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The recent collapse of a rapid phase-shift reversal on a Jamaican north coast coral reef after the 2005 bleaching event

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The community structure of most Caribbean reefs has changed dramatically since the 1980s. Invoking a chemistry analogy, in 1994 Hughes termed the change a "phase shift" to describe the change from a coral dominated habitat to one dominated by macroalga on the north coast of Jamaica over a period of 17 years. The loss of live coral cover is exemplified by the demise of Acropora spp. in Discovery Bay, Jamaica. Dense, monospecific high relief thickets of Acropora palmata (elkhorn coral) and A. cervicornis (staghorn coral) were characteristic of shallow and intermediate depth coral communities in the Caribbean prior to the late 1970s. In the early 1980s, A. cervicornis live coral cover was >21% at several sites around Discovery Bay, by 1987 it had declined to less than 1%. No large population of Acropora reestablished in the vicinity of Discovery Bay for nearly two decades. In 1995, A. cervicornis cover at Dairy Bull Reef (DBR) was only 0.6%. By 2004, A. cervicornis cover had increased to about 10.5% . In 2005, A. cervicornis cover further increased to 44%. This increase in A. cervicornis cover was part of the important "phase shift reversal" reported by Idjadi et al. in 2006. An isolated population of A. cervicornis exhibited similar coral cover at East Rio Bueno (ERB). In 2005, both populations of A. cervicornis bleached; however, the two populations responded differently. The population at DBR was decimated by the bleaching event and the surviving remnants were attacked by Coralliophila. At DBR, cover declined to
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Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (Suppl. 1): 149-159, May 2008
The recent collapse of a rapid phase-shift reversal on a Jamaican
north coast coral reef after the 2005 bleaching event
N.J. Quinn
1,2
& B.L. Kojis
1,3
1. Discovery Bay Marine Lab, University of the West Indies, Discovery Bay, St. Ann, Jamaica
2. Present address:, Department of Planning and Natural Resources, St. Croix East End Marine Park, 45 Mars Hill,
Frederiksted, U.S. Virgin Islands 00841 FAX 340-777-8151. norman.quinn@dpnr.gov.vi
3. Present Address: Division of Science and Math, University of the Virgin Islands, #2 Brewers Bay, St. Thomas, U.S.
Virgin Islands 00802
Received 15-VII-2007. Corrected 01-XI-2007. Accepted 13-II-2008.
Abstract: The community structure of most Caribbean reefs has changed dramatically since the 1980s.
Invoking a chemistry analogy, in 1994 Hughes termed the change a “phase shift” to describe the change from a
coral dominated habitat to one dominated by macroalga on the north coast of Jamaica over a period of 17 years.
The loss of live coral cover is exemplified by the demise of Acropora spp. in Discovery Bay, Jamaica. Dense,
monospecific high relief thickets of Acropora palmata (elkhorn coral) and A. cervicornis (staghorn coral) were
characteristic of shallow and intermediate depth coral communities in the Caribbean prior to the late 1970s. In
the early 1980s, A. cervicornis live coral cover was >21% at several sites around Discovery Bay, by 1987 it
had declined to less than 1%. No large population of Acropora reestablished in the vicinity of Discovery Bay
for nearly two decades. In 1995, A. cervicornis cover at Dairy Bull Reef (DBR) was only 0.6%. By 2004, A.
cervicornis cover had increased to about 10.5% . In 2005, A. cervicornis cover further increased to 44%. This
increase in A. cervicornis cover was part of the important “phase shift reversal” reported by Idjadi et al. in 2006.
An isolated population of A. cervicornis exhibited similar coral cover at East Rio Bueno (ERB). In 2005, both
populations of A. cervicornis bleached; however, the two populations responded differently. The population at
DBR was decimated by the bleaching event and the surviving remnants were attacked by Coralliophila. At DBR,
cover declined to <0.5% by June 2006. The population at ERB recovered from the bleaching event with little
decrease in cover. Coral recruitment was examined over three spawning periods on DBR. Only two Acropora
spat recruited to settlement tiles – one in 2003 and one in 2005. Acropora recruitment represented only 0.7% of
the total spat recruiting to the tiles during the entire sampling period. Rev. Biol. Trop. 56 (Suppl. 1): 149-159.
Epub 2008 May 30.
Key words: Acropora cervicornis, restoration, recruitment, planulae.
Four decades ago Goreau and Goreau
(1973) documented Caribbean coral reef com-
munity structure on north coast Jamaican
reefs. The importance of their studies became
increasingly recognized as shallow water live
coral cover declined throughout the region and
the acroporids became locally extinct on many
reefs (Knowlton et al. 1990). In fact, their work
was used as a standard by which coral reefs
were measured throughout the Caribbean.
