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Idjadi et al 06
Coral Reefs (2006) 25: 209–211
DOI 10.1007/s00338-006-0088-7
NOTE
Joshua A. Idjadi Æ Sarah C. Lee Æ John F. Bruno
William F. Precht Æ Laurie Allen-Requa
Peter J. Edmunds
Rapid phase-shift reversal on a Jamaican coral reef
Received: 27 April 2005 / Accepted: 22 November 2005 / Published online: 15 March 2006
Ó Springer-Verlag 2006
Abstract Many Caribbean reefs have experienced a
phase-shift in community structure, the principle
Introduction
features being a decline in coral cover and an increase
Since the 1980s, the community structure of most
in macroalgal biomass. However, one Jamaican
Caribbean reefs has changed dramatically (Gardner
reef—Dairy Bull on the north shore near Discovery
et al. 2003). A phase-shift from coral- to macroalga-
Bay—is once again dominated by scleractinian corals
dominated benthic reef communities has been caused by
and several key species have returned. Living coral cover
a variety of factors, of which two of the most important
at 6–8 m depth at Dairy Bull has doubled over the past
have been the drastic reduction in abundance of Acro-
9 years and is now $54%. The absolute cover of Acro-
pora spp. (Aronson and Precht 2001) and the decline in
pora cervicornis was <1% in 1995, but increased to
herbivory (Hughes 1994).
$11% by January 2004. During this time the cover of
This phase-shift from coral to macroalgal dominance
macroalgae decreased by 90%, from 45 to 6%. We
is widely accepted as a model for the recent dynamics of
speculate that long-lived colonies of Montastraea
Caribbean reefs in general and Jamaican reefs in par-
annularis may have facilitated the recovery of this reef
ticular (Hughes 1994). Indeed, Jamaican coral reefs of-
by providing structural refugia.
ten are depicted as a classic example of reef degradation
in the Caribbean, in part because multiple agents have
Keywords Acropora cervicornis Æ Coral reef Æ
acted in concert to create widespread and persistent
Montastraea annularis Æ Phase-shift
macroalgal dominance.
Materials and methods
Communicated by Ecology Editor P.J. Mumby Surveys were performed on the fringing reef at Dairy
J. A. Idjadi
Bull, located $2 km east of Discovery Bay, Jamaica.
Department of Biological Science, University of Delaware, This reef is $500 m long and $100 m wide and is part of
Newark, DE 19716, USA a discontinuous reef system that spans most of Jamaica’s
north coast. We conducted our surveys on this reef in
S. C. Lee Æ J. F. Bruno 1995, 2003, and 2004.
Department of Marine Sciences,
The University of North Carolina at Chapel Hill, To estimate the coverage of benthic components
Chapel Hill, NC 27599-3300, USA such as corals and macroalgae, three transects each
30 m in length were haphazardly placed parallel to
W. F. Precht (&) depth contours at 6–8 m depth. Benthic community
Ecological Sciences Division, PBS&J,
2001 NW 107th Avenue, structure was quantified with standard photographic
Miami, FL 33172, USA survey techniques. We photographed the substratum
E-mail: bprecht@pbsj.com within 0.25 m2 quadrats positioned at random locations
Tel.: +1-305-5143488 along transects (n=37 quadrats in 1995, 73 in 2003,
Fax: +1-305-5949574
and 38 in 2004). Statistical comparisons of benthic
L. Allen-Requa Æ P. J. Edmunds cover components were made between 1995 and 2004
Department of Biology, California State University, data using unpaired t-tests. All data are reported
18111 Nordhoff Street, Northridge, CA 91330-8303, USA ±1 standard error.
210
study, massive corals of the M. annularis species com-
Results and discussion plex have been shown to withstand hurricane damage
(Woodley et al. 1981), and contribute substantially to
Benthic cover surveys revealed drastic changes in coral topographic complexity (Aronson and Precht 2001).
