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The Effects of Habitat on Coral Bleaching Responses in Kenya

Author(s): Gabriel Grimsditch, Jelvas M. Mwaura, Joseph Kilonzo, Nassir Amiyo


Source: AMBIO: A Journal of the Human Environment, 39(4):295-304. 2010.
Published By: Royal Swedish Academy of Sciences
URL: http://www.bioone.org/doi/full/10.1007/s13280-010-0052-1

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AMBIO (2010) 39:295–304
DOI 10.1007/s13280-010-0052-1

REVIEW PAPER

The Effects of Habitat on Coral Bleaching Responses in Kenya


Gabriel Grimsditch, Jelvas M. Mwaura,
Joseph Kilonzo, Nassir Amiyo

Received: 30 April 2009 / Revised: 8 February 2010 / Accepted: 15 March 2010 / Published online: 13 May 2010

Abstract This study examines the bleaching responses of lead to mortality of colonies and even a phase shift from a
scleractinian corals at four sites in Kenya (Kanamai, coral-dominated to an algal-dominated reef with a sub-
Vipingo, Mombasa and Nyali) representing two distinct sequent loss of biodiversity (Glynn 1993; Nystrom et al.
lagoon habitats (relatively shallow and relatively deep). 2000).
Bleaching incidence was monitored for the whole coral However, it is possible for corals to either resist con-
community, while zooxanthellae densities and chlorophyll ditions that cause coral bleaching or to recover from
levels were monitored for target species (Pocillopora bleaching events (resilience). Resistance can be defined as
damicornis, Porites lutea, and Porites cylindrica) during a the ability of an ecosystem to withstand disturbances
non-bleaching year (2006) and a year of mild-bleaching without undergoing a phase shift or losing neither structure
(2007). Differences in bleaching responses between habi- nor function. Resilience can be defined as the ability of a
tats were observed, with shallower sites Kanamai and system or an individual to absorb or recover from distur-
Vipingo exhibiting lower bleaching incidence than deeper bance and change, while recovering its functions. Several
sites Nyali and Mombasa. These shallower lagoons display ecological and environmental factors can affect a coral
more fluctuating thermal and light environments than the reef’s resistance and resilience to bleaching events. These
deeper sites, suggesting that corals in the shallower lagoons include the coral reef’s community species composition,
have acclimatized and/or adapted to the fluctuating envi- herbivory levels, zooxanthellae population, temperature
ronmental conditions they endure on a daily basis and have and irradiance history, connectivity to other reefs, water
become more resistant to bleaching stress. In deeper sites movement, and shading or screening properties (Grims-
that did exhibit higher bleaching (Mombasa and Nyali), it ditch and Salm 2006; Hoegh-Guldberg 1999; West and
was found that coral recovery occurred more quickly in the Salm 2003).
protected area than in the non-protected area. Furthermore, it has been shown that it is possible for
colonies to acclimatize to increased temperatures and high
Keywords Coral reef  Bleaching  Resilience irradiance levels so that they are able to resist bleaching
events when they occur. Acclimatization can be defined as
the ability of an organism to undergo phenotypic changes
INTRODUCTION in response to stress in the natural environment that result
in the readjustment of the organism’s tolerance levels to
Climatic changes linked with global warming are causing that stress (Coles and Brown 2003). Threshold tempera-
increased frequency of coral bleaching events worldwide. tures that induce coral bleaching-related mortality vary
Coral bleaching events are usually associated with above- worldwide (from 27°C in Easter Island; Wellington et al.
average water temperatures and high irradiance levels that 2001, to 36°C in the Arabian Gulf; Riegl 1999) according
cause the disruption of the obligate symbiosis between to the maximum water temperatures that are normal in the
scleractinian corals and micro-algal zooxanthellae (Hoegh- area, implying a capacity of corals and/or zooxanthellae to
Guldberg 1999). The symbiotic zooxanthellae are expelled acclimatize to high temperatures depending on their envi-
and the coral is left in a weakened state that can eventually ronment. Furthermore, corals that are regularly exposed to