Prior to 1980, Acropora cervicornis was
abundant at the West Fore Reef at Discovery
Bay (Goreau and Wells 1967, Tunnicliffe 1983).
However, in the 1980’s, the cover of A. cervi-
cornis was reduced from 53% + 10% to a negli-
gible presence by Hurricanes Allen (1980) and
Gilbert (1988) (Woodley et al. 1981, Crawford
1995, Woodley 1991). Recovery of A. cervi-
cornis after Hurricane Allen was impeded by
high mortality of surviving fragments caused
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by disease (Knowlton et al. 1981). During the
past 25 years, there has been no wide spread
increase in A. cervicornis due to disease and
predation (Bruckner et al. 1997) and the low
rate of sexual recruitment characteristic of this
species (Sammarco 1980, Tunnicliffe 1981).
This naturally low sexual recruitment rate has
been exacerbated by the decline in the abun-
dance of the reproductive adult population
(Kojis and Quinn 1993, 2001, Quinn and Kojis
2005, 2006a, 2006b).
Hughes (1994) used the term “phase shift”
to describe the change from a coral dominated
to an algal dominated reef community on the
north coast of Jamaica near Discovery Bay. The
recent rapid increase in live coral cover, par-
ticularly in the abundance of A. cervicornis, at
Dairy Bull Reef on the north coast of Jamaica,
suggested that a “phase shift reversal” from
an algal dominated to a coral dominated reef
community was possible (Idjadi et al. 2006).
However, in 2005 a severe bleaching event
impacted reefs on the north coast of Jamaica.
This paper describes the impact of the 2005
bleaching event on the Dairy Bull Reef corals,
with an emphasis on the impact on A. cervi-
cornis, and compares these results to those at
the nearby east Rio Bueno reef.
MATERIALS AND METHODS
Water Temperature: A Hugrun Seamon
brand underwater temperature recorder with an
absolute accuracy of + 0.05 ºC was deployed at a
depth of 8 m at Dairy Bull Reef (DBR) (18º28.04’
N; 77º23.10’ W), < 2 km from the West Fore Reef
(WFR) site (18
o
28.17’ N; 77
o
24.49 W), from
21 December 2000 to 15 May 2002. It recorded
11,707 hourly subsurface seawater temperature
(S3T) observations. The UTR was placed adja-
cent to a rich coral reef community ~ 5 cm above
the seabed to allow for a good flow around
the recorder. The recorder was moved to the
WFR (depth: 11 m) and recorded 27,735 hourly
S3T observations from 2 November 2002 to
31 December 2005. The sites were considered
similar and the data pooled. Gaps in both data sets
were the result of equipment failure.
Coral Recruitment: Coral recruitment
arrays were constructed by attaching four 208
cm
2
unglazed terracotta tiles to a PVC array.
The tiles were smooth on one side and had 12
ridges on the other side. Two tiles were ori-
ented horizontally and two vertically on each
array. Paired arrays were installed at DBR at
9 m depth with the tiles ~ 0.8 m above the sub-
strate in early April 2003. They were replaced
eight months later in December 2003 (“summer
2003” sampling period), ten months later in late
October 2004 (“winter / summer 2004”), nine
months later in July 2005 (“winter 2005”) and
finally collected four months later in November
2005 (“summer 2005”). After removal from
the site, the tiles were fixed in formalin and
bleached. Scleractinian corals were counted
(standardized to number recruits m
-2
) and iden-
tified to family where possible using a binocu-
lar microscope.
Coral Surveys and Bleaching: Structured
quantitative surveys were conducted between
September 2005 and June 2006 along 20 m
transect lines parallel to the depth contours at
the 7 - 10 m depth range at DBR (n=10) and
at East Rio Bueno (ERB) (n=8) (18
o
28.47’
N; 77
o
25.50’W). The structured quantitative
surveys consisted of recording data at 0.5 m
intervals under the transect line in accordance
with standard Reef Check protocol (Hodgson
and Liebeler 2002). Additionally, when corals
were encountered, the species was identified
and the following attributes recorded: presence
or absence of bleaching, percent of colony
bleached, and evidence of recent presence of
predators. If the coral was recently killed, this
was recorded this as well. The percentage of
coral colonies bleached and the average per-
centage of bleaching 1 m on each side of each
transect line was estimated. Qualitative assess-
ment of overall bleaching was carried out from
September 2005 to June 2006 during 60-75
min dives (n=25) by estimating the percentage
of colonies bleached at depths 2-35 m. Photos
representative of the coral community at each
survey location were taken both along the
Reef Check transects and during the random
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surveys and subsequently used in the analysis.
From the photos the percent bleaching of the
dominant coral present in each photo was esti-
mated. Based on all three data sets, the extent
of bleaching was determined.
