyfish, but
not for the other species of butterflyfishes. However,
foureye butterflyfish abundance was better predicted by the
surface area of diseased tissue than the surface area of live
tissue. This suggests that foureye butterflyfish were
attracted to tall, diseased coral heads. The species of coral
had no influence on total reef fish abundance (F= 1.48,
df = 4.35, p= 0.1987) or foureye butterflyfish abundance
(F= 0.61, df = 4.35, p= 0.6863).
During 2018, when corals had active SCTLD infections,
the diseased and healthy corals had significantly higher
occurrences of foureye butterflyfish than did dead corals
(F= 3.31, df = 1.19, p= 0.0434) (Fig. 3a), but other spe-
cies of butterflyfish did not show any difference in occur-
rences due to coral status (F= 0.25, df = 1.19,
Table 1 Percent coral cover
(averaged among quarterly
censuses), disease (2018 disease
prevalence surveys), and pre-
and post-disease fish abundance
data for nine reef sites across the
middle Florida Keys (Long Key
Ledge A and B were combined
for the purpose of analyses)
Site Region %Coral cover %Disease Pre-disease fish count Post-disease fish count
E Turtle Patch Nearshore 21.35 43.40 264 456
Coral Gardens Nearshore 16.31 23.08 1238 529
Karas Reef Nearshore 12.75 27.27 NA 572
Turtle Shoals Nearshore 10.60 39.73 157 198
Elbow Offshore 8.25 45.83 508 569
11’ Mound Offshore 6.63 41.38 467 290
Stag Party Offshore 4.62 37.50 97 185
LK Ledge Offshore 2.02 32.05 875 848
Out Spa Offshore 1.96 41.67 277 184
1584 Coral Reefs (2020) 39:1581–1590
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p= 0.7728). This association was no longer observed when
the same corals no longer had active infections of SCTLD
in 2019 (F= 0.03, df = 1.19, p= 0.9661) (Fig. 3b).
In 2018, during the active infections of SCTLD, foureye
butterflyfish were observed feeding more often on diseased
corals than on healthy corals (F= 2.67, df = 1.19,
p= 0.0980) (Fig. 4), but these differences were no longer
present in 2019 after SCTLD infections had cleared
(F= 0.80, df = 1.19, p= 0.4718). However, foureye but-
terflyfish feeding intensity was unrelated to the percent
decrease in the surface area of healthy coral tissue between
2018 and 2019 (F= 0.17, df = 1.19, p= 0.9864). When
analyzing healthy and diseased coral heads, foureye
Fig. 2 a Stony coral tissue loss
disease infected more than 35%
of all corals in June 2018
including Colpophyllia natans,
Orbicella faveolata,
Montastraea cavernosa, and
Siderastrea siderea.bDisease
abundance (indicated in black)
was significantly higher
(F
2,26
= 150.9, P\0.0001) in
2018 than in 2017 and 2019.
Butterflyfish fish abundance
(Chaetodon spp.) (indicated in
orange) was also higher
although not significantly
(F
2,26
= 2.92, P= 0.0716) in
2018 than in 2017 and 2019
Table 2 Analysis of variance of year (2018 vs 2019) and coral status
(healthy, disease/healed, dead) on fish abundance (n= 120)
Fish group Source df FP
All reef fish Year 1.27 0.23 0.6297
Coral status 2.26 1.93 0.1484
Other butterflyfish Year 1.27 3.32 0.0749
Coral status 2.26 0.25 0.7749
Foureye butterflyfish Year 1.27 11.07 0.0012
Coral status 2.26 0.10 0.9015
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butterflyfish have no relationship to percent morbidity
(F= 0.51, df = 1.19, p= 0.4795). Furthermore, 19 of 20
diseased corals survived their SCTLD infection regardless
of the intensity of foureye butterflyfish feeding.
Discussion
Stony coral tissue loss disease reached the reefs in the
middle Florida Keys in 2017, peaking in 2018, and
declining significantly in 2019 (Florida Keys National
Marine Sanctuary 2018). In 2018, location-specific infec-
tion rates across all available corals ranged from 23.1
percent to 45.8 percent with no correlation between percent
coral cover and disease prevalence. We did not find a
significant difference in coral infection rates between our
inshore and offshore reefs as previously reported (Rippe
et al. 2019). However, these estimates of disease preva-
lence may be lower than actual disease prevalence due to
limitations imposed by estimating disease from video
transects.
