Assessing the Potential for Natural Recovery and Coral Restoration Techniques for Enhancing Coral Habitat in Jamaica

Norman J. Quinn
Discovery Bay Marine Lab, University of the West Indies, Discovery Bay, St. Ann, Jamaica

Barbara L. Kojis
Division of Fish and Wildlife, St. Thomas, US Virgin Islands 00802

W. Austin Bowden-Kerby
CounterPart International, 1200 18th Street NW, Suite 1100, Washington, DC 20036

Abstract - The reduced biodiversity of Caribbean coral reefs has been attributed to disturbances of various types. Many of the once abundant coral species have disappeared from Jamaican coral reefs with algae taking their place resulting in a less attractive reef habit with fewer fish which has affected the Jamaican economy. The mortality of Acropora species has been a particularly important driving force in shaping the transition. The abundance of Acropora spat settling in the Caribbean are lower than other coral families and much lower than on South Pacific reefs. A natural recovery through sexual reproduction is unlikely in the next decade. The long-term survival of remnant A. cervicornis populations is threatened unless successful sexual reproduction is restored. This study seeks to assess methods of restoring A. cervicornis populations by creating pockets of greater reef health around surviving populations. Experi- mental transplants using several techniques are underway to develop a suitable technology for restoring A. cervicornis populations in Jamaica. In one technique the mean survivorship ranged from 4% - 78% after 55 weeks. Mortality associated with damselfish (Stegastes) and fireworms (Hermodice carunculata) were observed. By increasing coral habitat biomass and complexity we can increase the abundance of fish populations which will result in greater fish catches for subsistence fishers. As well, more coral and fish biodiversity improves the attractiveness of the reef community for divers and snorkelers making Jamaica a more appealing destination for tourists.

provides an attraction for travelers to Jamaica. The Jamaican coral reef is an active ecosystem, which despite experiencing major damage is still functional and fascinating. The major culprits of coral reef decline are hurricanes, pollution, disease, chronic overfishing, bleaching episodes and physical damage from anchors. In 1980, Hurricane Allen produced >10 m waves which slammed into the coral reefs and began a period where Jamaican reefs declined in diversity and abundance. B. Coral Reefs in Decline Only two to three decades ago, Acropora cervicornis was one of the most important reef-building corals on Jamaican reefs [1]. However, this species has declined throughout the region, becoming locally extinct on many reefs [2]. The Caribbean wide decline of Acropora corals in recent decades has serious consequences to coral reef biodiversity, coastal geology, and to the fisheries and tourism economies of the region. Such is the present situation that the three described Caribbean species are being considered for listing as endangered or threatened species [3], a first for reef-building scleractinian corals globally. There are many factors that are considered to have influenced the rapid reduction in populations. Waste runoff from land contributes to algal overgrowth which competes with the corals for oxygen and light and space and may actually coral fecundity [4] prevent settlement of coral larvae [5]. In the Caribbean, chronic over fishing is an ever increasing threat to coral reefs [6]. The effect of over fishing is to promote algal overgrowth and thereby reduces the space for new coral to settle [7]. The decline has also been linked to the lack of predators, resulting in increases in corallivorous gastropods [8], increases in Acropora harming Stegastes fish, [9] decline of grazing Diadema sea urchins [10], algal overgrowth [11], and associated coral disease [12]. Serveral decades ago, the Acropora cervicornis population at the West Fore reef at Discovery Bay was

I. INTRODUCTION A. Coral Reef Ecology Coral reefs are often called the rainforests of the marine realm. The beauty and biodiversity of its plants and animals attracts snorkelers and divers from around the world helping to support the Jamaican tourist industry. The reefs also provide habitat and shelter that support fish populations which provide a livelihood for subsistence fishermen. As well, the shore is protected from the action of waves by the physical presence of the reef. The reef builds and regenerates the beaches, which