RESULTS
Water temperature: The daily subsur-
face seawater temperature (S3T) was higher
from mid May to mid October 2005 than for
this period in each of the years 2001-2004
(Fig. 1). The mean monthly S3T from April to
November 2005 was above the mean monthly
temperature for April to November 2001-2004
(Fig. 2). Mean monthly S3T in 2005 remained
at least 0.69
o
C warmer than the mean for
the previous four years from April through
October (Table 1) with the greatest differ-
ence between the mean monthly S3T in May
(1.05
o
C) and September (0.88
o
C). The warm-
est S3T recorded during the five years of this
study was 30.45
o
C, which occurred at 17:00hr
on 29 September 2005, a couple of weeks after
the initial observation of the onset of bleach-
ing. The period mid May to mid October was
characterized by calm seas and clear to partly
cloudy days (pers. obs.).
Coral Recruitment: Spat from the family
Poritidae were the most common (39.7%) at
Dairy Bull Reef (DBR) followed by uniden-
tifiable spat (25.2%), Agariciidae (23.2%),
Milleporidae (11.3%) and Acroporidae (0.7%).
Of the 302 Scleractinia and Millepora spat
recorded on settlement plates at DBR, two
acroporid recruits were observed, only one of
which was large enough to identify to species.
The identifiable recruit was an Acropora cervi-
cornis 8.5 cm in length. It recruited to a tile that
was deployed for 8.5 months (from 25 Oct 04
to 11 July 05), exhibiting an annualized growth
rate of at least 12 cm year
-1
.
Coral Bleaching: At the beginning of
2005, the scleractinian live coral cover at
DBR was high, creating a structurally com-
plex and visually appealing reef. Of the 56%
total live coral cover, 44% was A. cervi-
cornis. In early September 2005, the onset
of bleaching was first observed in colonies
of the Montastraea annularis complex when
portions of colonies began to pale or turn
white (Fig. 3). By mid September all spe-
cies of scleractinan coral at DBR (except
Dendrogrya cylindrus) showed some signs
of bleaching.
2000 2001 2002 2003 2004 2005
31
30
29
28
27
26
25
Temperature (ºC)
One month
Bleaching rst noticed
J F M A M J J A S O N D
Fig. 1. Mean monthly subsurface sea water temperatures 21 December 2000 31 December 2005 (modified from Quinn
and Kojis 2003a).
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2 5
2 6
2 7
2 8
2 9
3 0
3 1
J F M A M J J A S O N D
2 0 0 0 - 2 0 0 4
2 0 0 5
Temperature (ºC)
Months
Fig. 2. Comparison of 2005 mean monthly subsurface sea water temperature with mean temperatures from 21 December
2000 - 31 December 2004.
Fig. 3. Montastrea annularis at initial stages of bleaching in September 2005 at 9 m at Dairy Bull Reef.
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From October to December 2005, the per-
centage and degree of colonies bleached of each
species varied widely (Table 2a, 2b). However,
more than 50% of colonies of most species
bleached at DBR and ERB with more than 95%
of colonies of A. cervicornis, Agaricia spp.,
Leptoseris cucullata, Meandrina meandrites,
M. annularis, Porites spp., Siderastrea siderea
and Acropora spp. bleaching. Only A. cervi-
cornis exhibited 100% bleaching at both sites
with100% of colonies bleached totally white
(Table 2a, b).
Mortality varied widely between species
and sites. In late December 2005 / January
2006, the highest mortality from bleaching was
exhibited by A. cervicornis at DBR with 90%
mortality (Table 2a). In contrast, A. cervicornis
at ERB reef exhibited only 10% mortality
(Table 2b). A. palmata colonies at West Rio
Bueno recovered from the bleaching, but were
observed being preyed upon by the coral snail,
Coralliophila abbreviata, in February 2006.
Mortality was readily noted because filamen-
tous algae were beginning to cover the skel-
etons of recently killed coral.
In February, 2006, surviving remnants of
A. cervicornis at DBR were observed to be
preyed upon by the Coralliophila abbreviata.
There was also evidence of predation by the
bearded fire worm (Hermodice carunculata).
Predation on A. cervicornis by these two spe-
cies was not observed at ERB.
From February to June 2006, the remain-
ing live tissue on surviving bleached colonies,
including A. cervicornis, began to progressive-
ly regain its natural color. However, by June
2006 at DBR, there was less than <0.5% live
coral cover of A. cervicornis (Fig. 4).