and macroalgal cover from 1995 to 2004. The rapid shift This is a striking contrast to intermediate depths
from macroalgal to coral dominance is demonstrated by (6–20 m) on the Discovery Bay west fore-reef, where
an inversion in relative percent cover (Fig. 1). Between topographic relief has been reduced by coral mortality
1995 and 2004 scleractinian cover nearly doubled from and bioerosion. At these intermediate depths, coral
23.0±3.5 to 53.5±3.5% while macroalgal cover was cover presently is 5–15%, and macroalgae occupy
reduced by nearly 90% from 44.9±2.8 to 5.7±1.6%. Of >60% of the substratum (Cho and Woodley 2002).
the scleractinians Acropora cervicornis showed the most M. annularis-generated high structural complexity of the
dramatic increase in cover from 0.6±0.4 to 10.5±3.2%, Dairy Bull reef is likely to be an important difference
appearing in thickets between massive colonies of compared to nearby reefs (Bechtel et al. 2006), and it
Montastraea annularis reminiscent of the fore-reef may be that reefs without it are less resilient. If this
community structure that was present at Discovery Bay scenario is accurate, M. annularis species complex may
prior to Hurricane Allen (Rylaarsdam 1983; Fig. 2a). function as a foundation taxon (sensu Bruno and
There were no significant differences among sampling Bertness 2001) that facilitated the recovery of the Dairy
periods in cover of M. annularis; however, the cover of
other reef-building coral species increased significantly,
especially Agaricia agaricites and Porites astreoides
(Fig. 2b).
The benthic community at Dairy Bull in 2004 is
similar to benthic structure on pre-phase-shift Jamaican
reefs of the 1970s when average coral cover was $55%
(Huston 1985). The dramatic reduction in the abun-
dance of macroalgae to only 5.7±1.6% cover in 2004 is
even more striking and is similar to the cover recorded
prior to the Diadema mortality of 1983–84 (Hughes
1994). The presence of residual populations of living
coral at Dairy Bull in the 1990s (Edmunds and Bruno
1996), especially long-lived colonies of M. annularis,
maintained a high degree of habitat complexity. While
habitat complexity was not directly measured in this
75
coral
50
% cover
25
macroalgae
Fig. 2 Underwater photographs of Dairy Bull Reef, Jamaica in
0 February 2003. a Underwater photograph of a Acropora cervicornis
thicket filling in the areas between large, long-lived colonies of
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
Montastraea annularis at $6 m depth at Dairy Bull. View is
reminiscent of the coral community of Jamaica in the 1960s and
1970s. b Abundance of brooding species Agaricia agaricites and
Fig. 1 Percent cover of dominant benthic components in 1995, Porites astreoides within and between large M. annularis colonies
2003, and 2004. Data are means ± 1 SE that form the foundation species of the Dairy Bull Reef complex
211
Bull reef. At present, the role of Diadema as a factor in Bruno JF, Bertness MD (2001) Habitat modification and facilita-
reef recovery at Dairy Bull is unclear because of high tion in benthic marine communities. In: Bertness MD, Hay ME,
Gaines SD (eds) Marine community ecology. Sinauer, Sun-
temporal and spatial variance of the data. However, the derland pp 201–218
possibility of high rugosity favoring Diadema popula- Cho LL, Woodley JD (2002) Recovery of reefs at Discovery Bay,
tions, and hence increased herbivory, could be usefully Jamaica and the role of Diadema antillarum. In: Proceedings of
investigated in future studies (see Bechtel et al. 2006). the 9th international coral reef symposium Bali 1:331–338
Carpenter RC, Edmunds PJ (2006) Local and regional scale
While our study is the first to report a phase-shift recovery of Diadema promotes recruitment of scleractinian
reversal on a Caribbean reef, our results are spatially corals. Ecol Lett 9:271–280
restricted to the reef at Dairy Bull. However, when Edmunds PJ, Bruno JF (1996) The importance of sampling scale in
interpreted in conjunction with other studies (i.e., ecology: Kilometer-wide variation in coral reef communities.