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Fig. 1 Map of study sites

stressful environmental conditions have been shown to The objective of this study is thus to determine whether
acclimatize and exhibit physiological tolerance to elevated corals in different habitats display different bleaching
temperatures and UV-radiation that exceed normal thres- responses (i.e. resistance and resilience) and to suggest
holds (Brown et al. 2000, 2002a, b; Coles and Brown which environmental characteristics are responsible for the
2003). variation in response.
In order to combat the worst effects of climate change
and to conserve this valuable ecosystem, it is important to
determine which factors affect coral reef bleaching resis- MATERIALS AND METHODS
tance and resilience and to apply this knowledge in man-
agement plans. Although bleaching events cannot be The study was carried out in 2006 and 2007 at four patch
prevented by managers, by implementing appropriate reefs within lagoons: Mombasa Marine Park (MMP),
management responses it is possible to help a coral reef Nyali, Kanamai, and Vipingo in Kenya at a latitude of
recover from bleaching and to mitigate the worst effects. *4°S (Fig. 1). The coral reefs in this area are coastal
This study examines the bleaching responses of corals at fringing crest reefs that form a natural barrier to the wave
four sites (Nyali, Mombasa Marine Park, Kanamai, and energy of the ocean creating lagoons where this study was
Vipingo, Fig. 1) representing two distinct lagoon habitats carried out. MMP is protected from all extractive uses,
on the Kenyan coast (deeper and shallower lagoons). The including fishing, but is a primary site for snorkeling and
sites were characterized using environmental parameters SCUBA diving for tourists. Nyali is a marine reserve where
such as depth, water flow, light, and temperature. Bleach- only traditional fishing methods are allowed and is also
ing responses were monitored for the general coral com- regularly visited by tourists. Kanamai and Vipingo are
munity and zooxanthellae densities and chlorophyll levels unprotected shallow sites characterized by extensive fish-
were monitored for target species (Pocillopora damicornis, ing practices, but have lower tourist pressure.
Porites lutea, and Porites cylindrica) during a non- Five large and healthy coral colonies for each target
bleaching year (2006) and a mild bleaching year (2007). species (Pocillopora damicornis, Porites cylindrica, and