DISCUSSION
Idjadi et al. (2006) described a “phase shift
reversal” on Dairy Bull Reef (DBR) in which
live coral cover increased to 1970 levels and
macroalgal cover decreased to levels recorded
prior to Diadema mortality of 1983-84. They
declared that the benthic community at DBR
in 2004 was reminiscent of the fore reef
TABLE 1
Mean monthly sea water temperatures in
o
C: Years 2001-2004 versus 2005
Year Jan Feb Mar Apr May June July Aug Sept Oct Nov Dec mean
2001-04 27.06 26.74 26.86 27.00 27.45 28.11 28.46 28.78 29.27 29.38 28.76 27.85 27.98
2005 26.42 25.76 26.78 27.77 28.50 29.12 29.15 29.55 30.14 30.09 29.54 28.11 28.41
Difference
o
C -0.64 -0.98 -0.08 0.76 1.05 1.01 0.69 0.76 0.88 0.71 0.78 0.26 0.43
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TABLE 2A
Percentage of colonies bleached and average degree of bleaching for some common scleractinians, gorgonians and
sponges observed from Sept. 2005 Feb. 2006 at Dairy Bull Reef
Species % of colonies
bleached
Degree of bleaching Colony mortality related to bleaching
Scleractinia
Acropora cervicornis
100% totally white 90%
Acropora palmata
100% Totally white Occasional colonies completely dead,
mostly PM50
Agaricia lamarcki
95% totally white PM50
Agaricia agaricites
95% totally white PM50
Colpophyllia natans
66% pale to totally white PM50
Diploria strigosa
66% pale to medium very rare
Dendrogyra cylindrus
50% slight paleness none
Leptoseris cucullata
95% totally white PM50
Madracis decactis
60% pale PM50
Meandrina meandrites
95% severe – mostly white none
Montastraea annularis
complex
99% very pale to totally white Some mortality in smaller colonies,
mostly PM25
Montastraea cavernosa
5% some colonies pale spots none
Porites astreoides
100% slight color change none
Porites porites
99% severe PM50
Siderastrea siderea
95% severe – colonies colored pale
blue
PM75
Siderastrea radians
50% severe – colonies colored pale
blue
PM25
Stephanocoenia intersepta
50% pale none
Gorgonians
Eunicea calculata
5% occasionally pale none
Plexaurella nutans
5% occasionally pale none
Pseudopterogorgia sp
0% no bleaching none
Sponges
No evidence of bleaching 0% no bleaching none
Data blended from surveys described in Materials and Methods.
Legend: Partial mortality in <25% of the colonies, PM25; partial mortality in >25% and <50% of the colonies, PM50; partial
mortality in >50% of the colonies, PM75.
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community structure present at Discovery Bay
prior to Hurricane Allen with dense thickets of
Acropora cervicornis lodged between colonies
of Montastraea annularis and surmised that
colonies of M. annularis provided topographic
complexity that may have facilitated the recov-
ery of the DBR reef. The term “phase shift”
was originally used to describe the overgrowth
of north coast Jamaica coral reefs in 1983 by
algae associated with the mass die off or the sea
urchin, Diadema antillarum (Hughes 1994).
Other work on long term changes of coral reefs
all around Jamaica concluded that the sites that
Hughes studied gradually became eutrophic
in the early 1980s, while coral reefs on other
parts of Jamaica also transitioned from coral
to algal dominated reefs between the 1950s
to the 1990s depending on the increase of the
local human population and influx of sewage
effluents (Goreau 1992).
In 2005, the reefs on the north coast of
Jamaica near Discovery Bay were heavily
impacted by a bleaching event. The summer
of 2005 was unusual in that there were exten-
sive periods of low wind velocity, clear skies,
calm seas and low turbidity. These conditions
TABLE 2B
Percentage of Acropora cervicornis and A. palmata colonies bleached observed from Sept. 2005 Feb. 2006 at Rio Bueno
Reef
Species % of colonies bleached Degree of bleaching Colony mortality related to bleaching
Acropora cervicornis 100% Totally white 10%
Acropora palmata 100% Totally white 15%
Data blended from line transects and random photo surveys.
Fig. 4. Mean percentage cover of: live coral, Acropora spp., and macro algae at Dairy Bull Reef, in 1995, 2003, 2004 (from
Idjadi et al. 2006), 2005 and 2006 this study. Bars represent SE.
0
1 0
2 0
3 0
4 0
5 0
6 0
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
% coral cover
c o ra l A c ropo r a
m a c ro a lga e
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favored localized warming and high penetra-
tion of ultra violet radiation (UVR). It is most
likely that the prolonged, unusually warm
seawater temperature, exceeding >29.3
o
C for
one month, the predicted threshold for induc-
ing bleaching (Goreau et al. 1993), was the
major cause of the 2005 bleaching event. In
2005, 29.3
o
C was first recorded on May 22,
which was the earliest that this temperature
had been observed in the period 2001 – 2005
(Fig. 1) (Quinn and Kojis 2003a). Although
the temperature remained above 29
o
C dur-
ing June and July, a month of consecutive
daily temperatures exceeding 29.3
o
C did not
accrue until September 7. Coral bleaching
was first observed on a dive conducted on
the same date.
