Computational Biology and Chemistry 32 (2008) 311–314

Contents lists available at ScienceDirect

Computational Biology and Chemistry

journal homepage: www.elsevier.com/locate/compbiolchem

Review Article

Climate change, global warming and coral reefs: Modelling

the effects of temperature
M. James C. Crabbe ∗
Luton Institute for Research in the Applied Natural Sciences, Faculty of Creative Arts, Technology and Science, University of Bedfordshire, Park Square, Luton LU1 3JU, UK

a r t i c l e i n f o a b s t r a c t

Article history: Climate change and global warming have severe consequences for the survival of scleractinian (reef-
Received 15 April 2008 building) corals and their associated ecosystems. This review summarizes recent literature on the
Accepted 16 April 2008 influence of temperature on coral growth, coral bleaching, and modelling the effects of high tempera-
ture on corals. Satellite-based sea surface temperature (SST) and coral bleaching information available on
Keywords: the internet is an important tool in monitoring and modelling coral responses to temperature. Within the
SST
narrow temperature range for coral growth, corals can respond to rate of temperature change as well as
Satellite
to temperature per se. We need to continue to develop models of how non-steady-state processes such as
Scleractinian
Coral growth
global warming and climate change will affect coral reefs.
Bleaching © 2008 Elsevier Ltd. All rights reserved.
Interdecadal
Symbiosis
IPCC
Weather
El Niño
Great barrier reef
Caribbean

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311
2. The Influence Of Temperature On Coral Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
3. Coral Bleaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
4. Modelling The Effects Of High Temperature On Corals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
5. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314

1. Introduction barriers to wave action and storms by reducing the incident wave
energy through wave reflection, dissipation and shoaling, protect-
Coral reefs, found predominantly between the tropics of Capri- ing the land and an estimated half a billion people who live within
corn and Cancer, provide an environment in which one third of all 100 km of reefs.
marine fish species and many thousands of other species are found, The growth and subsistence of corals depend on many variables,
and from which 6 million tons of fish are caught annually. This not including temperature, irradiance, calcium carbonate saturation,
only provides an income to national and international fishing fleets, turbidity, sedimentation, salinity, pH, and nutrients. These vari-
but also for local communities, which in addition rely on the local ables influence the physiological processes of photosynthesis and
fish stocks to provide nutritional sustenance. The reefs also act as calcification as well as coral survival, and as a result coral reefs occur
only in select areas of the world’s oceans. Meteorological processes
can alter these variables, and Fig. 1 summarizes their influences on
∗ Tel.: +44 1582 489265; fax: +44 1582 489212. global and synoptic scales on coral requirements for growth and
E-mail address: [email protected]. survival (Walker, 2005; Crabbe et al., 2008a). Coral reefs are cur-

1476-9271/$ – see front matter © 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.compbiolchem.2008.04.001
312 M.J.C. Crabbe / Computational Biology and Chemistry 32 (2008) 311–314

Fig. 1. Schematic diagram summarizing key meteorological processes and coral requirements controlling calcification, photosynthesis, and survival. Adapted from Crabbe et
al. (2008a) and Crabbe et al. (2008b).