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AMBIO (2010) 39:295–304 297

Porites lutea) were mapped at the two shallower lagoons percentages of each bleached colony in 20 haphazard 2 m2
(Kanamai and Vipingo, 0.4 m depth and 0.6 m at low tide, areas at each site.
respectively) and the two deeper lagoons MMP and Nyali, Water temperature was monitored using underwater
1.4 m and 1.8 m depth at low tide, respectively). Target temperature loggers that were left at each site throughout
species were chosen according to general abundance the whole year for 2 years (Fig. 2). Light was monitored
and bleaching susceptibility, with one highly susceptible using a light meter for only 24-h periods on the same day
(Pocillopora damicornis), one moderate (Porites cylindri- and time at each site to avoid theft and algae over growth
ca), and one resistant (Porites lutea) target species (Obura of the light meter. Water flow was measured using a clod
2001). Coral size class data for all genera were collected card method (McClanahan et al. 2005). Plaster-of-Paris
for two fixed 25 9 2 m transects at each site. The coral clod cards were made in an ice-cube tray, dried and
size classes recorded were 0–5, 5–10, 10–20, 20–40, weighed before being deployed in the field in small net
40–80, 80–160, and [160 cm. Line intercept transect data bags that were tied to the substrate. They were collected
for substrate types (hard coral, soft coral, macroalgae, turf 24 h later, dried, and weighed again in order to determine
algae, sponges, rock, sand, other invertebrates, etc.) were how much had dissolved.
also collected on the same 25 m transects.
When bleaching occurs in Kenya, it occurs during the
late northeast monsoon from mid March when doldrum Analysis
conditions become most intense to late April when the first
storms of the southeast monsoon induce cooling of surface Current speed in g d-1 (24 h) was obtained by dividing
waters. In the 2 years of the study, sampling began before weight loss by deployment time. Weight loss (g) was then
bleaching during the northeast monsoon (February and converted to flow speed (Vw = cm s-1) using the equation
early March), continued during bleaching season of the Vw = (DWclod - 0.932)/2.357, which was obtained by
northeast monsoon (late March and April), and finished studies of clod dissolution in a flume with controlled water
during the recovery period of the southeast monsoon (May, flow (Anzai 2001).
June and July). Statistical analyses were performed using the program
Coral fragments were collected monthly from the Minitab (version 10.1). Differences in zooxanthellae den-
mapped colonies using a chisel. The fragments were sity were investigated using the Man–Whitney U-test and
transported to the laboratory submerged in seawater in the Kruskal–Wallis test.
small plastic bottles and were held in an aerated seawater
tank. Coral tissue was removed from the skeleton using a
water jet. Seawater was then added to the tissue to make up RESULTS
a practical volume, usually 500 ml. The tissue slurry was
homogenized using a hand-held homogenizer and 1.0 ml of Temperature
homogenate was loaded into a Sedgwick-Rafter chamber.
Using a compound binocular microscope and a magnifi- The shallower lagoons, Kanamai and Vipingo, displayed
cation of 9400, the number of zooxanthellae in 10 random the most extreme temperature conditions (Fig. 2), with
quadrats was recorded. The counting chamber was then higher standard deviations (±1.9°C and ±1.5°C, respec-
reloaded and another 10 random quadrats were counted. tively), higher maximum temperatures (36.2°C and 33.8°C,
The area of the coral skeleton was measured using the respectively), lower minimum temperatures (24.0°C
aluminum foil method; the area/weight ratio of aluminum and 23.7°C, respectively), and bigger differences between
foil was determined and coral skeletons were then wrapped diurnal and nocturnal temperatures (?0.9°C and ?0.4°C,
in this foil. The foil was then trimmed to fit the skeleton respectively) than the deeper lagoons. Between the two,
area and weighed. Using the weight of the trimmed foil and Kanamai displayed the more extreme fluctuations, as well
the area/weight ration for the foil, the skeleton area could as the highest average temperature overall (27.8°C)
be determined. (Fig. 3).
To measure chlorophyll a levels, a known volume of the The deeper lagoons of MMP and Nyali displayed less
homogenate was filtered through a GF/F filter paper and pronounced temperature variations with lower standard
the filtrate was dissolved in 10 ml of 90% acetone. It was deviations (both ±1.4°C) as well as smaller maximum–
then centrifuged for 10 min at 2,000 rpm and the chloro- minimum (?8.9°C and ?8.1°C, respectively) and diurnal–
phyll absorbance was read using a spectrophotometer. nocturnal (both ?0.2°C) differences than the shallower
If it was observed, bleaching incidence was recorded in lagoons. However, the deeper lagoons displayed bigger
the water using a random swim method. This consists of temperature increases from the 2006 to 2007 northeast
noting the size class, genus, and bleached/pale/dead monsoon bleaching seasons, with a ?0.9°C increase in

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Fig. 2 Temperature graphs for


all four study sites during the
period of the study (March 2006
to June 2007). Temperature was
logged every half an hour

Nyali and a ?0.8°C increase in MMP compared to a lagoon MMP. Unfortunately, light data for Nyali was not
?0.7°C increase in both Kanamai and Vipingo. available due to technical problems. Kanamai (4913 ±
14,315 lux) and Vipingo (4758 ± 10,489 lux) both dis-
Light played much higher average light levels, higher standard
deviations, and higher maxima (70,846 lux and 110,223 lux
As expected, the shallow lagoons, Kanamai and Vipingo, respectively) than MMP (average of 518 ± 879 lux,
also displayed more extreme light regimes than the deeper maximum of 5167 lux). Although light data is unavailable