Edmunds and Carpenter 2001; Carpenter and Edmunds Mar Ecol Prog Ser 143:165–171
Edmunds PJ, Carpenter RC (2001) Recovery of Diadema antilla-
2006), they appear to provide some important clues rum reduces macroalgal cover and increases abundance of
about the factors responsible for promoting coral reef juvenile corals on a Caribbean reef. Proc Natl Acad Sci USA
recovery along the north coast of Jamaica and 98:5067–5071
throughout the Caribbean. This is Discovery Bay Mar- ˆ ´
Gardner TA, Cote IM, Gill JA, Grant A, Watkinson AR (2003)
Long-term region-wide declines in Caribbean corals. Science
ine Lab publication number 715, and contribution 301:958–960
number 128 of the marine biology program of California Hughes TP (1994) Catastrophes, phase-shifts, and large-scale
State University, Northridge. degradation of a Caribbean coral reef. Science 265:1547–1551
Huston MA (1985) Patterns of species diversity in relation to depth
at Discovery Bay, Jamaica. Bull Mar Sci 37:928–935
Rylaarsdam KW (1983) Life histories and abundance patterns of
References colonial corals on Jamaican reefs. Mar Ecol Prog Ser 13:249–
260
Aronson RB, Precht WF (2001) Evolutionary paleoecology of Woodley JD, Chornesky EA, Clifford PA, Jackson JBC, Kaufman
Caribbean coral reefs. In: Allmon WD, Bottjer DJ (eds) LS, Knowlton N, Lang JC, Pearson MP, Porter JW, Rooney
Evolutionary paleoecology: the ecological context of macro- MC, Rylaarsdam KW, Tunnicliffe VJ, Wahle CM, Wulff JL,
evolutionary change. Columbia University Press, New York, Curtis ASG, Dallmeyer MD, Jupp BP, Koehl MAR, Neigel J,
pp 171–233 Sides EM (1981) Hurricane Allen’s impact on Jamaican coral
Bechtel JD, Gayle P, Kaufman L (2006) The return of Diadema reefs. Science 214:749–755
antillarum to Discovery Bay: patterns of distribution and
abundance. In: Proceedings of the 10th international coral reef
symposium, Okinawa (in press)
DOI 10.1007/s00338-006-0088-7
NOTE
Joshua A. Idjadi Æ Sarah C. Lee Æ John F. Bruno
William F. Precht Æ Laurie Allen-Requa
Peter J. Edmunds
Rapid phase-shift reversal on a Jamaican coral reef
Received: 27 April 2005 / Accepted: 22 November 2005 / Published online: 15 March 2006
Ó Springer-Verlag 2006
Abstract Many Caribbean reefs have experienced a
phase-shift in community structure, the principle
Introduction
features being a decline in coral cover and an increase
Since the 1980s, the community structure of most
in macroalgal biomass. However, one Jamaican
Caribbean reefs has changed dramatically (Gardner
reef—Dairy Bull on the north shore near Discovery
et al. 2003). A phase-shift from coral- to macroalga-
Bay—is once again dominated by scleractinian corals
dominated benthic reef communities has been caused by
and several key species have returned. Living coral cover
a variety of factors, of which two of the most important
at 6–8 m depth at Dairy Bull has doubled over the past
have been the drastic reduction in abundance of Acro-
9 years and is now $54%. The absolute cover of Acro-
pora spp. (Aronson and Precht 2001) and the decline in
pora cervicornis was <1% in 1995, but increased to
herbivory (Hughes 1994).
$11% by January 2004. During this time the cover of
This phase-shift from coral to macroalgal dominance
macroalgae decreased by 90%, from 45 to 6%. We
is widely accepted as a model for the recent dynamics of
speculate that long-lived colonies of Montastraea
Caribbean reefs in general and Jamaican reefs in par-
annularis may have facilitated the recovery of this reef
ticular (Hughes 1994). Indeed, Jamaican coral reefs of-
by providing structural refugia.
ten are depicted as a classic example of reef degradation
in the Caribbean, in part because multiple agents have
Keywords Acropora cervicornis Æ Coral reef Æ
acted in concert to create widespread and persistent
Montastraea annularis Æ Phase-shift
macroalgal dominance.
