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.
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
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

Page 1
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?
Page 2
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
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.
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
Page 3
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. 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
with rubble. The A-frames at Blue Hole are located in a The cultures were monitored for both growth and
back reef, lagoon environment with slightly turbid water survivorship. Growth rates and survivorship was
and mixed sand, rubble substrate. The Canoe Channel determined as in the previous experiment.
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.
Fig. 4. Acropora cervicornis attached to nylon
3) Cement Base Experiment monofilament line in line restoration experiment.
A 50/50 cement- sand mixture was flattened, pressed
and dried into the shape of a ~ 5 cm circular disk. Two III. RESULTS
holes were punctured through the disk before drying. A
nylon monofilament line would then be looped through A. Coral Recruitment
the holes to secure small (< 7 cm) fragments of A. Over a 30 month period only seven acroporid spat
cervicornis and A. prolifera. Two hundred and eleven settled – 3 in summer 2001 (16 spat m-2) and 4 in
disks were constructed. Disks were secured to a wire summer 2003 (23 spat m-2) (Fig. 5). No Acropora
mesh sheet with the fishing (Fig. 3). The fragments recruitment occurred at 14 m, 19 m, 26 m or 33 m.
were monitored for survivorship and growth. 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%).

10000

Acropora Others
1000
Fig 3. Coral framents on cement disks on wire mats (left) and
-2

on A-frames (right) in transit from construction site to reef site.

log scale
Spat m

100

4) Line Experiment
On June 16, 2004, sixteen A. cervicornis fragments 10
were collected from colonies at Dairy Bull Reef. After
~ 4 hrs in the shaded buckets, the fragments were
1
transported to the Canoe Channel, where they were
attached to monofilament nylon lines to create a set of ' 01 '02 ' 02 ' 03 '0 3
line cultured corals suspended underneath the mesh of m m er t e r '0 1 - m mer er '02 - ummer
S u Win S u t S
the underwater culture tables. Win
On September 3, 2004, the experiment was repeated
with 40 fragments also collected at Dairy Bull Reef. Fig. 5. Density of recruited Acropora spat
This time the fragments were deployed on lines in the normalized to m-2 over time.
Canoe Channel within 30 min of being collected (Fig. 4).

Page 4
 Acropora Others
180
1000 160
140

% Net Growth
120
-2

100
log scale
Spat m

100 CP
EBR
80
BH
10 60 CC
40
20
1
0
3m

9m

m
14

19

26

33
Jul. 2 Aug.20 Nov. 17 Feb. 28
Time Post transplantation
Fig. 6. Density of recruited Acropora spat normalized
to m-2 over depth. Log scale (n + 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).
B. Coral Restoration
1) Sampling Effect Experiment. 3) Cement Disk Experiment
Of the 40 tagged colonies in the source populationn Initially the survival and growth of the colonies was
where fragments were collected, only 19 were found on very good during the first couple of months. Colonies
after 3 months. Of these 19 source colonies, six were quickly grew and established multiple growing branches
dead (32% mortality) and 13 were alive (68% survival) (Fig. 9). The survivorship after 37 weeks was 19% - 40
with an apical polyp. After 3 months 32 control colonies of the original 211 fragments were still alive.
colonies were located. Eight colonies were dead (25% Many of the colonies were affected by Huricane Ivan
mortality) and 24 colonies were alive (75% survival). A which passed south of Jamaica in September 2004.
survey of the Acropora cervicornis population at Dairy After one year (June 2005) few complete colonies are
Bull Reef found a total of 106 dead tips on colonies alive. Several colonies have live areas, but no apical poly.
otherwise alive. This mortality is likely the result from Based on eight weeks of observations the annulaized
fireworm predation and damselfish bites and is rather growth rate was 23 cm year -1.
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. 9. Measuring growth on Acropora cervicornis
100%
fragment transplanted on cement disk.
80%
4) Line Experiment
60% Within two days of the start of the experiment most
% Survival

of coral had bleached. After five days we observed

40%
100% bleaching and the start of algal growth over the
20% white tissue. None of the colonies recovered. Five days
after the September transplantation, none of the corals
0%
1 6 11 16 21 26 31 36 41 46 51 56
had any bleaching inspite of water temperatures being at
Columbus Park
Weeks
East Back Reef Blue Hole Canoe Channel
their annual maximums [36]. After 26 weeks the
survivorship of the second attempt had a 32% survival.
Fig. 7. % survival of fragments over time. Weeks on x axis.
Considering that Hurricane Ivan passed two weeks into

Page 5
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
Page 6
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
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, high-
lighting 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.
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Page 7
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