Coral Reef Health in the
Survey and Recommendation
By Thomas J. Goreau, Ph.D., President, Global Coral Reef Alliance
Protecting Jamaica's Coral Reefs, 1991
Final Report Of The Negril Reef Mooring Buoy Workshop & Installation Project
Reef Relief and the Negril Coral Reef Preservation Society "for our children's children"
NOTE: This paper describes the environmental situation of coral reefs in the Negril area of western Jamaica during 1991,and contrasts it with previous studies going back to 1960. It documents the extent of deterioration in that interval, assesses the likely causes, and makes recommendations for steps to reverse the deterioration. Since it was written, all sites described have deteriorated significantly. Detailed lists of coral and algae species for the area are given, however there has since been a large-scale proliferation of many algae species that were rare or not present in 1991.
Surveys of reefs in the Negril area during November 1991 found that corals throughout the Bay were under moderate to severe stress from algal, sponge, and soft coral overgrowth, sediments, nutrients, coral bleaching, and boat and diver damage. The extent of algal overgrowth markedly accelerated in the previous five years, while coral bleaching had severely affected the growth rate of many or most corals. Many corals may die within a few years unless current threats to the reef are abated. Probably the most urgent step required is reduction of excessive nutrient inputs to the Bay from rivers, canals, and sewage outfalls. Tertiary wastewater treatment to remove phosphorus and nitrogen will be necessary to halt excessive algal growth which is smothering most of the reef. Environmental monitoring, expanded education of reef users, establishment of protected areas, control of over harvesting, and implementation of local, national, regional, and international policies which abate external threats, will all be needed to protect Negril reefs.
PREVIOUS WORK ON ECOLOGICAL STATUS OF NEGRIL REEFS
The first ecological survey of the Negril area, conducted from the Lighthouse to Green Island by the late T. F. Goreau, is still the largest ranging study to date. It established the baseline for all subsequent changes because it took place just after the road to the beach was built, canals began draining the Negril Morass, and before any tourist development in Negril. The entire 7 mile beach area was unpopulated: during one entire day in 1960 I saw only two other people, walking from Green Island with a wheelbarrow to collect coconuts from the Jamaica Tall palms that then covered the entire beach strand. Shallow beach waters had clean white sand entirely barren of algae. Most reefs were highly diverse and were in good condition. For background on the natural marine ecosystems of the Negril area readers are referred to Appendix 1, which summarizes the 1960 ecological evaluation and the management and conservation recommendations. Recommendations for protection of Negrilís unique marine and terrestrial habitats were made at that time to the Negril Area Land Authority and to the Beach Control Authority, urging that designated parts of the area be the third national marine park, after those proposed to the Government for Ocho Rios and Montego Bay in the mid 1950s. All recommendations appear to have been ignored, except a marine park in Montego Bay that took about 35 years to implement. Had they been followed, much of the current environmental damage might have been avoided.
2. Petroleum Corporation of Jamaica, 1980 - 1983
During the Petroleum Corporation of Jamaica's studies of proposals to excavate and burn the Negril Morass peat, environmental studies of the area were made. Chemical analyses of canal and river waters draining the morass, upstream from Negril resort area sewage inputs, showed high phosphorus and nitrogen concentrations, largely in organic forms. Enell concluded these elements were sufficiently abundant that they were not likely to limit productivity in those fresh waters. To determine if drainage of the Morass through man-made canals after 1960 had affected coral growth, 7 corals growing at 30 feet depth were cored by underwater drilling apparatus by Hendry et al. Annual growth rates determined from X-Ray studies of skeleton bands showed a very small, but clear, decrease in growth rates from 1960 to 1980 compared to those from 1940 to 1960.