Materials and methods
Communicated by Ecology Editor P.J. Mumby Surveys were performed on the fringing reef at Dairy
J. A. Idjadi
Bull, located $2 km east of Discovery Bay, Jamaica.
Department of Biological Science, University of Delaware, This reef is $500 m long and $100 m wide and is part of
Newark, DE 19716, USA a discontinuous reef system that spans most of Jamaica’s
north coast. We conducted our surveys on this reef in
S. C. Lee Æ J. F. Bruno 1995, 2003, and 2004.
Department of Marine Sciences,
The University of North Carolina at Chapel Hill, To estimate the coverage of benthic components
Chapel Hill, NC 27599-3300, USA such as corals and macroalgae, three transects each
30 m in length were haphazardly placed parallel to
W. F. Precht (&) depth contours at 6–8 m depth. Benthic community
Ecological Sciences Division, PBS&J,
2001 NW 107th Avenue, structure was quantified with standard photographic
Miami, FL 33172, USA survey techniques. We photographed the substratum
E-mail: bprecht@pbsj.com within 0.25 m2 quadrats positioned at random locations
Tel.: +1-305-5143488 along transects (n=37 quadrats in 1995, 73 in 2003,
Fax: +1-305-5949574
and 38 in 2004). Statistical comparisons of benthic
L. Allen-Requa Æ P. J. Edmunds cover components were made between 1995 and 2004
Department of Biology, California State University, data using unpaired t-tests. All data are reported
18111 Nordhoff Street, Northridge, CA 91330-8303, USA ±1 standard error.
210
study, massive corals of the M. annularis species com-
Results and discussion plex have been shown to withstand hurricane damage
(Woodley et al. 1981), and contribute substantially to
Benthic cover surveys revealed drastic changes in coral topographic complexity (Aronson and Precht 2001).
and macroalgal cover from 1995 to 2004. The rapid shift This is a striking contrast to intermediate depths
from macroalgal to coral dominance is demonstrated by (6–20 m) on the Discovery Bay west fore-reef, where
an inversion in relative percent cover (Fig. 1). Between topographic relief has been reduced by coral mortality
1995 and 2004 scleractinian cover nearly doubled from and bioerosion. At these intermediate depths, coral
23.0±3.5 to 53.5±3.5% while macroalgal cover was cover presently is 5–15%, and macroalgae occupy
reduced by nearly 90% from 44.9±2.8 to 5.7±1.6%. Of >60% of the substratum (Cho and Woodley 2002).
the scleractinians Acropora cervicornis showed the most M. annularis-generated high structural complexity of the
dramatic increase in cover from 0.6±0.4 to 10.5±3.2%, Dairy Bull reef is likely to be an important difference
appearing in thickets between massive colonies of compared to nearby reefs (Bechtel et al. 2006), and it
Montastraea annularis reminiscent of the fore-reef may be that reefs without it are less resilient. If this
community structure that was present at Discovery Bay scenario is accurate, M. annularis species complex may
prior to Hurricane Allen (Rylaarsdam 1983; Fig. 2a). function as a foundation taxon (sensu Bruno and
There were no significant differences among sampling Bertness 2001) that facilitated the recovery of the Dairy
periods in cover of M. annularis; however, the cover of
other reef-building coral species increased significantly,
especially Agaricia agaricites and Porites astreoides
(Fig. 2b).