After the bleaching event of 2005, live
coral cover at DBR decreased dramatically to
well below the pre “phase shift reversal” lev-
els recorded by Idjadi et al. (2006). Total live
coral cover declined from 56% in 2005 to 11%
in 2006. In fact, the 2006 level was 12% less
than the 23% cover recorded in 1995 by Idjadi
et al. (2006).
Acropora cervicornis was a major contrib-
utor to the increase in live coral cover reported
by Idjadi et al. (2006) at DBR, comprising
nearly 20% of the total live coral cover. In 2005
(this study), total live coral cover was similar
to 2004, while A. cervicornis now comprised
>75% of the live coral cover. Although, the dif-
ferent survey methodology used in this study
may have contributed to the >50% increase
in A. cervicornis abundance recorded in 2005
(Fig. 4), there are other factors that may have
contributed to the decline in the contribution
of other coral species to total live coral cover
and the rapid increase in A. cervicornis. During
2003-2004, black band disease infected many
M. annularis colonies and killed large sec-
tions of many colonies (Quinn, personal obs.).
This event was overshadowed by the massive
increase in A. cervicornis at DBR. With its
high growth rate and branching structure A.
cervicornis was able to rapidly cover large
areas of the reef obscuring smaller species dur-
ing transect surveys and giving the impression
of a dynamic, healthy reef while masking the
decline in the M. annularis population.
Surveys conducted in late June 2006
revealed that <0.5% of the A. cervicornis popu-
lation at DBR was still alive. The high mortal-
ity of A. cervicornis from the 2005 bleaching
event and subsequent predation of the scattered
remnants of live tissue by Coralliophila and
Hermodice explains most of the decline in live
coral cover at DBR, since A. cervicornis pro-
vided the majority of live coral cover.
In contrast, the population of A. cervicornis
at East Rio Bueno (ERB) had much less mor-
tality. Live coral cover declined only 8%, from
47% in April 2004 to 39% in January 2006. The
intermittent high levels of turbid water from the
Rio Bueno River may have protected the colo-
nies at ERB from UV radiation, which possibly
worked synergistically with temperature to
cause bleaching of corals in clear water (Jokiel
1980, Gleason and Wellington 1993). As well,
the cooler river water may have helped to mod-
erate an increase in S3T at this site. Bleaching
could also have been mitigated by the high
algal cover at this site. Algal cover dominated
by a Rhodophyta assemblage at ERB was high
in 2005 and remained high in 2006.
While the 2005 bleaching event around
DBR was extensive, it was unlike the events
occurring on the bleached reefs of the U.S.
Virgin Islands as no outbreak of disease (Miller
et al. 2006) or overgrowth of the encrusting
tunicate Trididemnum solidum were observed
(pers. obs.). Also, at the Jamaican study sites
total mortality was low for many coral species
(Table 2a, b).
This study documents a very different
interpretation of the factors influencing the
“phase shift” changes. Elevated subsurface sea
water temperatures apparently alter the popula-
tion structure and diversity of a coral reef much
fast than eutrophication, over fishing and her-
bivore mortality. As well, the recovery of DBR
will likely be inhibited by the severe impact of
bleaching on key species associated with the
“phase shift reversal” described by Idjadi et
al. (2006). Because A. cervicornis reproduc-
tive strategy emphasizes asexual fragmentation
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over sexual recruitment (Tunnicliffe 1981) and
the remaining populations are so small and
subject to intense predation, it is considered
unlikely that acroporids will rapidly recover at
DBR. A site inside Discovery Bay with a for-
merly large population of A. cervicornis has not
recovered in almost three decades (Wapnick et
al. 2004).
The reasons for the lack of recovery of
the acroporids on the north coast of Jamaica
are myriad but doubtlessly include the severe
reduction in gamete producing adults (Quinn
and Kojis 2006a) and the low survival of frag-
ments due to predation. During a 49 month
period only seven acroporid spat recruited to
terracotta tiles deployed at 3m and 9m on the
WFR (Quinn and Kojis 2006b) and during 32
months only a two acroporid spat recruited at
DBR. Acroporid settlement represented 0.7% of
the total coral settlement during the entire sam-
pling period. In contrast, recruits of Poritidae
and Agariciidae were abundant on settlement
plates and these two species can be expected to
rebound fairly quickly after a bleaching event
at the Jamaican sites, if a sufficient number of
colonies survive.