rently under severe threat from climate change (Lough, 2008), as allow them to persist in what is becoming an increasingly hostile
well as from many other anthropogenic influences, such as pollu- environment.
tion and overfishing (Mumby et al., 2007; Crabbe et al., 2008b). Growth rates also depend upon minimum seasonal tempera-
This review will concentrate on the effects of temperature on tures. Changes in average winter air temperature at the flower
modern scleractinian (reef-building) corals, and the recent insights garden banks in the Gulf of Mexico were found to correspond to
that modelling can provide to increase our understanding of coral changes in Montastea annularis growth rates (Slowey and Crowley,
ecology and survival in a period of climate change. 1995). Interdecadal changes in the growth rate of the corals cor-
responded to changes in average minimum winter season air
2. The Influence Of Temperature On Coral Growth temperatures at New Orleans. Slowey and Crowley (1995) acknowl-
edged that the correspondence between the changes in the two
Many studies have examined the impact of air temperature vari- were not one to one because the influence of air temperature on
ations on coral growth rates. In a study of the correlation between water temperature depends on a number of meteorological and
air temperature and growth rates of coral from a colony of Porites oceanographic factors. The minimum temperatures over the Gulf
lutea in the Great Barrier Reef in the district of Haapiti, Australia of Mexico can be caused by the passage of fronts bringing cold dry
(Bessat and Buiges, 2001), measurements were taken from coral air from Canada, and probably this process is primarily responsible
cores. For the period from 1958 to 1990, linear regression gave a cor- for stressing corals at the flower gardens and reducing their winter
relation between air temperature 25 km from where the core was growth rate. There was a major shift towards colder winters during
taken and coral skeletal density of r = 0.37; for the same period, the the 1950s and this coincided with the decline of coral growth at the
correlation between air temperature and annual calcification rate flower gardens.
was lower at r = 0.28. These results indicated that a 1 ◦ C rise in tem-
perature would lead to an increase in the density rate of about 10.5% 3. Coral Bleaching
and an increase in the calcification rate of about 4.5%. Coral calcifi-
cation rates and extension rates have been highly correlated with Most of the pigmentation within corals is within the symbi-
sea surface temperatures (SSTs) and to a lesser extent with incom- otic algal cells—the zooxanthellae. Coral bleaching is caused by
ing solar radiation (Nie et al., 1997; Lough and Barnes, 2000). In corals losing their zooxanthellae. Thermal bleaching occurs when
juvenile corals, temperature causes a transition between isomet- the coral is exposed to prolonged above-normal (or below-normal)
ric and allometric growth scaling in warmer versus cooler years, temperatures, resulting in additional energy demands on the coral,
respectively (Edmunds, 2006; Edmunds, 2008). depleted reserves, and reduced biomass (Muller-Parker and D’Elia,
Interestingly, on the Great Barrier Reef (GBR), calcification rates 1997). The effect of high temperatures can be aggravated by high
in massive Porites colonies declined by approximately 21% in two levels of irradiance (Gleason and Wellington, 1993), although high
regions 450 km apart. This was a function primarily of a decrease UV radiation is not a primary factor in causing mass bleaching
in linear extension (16%) with a smaller decline in skeletal density (Hoegh-Guldberg, 1999). Coral reefs within or near the western
(6%) (Cooper et al., 2008) and contrasts with previous studies on the Pacific warm pool (WPWP) have had fewer reported bleaching
environmental controls on growth of massive Porites on the GBR. In events relative to reefs in other regions. Analysis of SST data indi-
a study on reefs of East Africa (McClanahan et al., 2007) it appeared cate that the warmest parts of the WPWP have warmed less than
that it was not just the high stability of tropical environments that elsewhere in the tropical oceans, which supports the existence of
creates high biological diversity but also large temperature fluc- thermostat mechanisms that act to depress warming beyond cer-
tuations that prepares the corals for the unexpected and this may tain temperature thresholds (Kleypas et al., 2008).
 M.J.C. Crabbe / Computational Biology and Chemistry 32 (2008) 311–314 313