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AMBIO (2010) 39:295–304 299

38 Max Acropora and Pocillopora only accounted for 3.7% of


Avg Kanamai’s coral cover and 0.7% of MMP’s coral cover.
36 Min
34 Bleaching and Mortality Incidence
32
Bleaching was first observed during late April of 2007. It
30
was measured using the random swim method described in
°C

28 the ‘‘Method’’ section. The shallower lagoons, Kanamai


26 and Vipingo, exhibited lower levels of combined paling,
24
bleaching, and mortality (4.4% and 6.3% of total coral
area, respectively) than the deeper lagoons, Nyali and
22 MMP (35.1% and 26.9% of total coral area, respectively).
20 In May, at the height of the bleaching event, in Kanamai
MMP Nyali Vipingo Kanamai 1.3% of corals surveyed were pale, 2.8% were bleached,
and 0.4% were dead. In Vipingo, 0.9% were pale, 5.1%
Fig. 3 Average (±standard deviation), maximum, and minimum
water temperatures during study period were bleached, and 0.3% were dead. In MMP, 5.4% were
pale, 20.2% were bleached, and 1.4% were dead. In Nyali,
14.7% were pale, 19.7% were bleached, and 0.9% were
for Nyali, it is likely that light levels are similar to MMP dead.
given the depth and sediment levels of the site. During the recovery period in June, MMP still exhibited
relatively high bleaching incidence (9.8%) but low mor-
Water Flow tality (0.9%). Nyali exhibited the highest mortality (11.7%)
and also relatively high bleaching (4.5%). In July, mortality
There was no clear trend between water flow at deeper and increased to 13.1% in Nyali and 2% in MMP, but bleaching
shallower lagoons. MMP consistently displayed the highest incidence in MMP decreased to 6.0%, indicating recovery
average water flow during both neap (6.4 cm/s) and spring (Fig. 4).
tide cycles (9.0 cm/s). Kanamai (5.4 cm/s) displayed sim- Overall, Pocillopora and Porites accounted for the
ilar water flow to Nyali (5.5 cm/s) and Vipingo (5.6 cm/s) highest proportion of bleached colonies, but differed
during neap tide and, although unfortunately data for spring greatly in their mortality rates. In MMP, Nyali, and Vi-
tide at Nyali is unavailable, it appears that MMP stands out pingo, Pocillopora accounted for 37, 47.1, and 33.3% of
as the site displaying highest water flow. Overall, Kanamai bleached colonies, respectively. In Kanamai, MMP, and
displayed the lowest water flow during both neap and Nyali, Porites accounted for 47.2, 28.3, and 13.8% of
spring tides (8.1 cm/s). bleached colonies, respectively (Fig. 5). However, during
the recovery period of June and July, Porites experienced
Substrate Cover low bleaching-related mortality while Pocillopora experi-
enced high mortality. Porites colonies only experienced
MMP and Kanamai displayed higher hard coral cover mortality in Vipingo, where they accounted for 12.5% of
(28.4% and 23.0%, respectively) while Nyali and Vipingo total bleaching-related mortality, whereas Pocillopora
displayed lower hard coral cover (14.7% and 10.6%, accounted for the highest proportion of bleaching-related
respectively), showing no trend with depth. Macroalgal mortality in all sites (from 37.5% in Vipingo to 90% in
cover was higher in deeper sites, MMP (25.5%) and Nyali MMP) (Fig. 6). Furthermore, of the three species only
(21.0%), than in shallower sites, Vipingo (8.3%) and Pocillopora damicornis suffered high mortality in both
Kanamai (3.7%). Hard corals and turf algae dominate the 2006 (50% overall mortality) and 2007 (47% overall
substrate at all sites. mortality) with colonies in Kanamai experiencing the
lowest mortality rate in both years.
Coral Cover