Corallivores (Chaetodontids) also showed a peak abun-
dance in 2018, twice as high than in 2017 or 2019. This
pattern was not seen in our other reef fish functional groups
and thus was not a general phenomenon affecting all reef
fish species. Across all censuses, there was a positive, but
not significant, relationship between corallivore abundance
and percent disease prevalence. A previous study found no
correlation between butterflyfish abundance and the
prevalence of coral disease but a positive response of
corallivorous drupellids in response to disease (Raj et al.
2016). What our study suggests is that response to
increasing disease prevalence is very rapid and unlikely
due to increases in butterflyfish recruitment or growth rates.
More likely butterflyfish are locally attracted to large dis-
eased coral heads from surrounding areas with few dis-
eased individuals. Whether or not butterflyfish track the
movement of the disease front long distances is not yet
known, but there are some studies that other corallivores,
e.g., drupellids, track resources using chemical cues
released by stressed corals (Stambler 2010; Tsang and Ang
2015; Kaullysing et al. 2016).
Diver-based fish surveys conducted in 2018 and 2019
found that butterflyfish were more abundant around large
coral heads in 2018 than in 2019, but other reef fishes were
not. This was mostly due to increases in one species of
butterflyfish, C. capistratus. Reef fish abundance was
positively correlated with coral height and area of live
tissue. However, for C. capistratus abundance was most
strongly correlated with area of diseased tissue and was
unrelated to coral species. The other three species of but-
terflyfishes show no correlations with abundance, coral
species, or size. Previous studies have found similar asso-
ciations of reef fishes with large coral heads and high live
tissue area due to the biological and physical structure that
coral provides (Bell and Galzin 1984; Roberts and Ormond
1987; Gratwicke and Speight 2005).
Time-lapse videophotography in the absence of divers
found no pattern in association for reef, spotfin, and banded
butterflyfishes which showed similar frequencies during
and after the disease outbreak across all coral types.
However, foureye butterflyfish associated more with dis-
eased corals than healthy or dead corals in 2018 and dis-
appeared one year later when the same corals no longer had
active SCTLD infections. While time-lapse videophotog-
raphy may overestimate occurrences due to multiple
observations of the same individual fish, they still corrob-
orate our diver-based visual surveys indicating higher
association by foureye butterflyfish. Foureye butterflyfish
were observed feeding significantly more often on diseased
corals than on healthy or dead corals with photographs
indicating frequent feeding on the active disease line.
These differences disappeared the following year when the
infected coral heads no longer had active SCTLD. This
parallels previous results where butterflyfishes were
attracted to diseased corals in the field and fed on disease
lesions (Aeby and Santavy 2006; Aeby 2007; Raymundo
et al. 2009; Chong-Seng et al. 2011).
Table 3 Correlations of fish
abundance to coral
characteristics (n= 120)
Coral characteristic All reef fish Other butterflyfish
a
Foureye butterflyfish
rP r P r P
Coral height 0.227 0.0128* 0.087 0.3437 0.270 0.0030*
Surface area (total) 0.188 0.0406 0.032 0.7228 0.268 0.0032*
Surface area (live) 0.317 0.0004*-0.011 0.9003 0.130 0.1582
Surface area (diseased) 0.067 0.4646 -0.046 0.6134 0.314 0.0005*
Surface area (dead) 0.044 0.6279 0.046 0.6167 0.249 0.0063*
a
Other butterflyfish includes all spotfin, banded, and reef butterflyfishes
Bold significant at a= 0.05
*Significant at global a= 0.05 by sequential Bonferroni correction
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With the attraction of corallivorous fishes to sloughing
coral tissue, the increase in foureye butterflyfish to diseased
colonies follows previously observed trends (Aeby and
Santavy 2006; Raymundo et al. 2009; Chong-Seng et al.
2011). Additionally, C. capistratus is a home-ranging
(Neudecker and Lobel 1982; Gore 1983), generalist
corallivore (Gore 1984; Cole et al. 2008; Bellwood et al.