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reduced to a mere 4% of its original cover by Hurricane Allen [13]. Eight years later when Hurricane Gilbert struck Jamaica, the few recovering stands of Acropora that had survived Allen were smashed again [14]. Subsequent incidents of predation and disease reduced the A. cervicornis population to <1% of their original levels [15]. Even on reefs where measures to address the root causes of coral decline have been implemented, A. cervicornis populations do not appear to be recovering, as larval recruitment is very sparse [5]. A likely hypothesis for a lack of larval-based recovery is that most of the surviving A. cervicornis populations are either too young or too small to produce planula that would normally re-colonize reefs [16]. Current observations suggest that the long-term survival of A. cervicornis is threatened unless root causes of decline are more effectively addressed throughout the region, and successful sexual reproduction occurs. C. Coral Restoration Technology Artificial reefs have been used for centuries for creating underwater habitat and increasing biological activity [17]. Not all reefs can be restored and some corals are more receptive to restoration than others [18, 19]. In the Pacific, the coral reefs in Fiji are much more resiliant than the reefs in the Marianas Islands [20]. Due to the high expenses involved in restoration actions and the variety of practices that could be used, we need innovative but general models that will guide us in restoring damaged habitats. The gardening of coral reefs with sexual and asexual recruits after their mariculture in situ within special nursery areas may serve as such a framework for developing restoration protocols suitable for sharing through a network of Marine Protected Areas throughout Jamaica. The Acropora cervicornis is a keystone species of critical importance to biodiversity, fisheries, and tourism interests. A. cervicornis is particularly vital as fisheries habitat due to it being the only large open-branched coral species of reef slope, back reef, and logoonal environments, so the loss of this species represents a loss to the biodiversity and essential fisheries habitat of Caribbean reefs. In the Indian Ocean transplanted corals have been shown to enhance fish abundance and diversity [21]. In Tobago, West Indies, Acropora cervicornis transplantation has been done with a survival of 30-35% of the colonies eight months after transplantation. Of the surviving colonies the growth rate ranged from 6.5 to 11.7 cm year -1 [22]. A. cervicornis is suitable as a transplanted coral because of its relatively fast growth rate. D. Is Natural Recovery Likely in the Next Decade?

Many hypotheses have been proposed to explain the lack of recovery of the coral assemblages [2, 6, 15]. One of the most likely is the lack of sexual recruitment of the major reef building corals in the family Acroporidae (5, 23, 24, 25]. The potential for coral to naturally recover should be examined before efforts to restore reefs are undertaken. Efforts to transplant or restore reefs are unnecessary if the population has the capacity to recovery through natural means. We have sought to determine the abundance, depth distribution and seasonal variability of sexual recruitment of acroporid corals along the West Fore Reef at Discovery Bay, Jamaica. It was hypothesized that owing to a paucity of mature acroporid colonies there would be low levels of larval recruitment. Recruitment patterns of juvenile corals are important to the overall community structure of coral reefs [26, 27]. Studies frequently focus on juvenile coral >1 cm diameter called “visible” recruitment. Since the post-settlement process may change the number of recruits [28], recruitment onto tiles is considered to be a better indication of the availability of planktonic planulae than the study of “visible” corals [27]. As well, early settlement stages are more vulnerable than adults to changes in nutrient [29] and sediment levels [30]. As part of this project were are investigating the settlement of planula on tiles placed on the reef. Small, fragmented colonies still occur and it is hoped that at least some of the remaining coral populations are now reasonably stable, being composed of the more resistant survivors of major bleaching and disease epidemics. However, Acropora cervicornis is not returning to reefs where it was formerly common [31], as sexual recruitment of Acropora is rare or absent in the Caribbean [25, 26, 32]. Given the low levels of successful sexual reproduction of A. cervicornis its long-term survival is threatened and a proactive approach may be needed to in initiate the restoration of healthy populations. E. Experimental Restoration Efforts on Jamaican Reefs With the implementation of no-take MPAs and measures to address the root causes of coral reef decline in several Caribbean countries, patches of increased reef health can be expected to return. Once the fish, crustaceans, and other species that positively influence coral health have become more abundant, corals should begin to fare better on the reefs. Abundant herbivorous fish populations have been shown to keep algae in check, helping enable corals to survive well even in nutrient enriched waters [33]. Discovery Bay Marine Laboratory (DBML) and Counterpart International are working with local stakeholders including the Montego Bay Marine Park, Northern Jamaica Conservation Association and several