All sites are dominated by massive and branching Porites Zooxanthellae Densities
colonies. 90.5% of MMP, 80.3% of Kanamai, 59.9% of
Nyali, and 42.2% of Vipingo coral cover was massive and Average zooxanthellae densities for marked colonies
branching Porites. Vipingo (10.2%) and Nyali (8.3%) (Pocillopora damicornis, Porites lutea, and Porites cyl-
displayed the highest relative coral cover of the more indrica) were higher for all species at all sites in 2006 than
bleaching-susceptible genera, Acropora and Pocillopora. in 2007. Average zooxanthellae densities were also higher

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300 AMBIO (2010) 39:295–304

Fig. 4 Bleaching, paling, and Kanamai Vipingo


mortality incidence during 2007 100% 100%
Dead
80% Bleached 80%

% of coral area
Pale
60% Normal 60%

40% 40%

20% 20%

0% 0%
March May June July March May June July

MMP Nyali
100% 100%

80% 80%
% of coral area

60% 60%

40% 40%

20% 20%

0% 0%
March May June July March May June July

n=36 n=15 n=46 n=87 in shallower sites, Kanamai and Vipingo, than in deeper
100% Other sites, MMP and Nyali, for all species. In general, Kanamai
90% Por Bra exhibited the highest average zooxanthellae densities while
% of bleached coral

80% Acr MMP displayed the lowest. Porites lutea colonies exhib-
70% Por Col
ited the highest zooxanthellae densities while Pocillopora
60% Por Mas
damicornis displayed the lowest densities, with Porites
50% Poc
cylindrica in between the two (Fig. 7).
40%
In 2006, colonies in MMP, Nyali, and Kanamai dis-
30%
played similar zooxanthellae density trends with decreases
20%
in the warm months of February and March, and recovery
10%
in the following months as temperature decreased. Colo-
0%
Kanamai Vipingo MMP Nyali
nies in Vipingo, on the other hand, exhibited increasing
densities for all species during February and March
Fig. 5 Proportion of bleached corals by genera. Por Bra, Porites followed by decreases from April to June. In 2007, aver-
branching; Acr Acropora; Por Col, Porites columnar; Por Mas, age zooxanthellae densities in Pocillopora damicornis
Porites massive; Poc, Pocillopora
remained very low (\1 9 106 cells cm-2) in all sites with
recovering densities in July. Porites lutea and Porites
cylindrica colonies showed a general increasing trend in
n=11 n=8 n=10 n=42
100% Other average zooxanthellae densities from March to July
Por Mas (Fig. 7).
80% Acr
% of dead corals

Poc
60% Chlorophyll a Levels
40%
Chlorophyll a concentrations were generally higher at
20% shallower sites, Kanamai and Vipingo, than at deeper sites,
Nyali and MMP, and Kanamai exhibited by far the highest
0% chlorophyll a concentrations while MMP exhibited the
Kanamai Vipingo MMP Nyali
lowest. Porites cylindrica consistently exhibited the
Fig. 6 Proportion of corals exhibiting bleaching-related mortality by highest chlorophyll a concentrations at all sites, while
genera. Por Mas, Porites massive; Acr, Acropora; Poc, Pocillopora Pocillopora damicornis consistently displayed the lowest

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AMBIO (2010) 39:295–304 301

Fig. 7 Average zooxanthellae


densities for selected species
(PD Pocillopora damicornis,
PC Porites cylindrica, PL
Porites lutea) during 2006
and 2007

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Fig. 8 Average mg chlorophyll Porites cylindrica Porites lutea


a per m3 for selected species 7.E-02 3.E-02
Kanamai
(Porites cylindrica, Porites 6.E-02 Vipingo
lutea, and Pocillopora Nyali 2.E-02

mg Chl a/m3
mg Chl a/m3
damicornis) during 2007 5.E-02 MMP
2.E-02
4.E-02
3.E-02 1.E-02
2.E-02
5.E-03
1.E-02
0.E+00 0.E+00
Mar May Jun Jul Mar May Jun Jul