Corals can die as a result of bleaching, though they may partially

or fully recover from bleaching events (Lough, 2000). Bleach-
ing causes a decrease in the growth rate of corals, and the time
taken for a coral to recover from a bleaching event may be sev-
eral years or decades. If the frequency of bleaching increases, then
the capacity for coral reefs to recover is diminished (Done, 1999).
This has been observed during a number of ‘natural’ events, e.g.,
1983–1984 (Glynn, 1990), 1997–1998 (Goreau et al., 2000; Glynn
et al., 2001) and 2005 (Wilkinson and Souter, 2008). Experimental
studies show that thermal history, in addition to light history, can
influence the response of reef-building corals to thermal stress, and
therefore, have implications for the modeling of bleaching events
(Middlebrook et al., 2008). Significant levels of mortality can occur
in a bleaching event before any chance for subsequent recombina-
tion of the host-symbiont unit (Jones, 2008). To better understand
factors affecting the potential evolution of bleaching resistance in
corals in response to increased average sea temperatures, a math-
ematical model of coevolutionary interactions between partners
in a coral-algae mutualistic symbiosis has been developed (Day Fig. 2. Influence of the rate of change of temperature on the growth rate of an Acro-
pora palmata coral colony from Curaçao (Bak, 1976). R2 = 0.53. For details see the
et al., 2008). This showed that traits in mutualistic symbioses,
text.
such as thermal tolerance in corals, are potentially subject to novel
kinds of evolutionary constraints and that these constraints are
mediated by ecological dynamics. The nature of interspecific con- provided alerts for the 2005 Caribbean mass-bleaching event, indi-
trol of bleaching resistance and the mode of sexual reproduction cating that average ocean temperatures during July–October 2005
interacted to strongly influence the rate of spread of resistance for the Caribbean exceeded temperatures seen at any time during
alleles. the past 154 years (NOAA, 2008). Similar systems are in use for the
Great Barrier Reef (Maynard et al., 2008).
4. Modelling The Effects Of High Temperature On Corals Modelling of growth rates (by weight; Bak, 1976) of the branch-
ing coral Acropora palmata with temperature on a Curaçao coral reef,
The frequency that corals will be bleached in the future has using a smoothing spline to produce a nonparametric fit to the data,
been estimated by using projections of future sea surface tem- suggested that the 30-day averaged maximum daily temperature
peratures from four different general circulation models (GCMs) could explain about 3% of the variability in the time-dependent
forced by the IPCC IS92a emission scenario (Hoegh-Guldberg, growth rate (Crabbe et al., 2008a). Interestingly, the temperature
1999). The SST projections were combined with thermal thresh- correlation was negative, suggesting that during the measurement
olds for corals, derived by using the Integrated Global Ocean period, temperatures rose to higher than optimum temperatures
Services System (IGOSS) dataset provided by the Joint World and for growth, thus inhibiting coral growth, but were not sufficiently
Scientific Meteorological Organization (WMO) and United Nations high to cause bleaching of this species. Thus what would normally
Educational, Scientific and Cultural Organization (UNESCO), Joint be a positive correlation became a slight but significant negative
Intergovernmental Oceanographic Commission’s (JCOMM) Techni- correlation.
cal Commission for Oceanography and Marine Meteorology, and For Acropora palmata colonies on fringing reefs off the north
from literature and Internet reports of bleaching events. All SST coast of Jamaica, over the period 2002–2007, the rate of growth
projections indicated that the frequency of bleaching events is set of Acropora palmata was largely proportional to rate of change of
to rise rapidly, with the highest rates in the Caribbean, Southeast SST, with R2 = 0.935 (Crabbe, 2007). If we now model the growth
Asia, and Great Barrier Reef, and the lowest rates in the central rate data of Acropora palmata from Curaçao (Bak, 1976) with rate
Pacific. The frequency of bleaching events was predicted to become of change of temperature (Fig. 2), then R2 = 0.53, rather than the
annual in most oceans by 2040, and the Caribbean and South- small negative correlation if only temperature, rather than rate
east Asia are projected to reach this point by 2020, triggered by of change of temperature, is taken into account. These modelling
seasonal changes in seawater temperature rather than by El Niño studies suggest that within the narrow temperature range for coral
events. growth, corals respond to rate of temperature change as well as to
The geographical patterns and the timing of probable repeat temperature per se.
occurrences of coral mortality in the Indian Ocean have been esti-
mated (Sheppard, 2003). Forecast sea surface temperatures at 33 5. Conclusion
sites in the Indian Ocean were blended onto historical sea surface
temperatures. The forecast temperatures were estimated by using Climate processes and extremes can influence the physiolog-
the IS92a scheme, which follows a median path. These studies indi- ical processes responsible for the growth of coral reef colonies.
cate a 50% probability of SSTs being warm enough by 2030 for the Coral growth takes place within narrow limits of temperature,
occurrence of coral bleaching events at Comoros and Chagos in the irradiance, salinity, pH, and turbidity, all variables that are influ-
Indian Ocean, and by 2070 in the Saudi Arabian Gulf. enced by climate and weather. In a number of empirical models
In order to predict imminent rises in SSTs, the U.S. National for coral growth, small changes in temperature and rates of tem-
Oceanic and Atmospheric Administration’s Coral Reef Watch (NOAA perature change can significantly influence coral colony growth
CRW) develops and operationally produces satellite-based coral rates. We need to continue to develop models of how non-steady-
bleaching nowcasts and alerts available on the Internet (Liu et al., state processes such as global warming and climate change will
2006). These products are based on nighttime-only Advanced Very effect coral reefs, and on whether corals or their symbiont algae
High Resolution Radiometer (AVHRR) sea surface temperatures will evolve to keep pace with the climate and environmental
from operational NOAA polar-orbiting satellites, and, for example, changes.
314 M.J.C. Crabbe / Computational Biology and Chemistry 32 (2008) 311–314

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