2009), making the repeated opportunistic feedings on dis-
ease lesions expected. The lack of response of the other
three species to diseased tissue may be due to differences in
dietary habits. C. sedentarius and C. striatus are considered
benthic invertebrate feeders (Bellwood et al. 2009), which
Fig. 3 Occurrence of butterflyfishes in time-lapse photographs of
focal hard corals either dead, diseased, or healthy. Number of
photographs per hour with foureye butterflyfish (orange) or reef,
banded, spotfin butterflyfish (blue) ain 2018 and bin 2019. Note that
all disease corals from 2018 were healed by 2019 and showed no
active stony coral tissue loss disease. Occurrence of foureye
butterflyfish on diseased corals in 2018 was significantly higher
(P\0.05) than on dead corals. Other butterflyfish occurrence was
unrelated to hard coral status for both years
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explains why there was not an influx in their abundances,
although spotfin butterflyfish were observed feeding on the
disease lesions across our ten sites. C. ocellatus has similar
dietary needs as C. capistratus, so it is unknown why they
did not have a similar trend to the foureye butterflyfish.
With direct feeding on coral lesions, some studies have
found that active feeding leaves corals more vulnerable to
infection (Aeby and Santavy 2006; Raymundo et al. 2009),
while other studies found that the active feeding and
removal of infected tissue allowed corals an opportunity to
recover from disease (Aeby 1983; Cole et al. 2009). We
found no relationship between the abundance or frequency
of occurrence of butterflyfishes and coral morbidity or
mortality from SCTLD, but of our twenty infected corals,
nineteen survived the infection. This may support previous
findings that the butterflyfish may be halting or at least
slowing the progress of disease by consuming the slough-
ing tissue (Aeby 1983; Cole et al. 2009). Previous work on
butterflyfish transmission in black-band disease found no
evidence of increased or decreased transmission (Nicolet
et al. 2018).
It is unknown whether the observed increase in butter-
flyfish feeding influenced the rate of disease spread to
uninfected corals. If butterflyfish were facilitating the
spread of SCTLD, we would expect to find faster rates of
new infection in areas with higher butterflyfish abundance,
but if coral disease is increasing butterflyfish recruitment
after initial infection, disease spread would not necessarily
be related to butterflyfish abundance. More frequent
observations would be needed to measure this relationship
Fig. 4 a Evidence of feeding
on stony coral tissue loss
disease by foureye butterflyfish.
bPercent of occurrences where
feeding was observed in 2018
and 2019 by foureye
butterflyfish on dead, diseased,
and healthy corals. More than
50% of all observations of
foureye butterflyfish on diseased
corals in 2018 indicated
feeding. By 2019, these same
corals no longer had active
SCTLD infections and feeding
was no more frequent than
observed on healthy corals
1588 Coral Reefs (2020) 39:1581–1590
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with appropriate butterflyfish-free corals as controls. Future
research needs to investigate the potential influences that
these butterflyfish may have on disease transmission, with
specific focus on the surface area of live coral tissue.
This study is the first to examine the relationship
between butterflyfish and stony coral tissue loss disease in
the Caribbean. Given the rate of SCTLD spread and the
lethality of this pathogen, understanding the role of foureye
butterflyfish in its transmission should be a top priority in
the efforts to protect corals and manage this disease out-
break (Shore and Caldwell 2019).
Acknowledgements This research was made possible by the Florida
Keys National Marine Sanctuary (Permit # FKNMS-2017-032 and
FKNMS-2018-119). Funding for this project was provided by
Clemson University’s Creative Inquiry Initiative, the International
Women’s Fishing Association, The Explorer’s Club Mamont Scholar,
American Museum of Natural History Lerner-Gray Memorial Fund,
and Clemson University. We thank Kylie Smith, Randi Sims, Sydney
Whitaker, Isaac Ingrum, Reanna Jeanes, Riley Garvey, Rachel
Radick, and Kristiaan Matthee for assistance in the field data col-
lection. Thomas Fair, Kelsey Sox, Riley Garvey, Kristiaan Matthee,
Emma Crowfoot, and Rachel Radick assisted with the data processing
and analysis.
Open Access This article is licensed under a Creative Commons
Attribution 4.0 International License, which permits use, sharing,
adaptation, distribution and reproduction in any medium or format, as
long as you give appropriate credit to the original author(s) and the
source, provide a link to the Creative Commons licence, and indicate
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use is not permitted by statutory regulation or exceeds the permitted
use, you will need to obtain permission directly from the copyright
holder. To view a copy of this licence, visit http://creativecommons.
org/licenses/by/4.0/.
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