The benthic community at Dairy Bull in 2004 is
similar to benthic structure on pre-phase-shift Jamaican
reefs of the 1970s when average coral cover was $55%
(Huston 1985). The dramatic reduction in the abun-
dance of macroalgae to only 5.7±1.6% cover in 2004 is
even more striking and is similar to the cover recorded
prior to the Diadema mortality of 1983–84 (Hughes
1994). The presence of residual populations of living
coral at Dairy Bull in the 1990s (Edmunds and Bruno
1996), especially long-lived colonies of M. annularis,
maintained a high degree of habitat complexity. While
habitat complexity was not directly measured in this
75
coral
50
% cover
25
macroalgae
Fig. 2 Underwater photographs of Dairy Bull Reef, Jamaica in
0 February 2003. a Underwater photograph of a Acropora cervicornis
thicket filling in the areas between large, long-lived colonies of
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
Montastraea annularis at $6 m depth at Dairy Bull. View is
reminiscent of the coral community of Jamaica in the 1960s and
1970s. b Abundance of brooding species Agaricia agaricites and
Fig. 1 Percent cover of dominant benthic components in 1995, Porites astreoides within and between large M. annularis colonies
2003, and 2004. Data are means ± 1 SE that form the foundation species of the Dairy Bull Reef complex
211
Bull reef. At present, the role of Diadema as a factor in Bruno JF, Bertness MD (2001) Habitat modification and facilita-
reef recovery at Dairy Bull is unclear because of high tion in benthic marine communities. In: Bertness MD, Hay ME,
Gaines SD (eds) Marine community ecology. Sinauer, Sun-
temporal and spatial variance of the data. However, the derland pp 201–218
possibility of high rugosity favoring Diadema popula- Cho LL, Woodley JD (2002) Recovery of reefs at Discovery Bay,
tions, and hence increased herbivory, could be usefully Jamaica and the role of Diadema antillarum. In: Proceedings of
investigated in future studies (see Bechtel et al. 2006). the 9th international coral reef symposium Bali 1:331–338
Carpenter RC, Edmunds PJ (2006) Local and regional scale
While our study is the first to report a phase-shift recovery of Diadema promotes recruitment of scleractinian
reversal on a Caribbean reef, our results are spatially corals. Ecol Lett 9:271–280
restricted to the reef at Dairy Bull. However, when Edmunds PJ, Bruno JF (1996) The importance of sampling scale in
interpreted in conjunction with other studies (i.e., ecology: Kilometer-wide variation in coral reef communities.
Edmunds and Carpenter 2001; Carpenter and Edmunds Mar Ecol Prog Ser 143:165–171
Edmunds PJ, Carpenter RC (2001) Recovery of Diadema antilla-
2006), they appear to provide some important clues rum reduces macroalgal cover and increases abundance of
about the factors responsible for promoting coral reef juvenile corals on a Caribbean reef. Proc Natl Acad Sci USA
recovery along the north coast of Jamaica and 98:5067–5071
throughout the Caribbean. This is Discovery Bay Mar- ˆ ´
Gardner TA, Cote IM, Gill JA, Grant A, Watkinson AR (2003)
Long-term region-wide declines in Caribbean corals. Science
ine Lab publication number 715, and contribution 301:958–960
number 128 of the marine biology program of California Hughes TP (1994) Catastrophes, phase-shifts, and large-scale
State University, Northridge. degradation of a Caribbean coral reef. Science 265:1547–1551
Huston MA (1985) Patterns of species diversity in relation to depth
at Discovery Bay, Jamaica. Bull Mar Sci 37:928–935
Rylaarsdam KW (1983) Life histories and abundance patterns of
References colonial corals on Jamaican reefs. Mar Ecol Prog Ser 13:249–
260
Aronson RB, Precht WF (2001) Evolutionary paleoecology of Woodley JD, Chornesky EA, Clifford PA, Jackson JBC, Kaufman
Caribbean coral reefs. In: Allmon WD, Bottjer DJ (eds) LS, Knowlton N, Lang JC, Pearson MP, Porter JW, Rooney
Evolutionary paleoecology: the ecological context of macro- MC, Rylaarsdam KW, Tunnicliffe VJ, Wahle CM, Wulff JL,
evolutionary change. Columbia University Press, New York, Curtis ASG, Dallmeyer MD, Jupp BP, Koehl MAR, Neigel J,
pp 171–233 Sides EM (1981) Hurricane Allen’s impact on Jamaican coral
Bechtel JD, Gayle P, Kaufman L (2006) The return of Diadema reefs. Science 214:749–755
antillarum to Discovery Bay: patterns of distribution and
abundance. In: Proceedings of the 10th international coral reef
symposium, Okinawa (in press)