3. T. J. Goreau et al., 1986
In 1986 T. J. Goreau et al. measured growth rates of corals growing at 10 feet depth at sites near to and away from fresh water inputs from the South Negril River and the North Negril Canal, and at undisturbed sites in Samuels Bay. Growth records studied spanned 1970 to 1985. Results showed most corals had significantly decreased their growth rates over this period, with the fastest growing corals being most affected. The slowest growing corals, which grew nearest to fresh water inputs, showed least change. Experiments by Dallmeyer et al. on Negril corals showed their growth was depressed by river-borne peat sediment and brown water which fouled coral tissues and reduced light for photosynthesis by their symbiotic algae. The increase in suspended peat resulting from increased drainage of the Morass for hotel construction and for slash and bum agriculture in the Morass is probably responsible for the overall decline in coral growth rates observed.
Enriched nutrient levels in fresh water, derived from peat decomposition and sewage, may have slightly stimulated growth rates of one coral which was growing (very slowly) in the canal mouth surrounded by peat sediment. Almost all large coral heads formerly growing in this area had been overgrown by fleshy (macrophytic) algae. Algae appeared to be greatly stimulated by river-borne nutrients despite turbidity-reduced light levels. Huge waving mats of algae were growing over peat sediments and over large dead coral heads near the river. These algae were exceptionally richly coloured and lushly developed, showing no sign of herbivory by fish. Species were sharply zoned in concentric rings around the river mouth. Algal biomass and coral overgrowth decreased sharply away from river and canal mouths, from over 90% to around 5 - 10% in half a mile. Sediments and peat water appeared to be generally depressing coral growth throughout the area. Reddish peat sediments were extremely light, poorly consolidated, and easily resuspended and transported by waves. In contrast, nutrients can be almost completely removed from the water by algal blooms.
4. ETH group, 1990.
In 1990 geologists from the Eidgenossische Technische Hochshule, Zurich, Switzerland, dived extensively at Negril reef sites they had visited 5 years before. They were "shocked to see the deteriorating conditions of the reef all along this sector of the coast", and reported they had "never seen a reef in such critical condition" despite experience in many parts of the Caribbean and Indian Ocean. They noted that, other than the large amount of dead coral and small size of fish, "one of the most striking symptoms of degradation that we observed is the enormous increase in algae growing on the corals and other reef-building organisms, such as sponges". They suggested that continuation of these trends would destroy the reef community within 5 to 10 years, and urged restrictions on fishing as a partial solution.
5. A. Greenaway, 1990
In 1990 Greenaway took water samples in nearshore areas from Bloody Bay to near the lighthouse. These were analyzed for nitrogen and phosphorus (as inorganic and total dissolved forms), and for fecal coliform bacteria. Elevated of coliform bacteria and nitrate (measured by the low-sensitivity brucine technique) were reported primarily near the South Negril River and NCB Plaza. He concluded that except for areas near South Negril River and the UDC sewage plant at Bloody Bay, pollution did not appear to be a problem, because they were low compared to the Kingston area. Sewage pollution has been so intense near Kingston that the Hellshire reefs were mostly killed in the 1950s and 1960s by algal overgrowth. Nitrate levels were well above open ocean values, similar to values from waters from North Coast reefs undergoing serious algal overgrowth (conditions much more similar to Negril), and in the range experimentally shown by B. Lapointe et al. to stimulate growth of many Jamaican seaweeds. The Negril data are therefore consistent, despite their author's conclusions, with nutrient overloading. Nutrients were measured only downstream of sewage inputs but the earlier PCJ study measured only upstream locations, and different methods were used, so results are not readily comparable. Simultaneous measurements should be made at locations sampled in both studies to determine how much nutrients enter the bay, and how much is derived from sewage, from swamp drainage, or from inputs from the surrounding watershed.
PRESENT STATUS OF NEGRIL REEFS
T. J. Goreau. November 1991
The massive increase of tourism in the Negril area has enormously elevated environmental stresses from boat traffic, diver and anchor damage, nutrients, and seaweeds. Anchor and diver damage was moderate to severe at the most heavily used dive sites. Many corals had gouges and scraping damage from anchors, ropes, and chains around basal portions and outer extremities. Many large coral heads had been overturned and fragmented. Some tall heads with relatively narrow bases may have been pulled over by boats using engines to back up to free snagged anchors, which can be common during rough weather unless great care is taken to anchor only in sandy areas.