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local hotels to help restore the biomass of Acropora cervicornis on select reefs sites on the Jamaican north coast. The strategy is to create pockets of greater reef health around surviving A. cervicornis populations, using techniques such as removal of coral predators, weeding excess seaweed, removing Stegastes damselfish, and experimental introduction of sea urchins to control disease-harboring algae [34] around Acropora populations. Another strategy involves propagation of A. cervicornis branches taken from healthy populations for experimental propagation in areas large population once existed. This paper reports on our transplantation experiments. II. METHODS A. Coral Recruitment Coral recruitment arrays were constructed by attaching four 208 cm2 unglazed terracotta tiles to a PVC array (Fig. 1). The tiles were smooth on one side and had 12 ridges on the other side. Two of the tiles were arranged horizontally and two vertically on the array. The tiles were ~ 0.8 m above the substrate.

depth of ~10 m. Buckets filled with sea water were used to temporarily house the fragments while they were transported by boat to the laboratory. 1) Sampling Effect Experiment. We examined source colonies where the fragments were taken from to see if collecting breaking fragements caused an increase in mortality. On September 2, 2004, colored cable ties were attached to 40 source colonies ~ 2 cm below where fragments were broken off. An additonal 40 source population colonies were used as a control and were tagged with different color ties at the sample approximate position where the cable tag ties were located on the donor colonies. Three months later the colonies were reexamined for mortality. 2) A frame Experiment. Some fragments were then attached to 14 wire mesh “A-frames” using plastic cable ties. The base for the coral fragments were wire mesh (0.8 m x 1.2 m) bent at 90o . The frames were tagged with numbered tags for identification. Two weights were attached to the base of each frame for stability. Five fragments were placed on the outer side of each frame (Fig. 2).

Fig. 1. Coral recruitment collector deployed on the West Fore Reef, Discovery Bay, Jamaica.

The arrays were initially installed in late March 2001 and were replaced in October 2001 (“summer 2001” sampling period), April 2002 (“winter 2001 / ‘02”), October 2002 (“summer 2002”), in April 2003 (“winter 2002/‘03”) and October 2003 (“summer 2003”). The scleractinian and milleporan corals were counted (standardized to number recruits m-2) and identified to family where possible using a binocular microscope. B. Coral Restoration Around Discovery Bay several healthy Acropora cervicornis populations have been located of sufficient size to be use for coral restoration experiments. On June 2, 2004, 160 fragments of Acropora cervicornis ranging in length from 5 to 14 cm were collected from colonies at Dairy Bull Reef (18º28.24 N; 77º24 W) at a

Fig. 2. M. Stennett monitors growth and survival of Acropora cervicornis fragments on A-frame. Dead tips of fragments are likely the result of Hermodice carunculata predation.

The experimental A-frames were then deployed at several sites around Discovery Bay: Columbus Park Reef (18º46.448 N; 77º41.428 W) - depth 6 m; East Back Reef (18º46.856 N; 77º40.454 W) - depth 4 m; the Blue Hole (18º47.252 N; 77º41.662 W) - depth 2 m, Back Reef Canoe Channel (18º28.305 N; 77º24.950 W) - depth 2 m; and outside the bay at West Fore Reef (18º47.252 N; 77º41.662 W) - depth 6 m (Fig. 3). At Columbus Park Reef, the water was quite turbid with a visibility frequently <10 m. The substrate was silt. A live, healthy reef with a dense A. cervicornis population had previously lived there for at least 6,000 year b.p. [31]. The East Back Reef is an area subjected to breaking waves and clear water. The substrate is sand mixed