Acroporidae spat settlement rates in the
Caribbean are commonly lower than other
coral families (Rylaarsdam 1983) in the region
and much lower than Acroporidae settlement
rates on reefs in the South Pacific (Kojis and
Quinn 2001, Quinn and Kojis 1999, 2003b).
Current observations suggest that the long-term
survival of A. cervicornis is threatened by a
number of factors including bleaching, disease,
predation, and a low rate of sexual recruitment
(Knowlton et. al. 1981, Woodley et al. 1981).
Given the overall severe decline and local
extinction of populations of A. cervicornis, it
is unlikely that this species will recover unless
successful recruitment of sexually produced
planulae occurs.
ACKNOWLEDGMENTS
This publication was made possible
through support provided by the National
Institutes of Health through the University
of Mississippi under the terms of agreement
No. R21 TW006645 funded by the Fogarty
International Center and the National Institute
for Research Resources for the International
Cooperative Biodiversity Groups. The opin-
ions expressed herein are those of the authors
and do not necessarily reflect the views of the
National Institutes of Health or the University
of Mississippi. We are grateful for the use of
the facilities of the Discovery Bay Marine
Laboratory, University of the West Indies.
The manuscript was greatly improved by the
constructive comments from T. J. Goreau, C.
Morrall and S.C. Steiner. Additional financial
support was provided by Tropical Discoveries
Fund. Peace Corps Volunteer C. Rhoads and T.
Downes kindly assisted on several of the dives.
This is Discovery Bay Marine Laboratory pub-
lication #751. John Farchette III is gratefully
acknowledged for translating the abstract.
RESUMEN
La estructura de la comunidad de la mayoría de los
arrecifes del Caribe ha cambiado dramáticamente desde
la década de 1980. Invocando una analogía de la química,
Hughes llamó en 1994 a este fenómeno un “cambio de
fase” para describir el cambio de un hábitat dominado
por corales a uno dominado por macroalgas en la costa
del norte de Jamaica. La pérdida de cobertura coralina es
ejemplificada por la muerte de Acropora spp. en Discovery
Bay, Jamaica. Antes de 1970, eran características de las
comunidades coralinas de profundidades bajas e interme-
dias caribeñas, las acumulaciones densas, altas y mono-
específicas de Acropora palmata (coral cuerno de alce) y
de A. cervicornis (coral cuerno de venado). A inicios de
la década de 1980, la cubierta coralina viva de A. cervi-
cornis era >21% en varios sitios alrededor de Discovery
Bay, pero para 1987 había declinado a menos de 1%. No
se reestablecieron poblaciones grandes de Acropora en la
vecindad del Discovery Bay por casi dos décadas. En 1995,
la cubierta de A. cervicornis en Dairy Bull Reef (DBR) era
de 0.6% y aumentó a cerca de 10.5% en 2004. Para 2005,
la cobertura de A. cervicornis aumentó hasta 44%. Este
aumento en la cobertura de A. cervicornis es parte de una
importante “reversión de fase” informado por Idjadi et al.
en 2006. Una población aislada de A. cervicornis exhibió
una cobertura coralina similar en Río Bueno (ERB). En
2005, ambas poblaciones del A. cervicornis se blanquea-
ron. Sin embargo, las dos poblaciones respondieron de
manera diferente. El blanqueamiento diezmó la población
de A. cervicornis en DBR; los sobrevivientes fueron
atacados por Coralliophilia, y la cobertura A. cervicornis
158
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (Suppl. 1): 149-159, May 2008
habia declinado a <0.5% en junio 2006. La población en
ERB se recuperó del blanqueamiento con una disminución
mínima en la cobertura de coral. El reclutamiento coralino
fue examinado durante tres períodos de reproducción en
DBR. Solamente hallamos dos reclutas de Acropora en
los azulejos - uno en 2003 y otro en 2005. Los Acropora
representaron solamente el 0.7% del total de reclutas en los
azulejos durante todo el período de muestreo.
Palabras claves: Acropora cervicornis, restauración,
reclutamiento, plánula.
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... D. cylindrus is considered to have medium resistance to white plague and bleaching FIGURE 6 | Colonies of D. cylindrus in different condition including: (a) healthy, (b) healthy with associated fauna, (c) fragmented due to anchors breakage, (d) with tissue mortality at the base likely to lead to fragmentation, and (e) dead and colonized by macroalgae and sponges. (Ward et al., 2006;Quinn and Kojis, 2008;Bruckner and Hill, 2009). For example, in Florida (Jaap, 1985), Honduras (Riegl et al., 2009), Jamaica (Quinn and Kojis, 2008) and Puerto Rico (Bruckner and Hill, 2009) from 1983 to 2008, 0-8% of D. cylindrus colonies experienced bleaching, whereas >40% of Acropora spp., Agaricia spp., Helioseris cucullata, O. annularis, Porites spp. ...