Pocillopora damicornis
6.E-02

5.E-02
mg Chl a/m3

4.E-02

3.E-02

2.E-02

1.E-02

0.E+00
Mar May Jun Jul

concentrations, with Porites lutea in between the two smaller volumes of water in the shallower lagoons proba-
(Fig. 8). bly explain the difference in temperature and light regimes;
Chlorophyll a concentrations in Pocillopora damicornis shallower lagoons absorb but also lose heat more quickly
colonies generally decreased progressively at all sites from and attenuate less light than in the deeper lagoons. That
March to July as colonies bleached and failed to recover. year no bleaching was observed on the deeper reef slope
Porites lutea and Porites cylindrica colonies in the shal- (anecdotal observations).
lower site Kanamai displayed peaks during March and then Moreover, differences in temperature and light regimes
decreasing concentrations to July. In MMP and Nyali, the could explain differences in bleaching responses between
concentration peaks for these two species were not as similar-depth sites. Kanamai is a shallower site than
pronounced and generally occurred in May or June Vipingo, displaying higher average temperatures, standard
(Fig. 8). deviation in temperatures, difference between maximum
and minimum temperature, diel temperature variation,
maximum light level, and standard deviation in light levels.
DISCUSSION This may explain why Kanamai also displayed lower
bleaching incidence and bleaching-related mortality than
Differences in bleaching responses during 2007 between Vipingo. Similarly, MMP is a shallower site than Nyali,
shallower and deeper lagoonal sites were observed, with displaying higher average temperatures and difference
shallower sites, Kanamai and Vipingo, exhibiting lower between maximum and minimum temperature. Nyali
bleaching and paling incidence than deeper sites, Nyali and experienced increased warming during the 2007 northeast
MMP, in both haphazard sampling and marked Pocillopora monsoon bleaching period (?0.9°C to 2006) compared to
damicornis colonies. These results suggest that corals in MMP (?0.8°C to 2006). These factors could explain why
the shallower lagoons have acclimatized to the more Nyali exhibited higher bleaching incidence than MMP.
extreme environmental conditions they endure on a daily On the other hand, water flow did not correlate well with
basis and have become more resistant to thermal stress than depth and bleaching incidence, and although corals in
corals in the deeper lagoons. Nyali suffered much higher bleaching incidence than
Shallower sites, Kanamai and Vipingo, displayed more Kanamai and Vipingo, all these sites displayed very similar
extreme temperature and light regimes than deeper sites, water flow velocities. It therefore appears that light and
Nyali and MMP, with higher maxima, lower minima, lar- temperature histories were more influential than water flow
ger standard deviations, and larger diel variations. The in determining the bleaching responses of corals.