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with rubble. The A-frames at Blue Hole are located in a back reef, lagoon environment with slightly turbid water and mixed sand, rubble substrate. The Canoe Channel site is just behind the reef crest at the West Fore Reef in a inter reef sandy area and receives clear ocean water. The substrate is sand with sea grass. Black plastic netting shade cloth was suspended over these A-frames at the shallow Canoe Channel site to reduce sun intensity. The fragments were monitored for survivorship and growth. Survivorship percentage was calculated by dividing the total number of live fragments by the total number of fragments. A fragment was considered live if some living tissue was observed. Missing fragments were counted as dead. Growth was measured from the fragment base to the apical polyp and then each branch from the branching point to the apical polyp. The lengths were added together and actual linear length and percentage growth calculated. 3) Cement Base Experiment A 50/50 cement- sand mixture was flattened, pressed and dried into the shape of a ~ 5 cm circular disk. Two holes were punctured through the disk before drying. A nylon monofilament line would then be looped through the holes to secure small (< 7 cm) fragments of A. cervicornis and A. prolifera. Two hundred and eleven disks were constructed. Disks were secured to a wire mesh sheet with the fishing (Fig. 3). The fragments were monitored for survivorship and growth.

The cultures were monitored for both growth and survivorship. Growth rates and survivorship was determined as in the previous experiment.

Fig. 4. Acropora cervicornis attached to nylon monofilament line in line restoration experiment.

III. RESULTS A. Coral Recruitment Over a 30 month period only seven acroporid spat settled – 3 in summer 2001 (16 spat m-2) and 4 in summer 2003 (23 spat m-2) (Fig. 5). No Acropora recruitment occurred at 14 m, 19 m, 26 m or 33 m. Settlement only occurred on tiles at a depth of 3 m and 9 m (Fig. 6). Acroporid settlement represented <1 % of the total coral settlement during the sampling period. Corals from the family Poritidae were the most common (50%) followed by Milleporidae (27%), Agaracidae (10%) and unknown spat (28%).



Spat m

log scale

Fig 3. Coral framents on cement disks on wire mats (left) and on A-frames (right) in transit from construction site to reef site.



4) Line Experiment On June 16, 2004, sixteen A. cervicornis fragments were collected from colonies at Dairy Bull Reef. After ~ 4 hrs in the shaded buckets, the fragments were transported to the Canoe Channel, where they were attached to monofilament nylon lines to create a set of line cultured corals suspended underneath the mesh of the underwater culture tables. On September 3, 2004, the experiment was repeated with 40 fragments also collected at Dairy Bull Reef. This time the fragments were deployed on lines in the Canoe Channel within 30 min of being collected (Fig. 4).



3 2 1 03 '02 e r '0 er ' 0 er ' 0 2-' '0 1 u mm umm inter '0 umm Winter S S S W
Fig. 5. Density of recruited Acropora spat normalized to m-2 over time.

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Acropora 1000


Spat m

log scale














180 160 140 120 100 80 60 40 20 0
Jul. 2 Aug.20 Nov. 17 Feb. 28
Time Post transplantation

% Net Growth



Fig. 6. Density of recruited Acropora spat normalized to m-2 over depth. Log scale (n + 1)