... (Ward et al., 2006;Quinn and Kojis, 2008;Bruckner and Hill, 2009). For example, in Florida (Jaap, 1985), Honduras (Riegl et al., 2009), Jamaica (Quinn and Kojis, 2008) and Puerto Rico (Bruckner and Hill, 2009) from 1983 to 2008, 0-8% of D. cylindrus colonies experienced bleaching, whereas >40% of Acropora spp., Agaricia spp., Helioseris cucullata, O. annularis, Porites spp. and Siderastrea siderea colonies experienced bleaching. ...
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... Most likely because the archipelago remains outside the main impact route of hurricanes and has not been subjected to coastal development (Zubillaga et al. 2005;. However, it is imperative to maintain constant surveillance across the whole MPA because A. cervicornis is known to be highly vulnerable to natural and anthropogenic disturbances such as diseases (Gladfelter 1982;Verde et al. 2016), recurrent epizootic events (Knowlton 1992;Williams and Miller 2005;Miller et al. 2014;Goergen et al. 2019), increase of storms frequency and habitat degradation (Hernández-Delgado et al. 2014;Goergen et al. 2019), and episodes of high thermal stress that can lead to loss of disease resistance (Quinn and Kojis 2008;Muller et al. 2018). Even in cases where the population's dynamic seems stable and without disturbances, the growth rate of patches is below equilibrium . ...
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Acropora cervicornis is one of the most important coral species in shallow reefs of the Caribbean as it provides habitat and structural complexity to several species of invertebrates and fish. However, the distribution range of A. cervicornis has shrunk and collapsed considerably in the last five decades, due to a combination of factors including the increase of disease prevalence, storm frequency, and anthropogenic threats. Despite being classed as “Critically Endangered” in the IUCN Red List, information regarding its population status and condition across large Caribbean coralline areas is limited. Herein we conducted the first Marine Protected Area (MPA) scale survey for this species at the Los Roques archipelago, which included visual census across 127 sites to determine the abundance, spatial distribution, habitat type, and patch morphology of A. cervicornis. We selected 11 sites, where this species was predicted and reported to be ubiquitous, to determine live A. cervicornis cover, its recent and old mortality cover, and white band disease prevalence as proxies for coral health. We found Acropora cervicornis in only 29% of the surveyed sites, with dispersed and scattered patches prevailing upon continuous patches. Moreover, the latter were located near the largest human population settlements, and inside the low protection zones of the MPA where fishing and touristic activities are permitted. The photomosaic survey showed that more than 75% A. cervicornis patches showed an average live cover above 27%, low prevalence of white band disease (<7%), and low macroalgal abundance (<10%); suggesting that Los Roques still holds healthy populations. Our results indicate that the persistence of this species urgently requires re-evaluating current MPA zoning, especially following recent evidence of overfishing and inadequate law enforcement. This study provides a baseline of A. cervicornis populations in Los Roques and Southern Caribbean that can be later used for local population management and conservation.
... Sincere thanks to three anonymous reviewers for critical feedback during review, Eugenia Sampayo, Chris Doropoulos, and Emma Kennedy for constructive discussions during the writing of the manuscript, and Nick Evensen for the branching, corymbose, and digitate Acropora photos in figure 2. Two clear outliers exist in the Caribbean dataset: an unusually high record of 7190.5 recruits per m 2 from Jamaica (Quinn & Kojis 2007), and a record of 1127.8 recruits per m 2 from the Bahamas (Avery & Liddell 1997). Revisiting the Kojis and Quinn (2007) study, a total of 302 Scleractinia and Millepora recruits were recorded from eight 208 cm 2 recruitment tiles (four per array) across 4 sampling periods, which equates to 453.7 recruits per m 2 . ...
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... Increases in the density of the grazing sea urchin Diadem antillarum, following its epidemic die-off in the 1980s, have been associated with greater coral recruitment in Jamaica [35,36]. After mortality from bleaching in 2005 reversed some of these gains [28,37], coral cover is once again increasing [28]. ...
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... The population decline that began in the early 1980s was triggered by a combination of disease (Aronson & Precht 2001) and predator outbreaks (Knowlton et al. 1990). Natural population recovery has been very rare, not only due to recurrent epizootic events (Williams & Miller 2005, Miller et al. 2014) but also because of low sexual and asexual recruitment (Knowlton et al. 1990, Quinn & Kojis 2005, degradation of critical habitats (Goreau 1992, Hernández-Delgado et al. 2014, episodes of high thermal stress (Quinn & Kojis 2008), and intense physical disturbances (e.g. hurricanes; Goreau 1992). ...