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AMBIO (2010) 39:295–304 303

Moreover, average zooxanthellae densities and chloro- genera occurs in Vipingo, which is a shallower site
phyll a concentrations are higher in Kanamai and Vipingo exhibiting low bleaching incidence. However, differences
than in Nyali and MMP, with highest densities in Kanamai. in community composition could partially explain varying
Studies in the region have shown that corals with higher bleaching responses between similar-depth sites. Nyali
zooxanthellae densities are more resistant to bleaching exhibited higher bleaching incidence than MMP and also a
(Grimsditch et al. 2008), a hypothesis that is further con- higher proportion of area covered by bleaching-susceptible
firmed by these results. In addition, Porites lutea displayed genera. The same applies to Vipingo compared to
the highest densities and was the most tolerant to bleach- Kanamai.
ing, while Pocillopora damicornis exhibited the lowest During the recovery period, colonies in Nyali displayed
densities and was the most susceptible to bleaching. Zoo- higher mortality than those in MMP, while a higher pro-
xanthellae population dynamics could thus also partly portion of corals in MMP remained bleached but did not
explain bleaching responses at different sites. die. It thus seems that colonies in Nyali colonies were
Interestingly, species that suffered less bleaching (Por- being overgrown by algae more quickly than those in
ites lutea and Porites cylindrica) did not exhibited MMP, which remained in a bleached state longer. This
decreases in chlorophyll a concentrations through time, and could be due to the protected status of MMP and the
actually exhibited increases in chlorophyll a during possibility that herbivorous fish populations are higher than
bleaching and recovery months in some cases. However, in Nyali. McClanahan and Arthur (2001) showed that fish
Pocillopora damicornis, which was more susceptible to densities (not including damselfish and wrasses) in non-
bleaching, displayed decreasing chlorophyll a concentra- fished sites (including MMP as study site) were 62.6
tions with time, indicating that bleaching events cause individuals per 100 m2 compared to 12.2 individuals per
disruptions in pigments as well as symbionts. 100 m2 in fished sites (including Nyali as a study site).
The genus most affected by bleaching at Nyali, MMP, In conclusion, the data from this study show that during
and Vipingo was Pocillopora, and it accounted for most of the 2007 bleaching season, corals in the shallow lagoons of
the bleaching-related mortality in all sites. Porites also Kanamai and Vipingo were more resistant to bleaching
made up significant proportions of bleached corals in stress than corals in the deeper lagoons of MMP and Nyali,
MMP, Nyali, and Kanamai. However, Porites colonies probably due to a history of higher light and temperature
hardly exhibited any bleaching-related mortality, showing extremes and variation at the shallower sites. Interesting
that although this genus does bleach and pale it is tolerant differences were also found between the deeper lagoons at
to bleaching. Despite their susceptibility to bleaching, Nyali and MMP—on the one hand MMP reefs experience
Acropora corals form a very small proportion of bleached slightly larger diel variation in temperature, and on the
corals in most sites because they are scarce, except for in other they experienced less warming in 2007. Whether this
Vipingo where they form a larger proportion of the total acclimatization is due to short-term phenotypic changes or
coral are and subsequently also a larger proportion of due to a longer-term process of adaptation through natural
bleached corals. The high susceptibility for Pocillopora is selection is a question that could be further researched.
consistent with their K-selected life history strategy as a Pocillopora was the genus most susceptible to bleaching-
fast-growing but short-lived genus (Obura, 2001). How- related mortality, especially at Nyali, and Porites branch-
ever, it could also be indicative of a longer term phase shift ing, massive and columnar all displayed high resistance to
in the scleractinian coral community, with more suscepti- bleaching. Among all these patterns of bleaching suscep-
ble genera being eliminated by consecutive bleaching tibility, it is not clear whether the acclimatization is host-
events. Acropora, another vulnerable coral genus, abun- based or symbiont-based, and monitoring of the zooxan-
dance in the region has already been dramatically reduced thellae clades in corals at these sites could help in
due to past bleaching events (Obura 2002). However, coral answering this question. Finally, bleached corals in MMP
recruitment in the region is overwhelmingly dominated by suffered lower levels of mortality than in Nyali, possibly
Pocillopora recruits (Obura, 2002), counteracting high associated with higher rates of herbivory due to MMP’s
Pocillopora mortality and thus perhaps mitigating Pocil- protected status. While herbivory is often related to resil-
lopora decline in the near future. ience of reefs and the influence of herbivores assisting
All sites were dominated by massive and branching recovery by coral recruits, this result suggests herbivory
Porites corals, discounting the possibility that large varia- may also mediate interactions between algae and bleached
tions in coral community compositions determined corals, and assist in the survival of bleached corals. Pri-
bleaching responses. Bleaching-susceptible genera such as orities for research should explore management strategies
Pocillopora and Acropora did not necessarily form a larger that could increase coral reef resilience to climate change,
proportion of the coral population in sites most affected by and whether marine parks are an effective tool in bolstering
bleaching. The highest area percentage covered by these ecological resilience.

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Acknowledgments This study was made possible by the generous Riegl, B. 1999. Corals in a non-reef setting in the southern Arabian
support of the MacArthur Foundation and the Swedish International Gulf (Dubai, UAE): Fauna and community structure in response
Development Agency. to recurring mass mortality. Coral Reefs 18: 63–73.
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