B. Coral Restoration 1) Sampling Effect Experiment. Of the 40 tagged colonies in the source population n where fragments were collected, only 19 were found on after 3 months. Of these 19 source colonies, six were dead (32% mortality) and 13 were alive (68% survival) with an apical polyp. After 3 months 32 control colonies were located. Eight colonies were dead (25% mortality) and 24 colonies were alive (75% survival). A survey of the Acropora cervicornis population at Dairy Bull Reef found a total of 106 dead tips on colonies otherwise alive. This mortality is likely the result from fireworm predation and damselfish bites and is rather intensive. 2) A-frame Experiment After 55 weeks Columbus Park Reef had the highest % survivorship - 78% with 7% at East Back Reef, 32% at Blue Hole, and 4% at Canoe Channel (Fig. 7). Although Hurricane Ivan passed south of Jamaica in September 2004, very large waves crashed over the reef and damaged the experiment at Canoe Channel. After 39 weeks, the overall mean net growth of corals on the A-frames was 7.3 cm (s.e.7.0 cm; n = 63), ranging from a mean of 1.1 cm (s.e.0.6cm; n = 16) at Canoe Channel to 15.3 cm (s.e. 4.6cm; n = 40) at Columbus Park (Fig. 8). That represents a mean growth rate of 0.19 cm week-1. or an annualized rate of 9.7 cm year-1.

Fig. 8. Growth rates of corals on A-frame experiment at Columbus Park reef (CP), East Back Reef (EBR), Blue Hole (BH) and Canoe Channel (CC).

3) Cement Disk Experiment Initially the survival and growth of the colonies was very good during the first couple of months. Colonies quickly grew and established multiple growing branches (Fig. 9). The survivorship after 37 weeks was 19% - 40 colonies of the original 211 fragments were still alive. Many of the colonies were affected by Huricane Ivan which passed south of Jamaica in September 2004. After one year (June 2005) few complete colonies are alive. Several colonies have live areas, but no apical poly. Based on eight weeks of observations the annulaized growth rate was 23 cm year -1.

Fig. 9. Measuring growth on Acropora cervicornis fragment transplanted on cement disk.


% Survival




0% 1 6 11 16 21 26 31 36 41 46 51 56

Columbus Park

East Back Reef


Blue Hole

Canoe Channel

Fig. 7. % survival of fragments over time. Weeks on x axis.

4) Line Experiment Within two days of the start of the experiment most of coral had bleached. After five days we observed 100% bleaching and the start of algal growth over the white tissue. None of the colonies recovered. Five days after the September transplantation, none of the corals had any bleaching inspite of water temperatures being at their annual maximums [36]. After 26 weeks the survivorship of the second attempt had a 32% survival. Considering that Hurricane Ivan passed two weeks into

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the experiment this is considered a rather low mortality considering the circumstances. By June 2005, the remaining colonies are doing extremely well with annualized percentage growth exceeding 400%. Colonies are vigorously growing from both sides and have numerous branches with no sign of Hermodice carunculata predation or Stegastes bites. IV. DISCUSSION A. Coral Recruitment The lack of recovery of acroporids can either be linked to localized factors or a general paucity of planktonic recruits. It is important to identify the mechanism at work. If high algal density is thwarting community recovery by occupying planulae settlement space, then a reduction in algal cover would free up settlement space and coral recruitment should increase. If there are few coral planulae being produced by acroporids, then available space will have little bearing on the potential for acroporid recovery. In that case, transplanting of acroporid fragments may provide the best chance of improving the rate of recovery in the immediate future [18]. It has been suggested that predators were preventing the recovery of staghorn populations as predation rates increased to levels beyond a predicted threshold as the result of a phase shift [2]. While the level of juvenile mortality from predation may be a key factor in the restructuring of some coral communities [37, 38], we suggest that the recruitment failure of acroporid planulae is more influential than juvenile mortality. Because Caribbean acroporid reproductive strategy emphasized asexual fragmentation over sexual recruitment [39] and the existing populations are so small it is considered unlikely that they will rapidly recover through growth and asexual fragmentation. The evidence from this study supports the hypotheses [25] that sexual recruitment of acroporids is currently low because there are few adult colonies to produce gametes and because of the potential lack of fertilization success when the distance between colonies is so great (Allee effect) [40]. With the additional impediment to settlement of a macro algal overgrown substrate, the possibility of successful sexual recruitment is further reduced. Currently, sexual reproduction is not likely to contribute to rapid recovery of acroporid communities, although the presence of some planulae recruits does indicate that sexual recruitment are available to reestablish populations on reefs with no acroporids. The lack of sexual recruits is likely a function of both the present paucity of large populations of Acropora and the poor health of the existing populations. Currently at the West Fore Reef at Discovery Bay, they represent < 1 % of the