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... The year 2005 was a record hurricane season, with four category 5 storms, including Katrina(Eakin et al. 2010). Even though disease prevalence in octocoral populations increased significantly, no major octocoral mass mortalities were reported, except for Plexaurella spp.(Quinn and Kojis 2007), which is the only Caribbean gorgonian coral genus carrying Symbiodinium clade C(Van Oppen et al. 2005). Clade B, which is prevalent in most Caribbean gorgonians, is more bleaching resistant(Lewis and Coffroth 2004; van Oppen et al. 2009). ...
Chapter
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... The increasing trend of sub-massive forms shows those coral reef ecosystems are slowly recovering from the pressures such as the bleaching in 1998 and 2010 and by the Indian Ocean Tsunami (Muley et al., 2000;Venkataraman, 2000;Thanikachalam, 2010) in the past. Despite high sedimentation, settlement of new recruits was observed in Fijian reefs due to the presence of higher coverage of Acroporidae branching forms (Quinn and Kojis, 2008). Similar scenario persisted in GOMMBR resulting in higher settlement of the Acropora spp. ...
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... The year 2005 was a record hurricane season, with four category 5 storms, including Katrina(Eakin et al. 2010). Even though disease prevalence in octocoral populations increased significantly, no major octocoral mass mortalities were reported, except for Plexaurella spp.(Quinn and Kojis 2007), which is the only Caribbean gorgonian coral genus carrying Symbiodinium clade C(Van Oppen et al. 2005). Clade B, which is prevalent in most Caribbean gorgonians, is more bleaching resistant(Lewis and Coffroth 2004; van Oppen et al. 2009). ...
Chapter
Octocoral animal forests (Gorgoniidae and Plexauridae: Octocorallia) at both sides of tropical America provide a unique and characteristic seascape. They can reach over 2 m in height and even form a closed “canopy” in the densest communities. As a functional forest, gorgonian corals provide feeding substrate and habitat for diverse associated biota. This shallow-water fauna was evidently affected by the closure of the Isthmus of Panama, which provided new and different ecological opportunities at both sides. The different ecological settings provided opportunities for these groups to undergo separate adaptive radiations. New ecological conditions could lead to diversification in this group. At the Tropical Eastern Pacific (TEP), new planktonic resources provided new niches for suspension-feeding organisms, such as azooxanthellated gorgonian corals, and could have driven an adaptive radiation to exploit the new food sources. In the Caribbean, there is evidence of ecological speciation in some genera, and the scenario of ecological divergence as a major driver of gorgonian coral diversification is very likely. Thus far, the developmental phenotypic plasticity that we see today in transisthmian gorgonian corals is not just the product of speciation but adaptive developmental plasticity, and it needs further study. Gorgonian corals are today affected by many of the stressors predicted by global change, such as an increase in the frequency and intensity of tropical storms, rising seawater temperatures, and invasive species, yet these cnidarians seem highly resilient to bleaching and ocean acidification conditions. However, there is a link between high thermal anomalies and gorgonian coral immunity, which is associated to disease outbreaks and mass mortalities in sea fans in the Caribbean since the 1980s and more recently in the TEP.
... Dairy Bull reef has for several years been the fringing reef with the highest coral cover in the Discovery Bay area [20]. After the 2005 bleaching event there was a major loss of live coral cover, particularly of A. cervicornis [8] [21]. Dancing Lady reef in the 1970s had colonies of A. palmata and A. cervicornis growing in close proximity to the reef crest, while in deeper water there was a similar picture for Orbicella annularis and luxurious stands of Agaricia sp. ...
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Chapter
Knowledge of factors that are important in reef growth and resilience helps us understand how reefs react following major environmental disturbances including overfishing, destructive fishing practices, coral bleaching, ocean acidification, sea-level rise, algal blooms, agricultural run-off, coastal and resort development, marine pollution, increasing coral diseases, invasive species, hurricane/cyclone damage and bleaching. Research in both the Indo-Pacific and in the Caribbean show how temperature and environmental extremes have influenced coral growth, recruitment and mortality. Three dimensional topography and complexity is important for reef vitality and viability in the face of environmental stressors. Within the narrow temperature range for coral growth, corals can respond to rate of temperature change as well as to temperature per se. A rational polynomial function model for coral colony growth appears as the best-fitting model for coral growth, closely followed by exponential logistic, Gompertz, and von Bertalanffy models. Models have also been developed for many varieties of coral morphologies, as well as for polyp spacing in those morphologies. The chapter concludes with the suggestion that developing large Marine Protected Areas (MPAs) as part of an overall climate change policy for a country may be the best way of integrating climate change into MPA planning, management and evaluation.
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