coral cover at < 10 m depth [5]. Furthermore, existing populations are under attack by various coral diseases [41], which are likely to reduce the fecundity of individual colonies. Considering the massive decline in acroporid populations Caribbean wide, it is likely that the larval pool has considerably decreased. It has been hypothesized that that recovery of acroporid populations in the Caribbean may be hindered by high growth rates and frequent fragmentation which may reduce the amount of energy available for sexual reproduction and result in low gamete production rates and, consequently, low larval recruitment rates [23]. Metabolic resources may be used to enhance growth at the expense of sexual reproduction. The abundance of Acropora spat settling in the Caribbean is commonly lower than other coral families and much lower than Acropora settlement rates on reefs in the South Pacific [25]. It appears that the long-term survival of remnant A. cervicornis populations is threatened unless successful sexual reproduction is restored. A recent Acropora workshop, recommended experimental acroporid mariculture and other propagation techniques, along with transplantation, and reattachment of dislodged Acropora fragments as a feasible strategy to rebuild degraded population [42]. The results from this study suggests that the success of restoration is very site specific. Local variations of abiotic conditions like waves exposure, salinity, depth, turbidity can influence survival. As well biotic influences such as abundance of fire worms and damselfish can influrence growth and survival. Frames on sand did considerably better that frames on rock or rubble, being away from lurking places of predatory Coraliophila snails and Hermodice fire worms, as well as being removed from Stegastes damselfish territories. However, if the frames were far from the reef, algal overgrowth became a problem due to a lack of grazing fish. The method of weighing the frames down with cement blocks at some sites resulted in the inadvertent creation of ideal Stegastes habitat, with subsequent negative impacts to the corals. The size of the wire mesh also affected this outcome, with small mesh becoming ideal shelter for the damselfish, plus allowing more surface for the algal farming activities of the fish. Solutions to these problems were implemented in the second phase of experiments in January and March 2005, and involve increasing the wire mesh apertures to 20 x 20 cm, and locating each frame on bare sand 1 - 3m from rocky areas, providing a barrier to predators but close enough for herbivorous fish to visit the frames regularly for cleaning. The specific outcomes of mortality and growth will be presented in detail in a subsequent paper. B. Management Implications Having a unique and special ecological role as well

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as great natural beauty, arguements for the restoration of Acropora cervicornis to reefs there it is expirated are compelling for many reasons. A. cervicornis is an essential fish habitat for many commercial fish, as it serves as vital nighttime resting places as well as provides shelter from predation during the day. The process of restoring staghorn corals served as a point of unification between sometimes antagonistic groups: government managers, NGOs, conservationists, fishermen, and the tourism industry. Restoring healthy breeding populations of staghorn coral at low-cost, even if for relatively limited areas, could have important reef conservation implications. This project supported existing management strategies and heightening coral reef conservation awareness during workshops, highlighting the vital importance of no-take marine protected areas in restoring healthy ecological balance to reefs. The research sites were located within existing coral reef conservation areas, taking advantage of the increased ecological health, while helping contribute to the recovery of biodiversity within the management areas. Important partnerships were formed with members of the tourism industry as their water sports staff participated in coral restoration workshops at DBML and some even hosted their own workshops on their properties. It is possible that the hospitality industry will appreciate the natural beauty of coral reefs and assist with reef conservation activities at local, national, and regional levels and specifically by sponsoring the Acropora restoration work. Acknowledgments We are grateful for the financial assistance provided by USAID through the CWIP2 program and for the use of the facilities of the Discovery Bay Marine Laboratory, University of the West Indies. Additional financial support was provided by Counterpart International, Tropical Discoveries Foundation, and 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 opinions 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. M Stennett deserves particular recognition for her assistance in the initial phase of the coral restoration experiment. This is DBML publication #710. REFERENCES [1] T.F. Goreau and J.W. Wells, “The shallow–water

Scleractinia of Jamaica: revised list of species and their vertical distribution range,” Bull. Mar. Sci., vol. 17, pp. 442-453, 1967. [2] N. Knowlton, J.C. Lang, and B.D. Keller, “Case study of natural population collapse: Post hurricane predation of Jamaican staghorn corals,” Smithson. Contrib. Mar. Sci., vol. 31, pp. 1-25, 1990. [3] W.F. Precht, M.L. Robbart, and R.B.Aronson, “The potential listing of Acropora species under the US Endangered Species Act,” Marine Pollution Bulletin vol, 29, pp. 534-536, 2004. [4] B.L. Kojis and N.J. Quinn, “Seasonal and depth variation in fecundity of Acropora palifera at two reefs in Papua New Guinea,” Coral Reefs vol. 3, pp. 165-172, 1984. [5] N.J. Quinn and B.L. Kojis, “Patterns of sexual recruitment of acroporid coral populations on the West Fore Reef at Discovery Bay, Jamaica,” Rev. Biol. Trop., vol. 53(1), pp. 83-90, 2005. [6] T.P. Hughes, 1994. “Catastrophes, phase shifts and large-scale degradation of a Caribbean coral reef,” Science vol. 265, pp. 1547-1551, 1994. [7] J.B.C. Jackson, “Reefs since Columbus,” Proc. 8th Int. Coral Reef Symp., vol. 1, pp. 97-106, 1997. [8] T.R. McClanahan, “Dynamics of Drupella cornus populations on Kenyan coral reefs,” Proc. 8th Int. Coral Reef Symp., vol. 1, pp. 633-638, 1997. [9] S.M. Lewis, “The role of herbivorous fishes in the organization of a Caribbean reef community,” Ecol. Monogr., vol. 56, pp. 183-200, 1986. [10] T.R. McClanahan, “Kenyan coral reef lagoon fish: effects of fishing, substrate complexity and sea urchins,” Coral Reefs vol. 13, pp. 231-241, 1994. [11] A.M. Szmant, “Nutrient effects on coral reefs: A hypothesis on the importance of topographic and trophic complexity to reef nutrient dynamics,” Proc. 8th Int. Coral Reef Symp. 2: 1527-1532, 1997. [12] A.W. Bruckner, R.J. Bruckner, and E.H. Williams Jr., “Spread of black-band disease epizootic through the coral reef system in St. Ann’s Bay, Jamaica,” Bull. Mar. Sci., vol. 61, pp. 919-928, 1997. [13] J.D. Woodley, E.A. Chornesky, P.A. Clifford, J.B.C. Pearson, J.W. Porter, M.C. Rooney, K.W. Rylaarsdam, V.J. Tunnicliffe, C.M. Wahle, J.L. Wulff, A.S.C. Curtis, M.D. Dallmeyer, B.P. Jupp, M.A.R. Koehle, J. Niegel and E.M. Sides, “Hurricane Allen’s Impact on Jamaican Coral Reefs,” Science vol. 241(4522), pp. 749-755, 1981. [14] J.D. Woodley, “The effects of Hurricane Gilbert on Coral reefs At Discovery Bay,” Appendix 9, in: UNEP Regional Seas Reports and Studies, No. 110, 1981. [15] N. Knowlton, J.C. Lang, M.C. Rooney and P. Clifford, “Evidence for delayed mortality in hurricane damaged Jamaican staghorn corals,” Nature,

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