Dramatic changes have taken place in the Negril area in the past five years. All regular divers reported marked recent deterioration of Negril reefs, with the last year or two being worst. Extensive overgrowth of corals by algae was reported in areas where this was previously never seen. In 1991 huge mats of fluffy green algae smothered large areas next to beaches and over seagrass beds, and long strands of algae covered corals in shallow reefs. After months of clear, hot weather, a severe Norther caused very heavy wave activity which swept all loose algal mats out to sea about one week before the study was made.
Mass coral bleaching, previously unknown, took place in Jamaica in four of the past five years, 1987, 1989, 1990, and 1991. 1990 and 1989 were the worst in Jamaica and Cayman, with 1987 and 1991 somewhat milder. These patterns are similar to sea surface temperature patterns seen in satellite data. Bleached corals cease growth for many months until they die or recover. Hurricane Gilbert, the strongest on record in the Atlantic, caused physical damage in 1988, the coolest year between 1987 and 1991.
In response to these concerns, a rapid ecological survey of Negril area reefs was undertaken during the Negril Coral Reef Preservation Society / Reef Relief Mooring Buoy Workshop. There was very limited time available, so ecological observations focused on corals and algae, with less attention paid to sponges, seagrasses, soft corals, other invertebrates, or fish. Unfortunately time did not permit surveying reefs north of Negril Bay. Areas examined in this and previous studies are listed below. Algae were collected at all sites except a and c, and drill cores of underlying substrate were taken at sites c, e, and g.
AREAS EXAMINED IN THIS AND PREVIOUS STUDIES
LOCATION 1960 1986 1991
South Side - - +
S.W. Cliffs. caves + - +
Negril River, south shore - + +
Outer Reef + - +
Long Bay + - +
Orange River canal + + -
Bloody Bay + - -
SamuelsBay + + -
Green Island + - -
Brief notes on 1991 sites:
a) South Side. Several miles past lighthouse. 0 to 30 feet. This area is unaffected by drainage from the Negril Morass or tourist areas. Best coral reefs seen in survey. Coral cover on hard ground around 90%.
b) Lighthouse to S. Negril River. 0 to 15 feet. Algae sampled at eight locations along the shore from the lighthouse to the River, along an environmental gradient from exposed marine conditions to heavy freshwater, sediment, and nutrient impact. Coral cover decreases from moderate to very poor approaching the river, and has decreased by around half since 1986.
c) Shallow central snorkel reefs. 0 to 10 feet. Area heavily impacted by anchor damage, diver damage, and turbidity.
d) Deep southern reef. "Throne Room" area. 70 to 90 feet. Among best reefs in Negril Bay. Moderate advanced diver activity and boat damage.
e) Medium depth southern reefs. "Treasure Reef". 35 to 65 feet. Considerable beginner diver activity.
f) Medium depth northern reefs. "Kingfish Point". 40 to 65 feet. Moderate advanced diver activity.
g) Shallow northem reefs. South of Booby Cay. 0 to 20 feet. Heavy beginner diver and snorkeler activity. Severe damage to reefs.
Brief notes on Negril reef species:
Comparing 1960 and 1991 surveys of coral abundance (Appendix 2) showed the most marked changes were:
1) The almost complete elimination of the two most common shallow water branching coral species throughout the area,
2) severe bleaching impact to the most abundant deeper water coral species,
3) overgrowing of all corals, bleached or not, by other organisms,
4) marked decrease of coral cover at inshore sites.
At all exposed shallow sites it was clear that the predominant coral species had until recent years been Acropora palmata (staghorn) and Acropora cervicornis (elkhorn) corals. Eroded and overgrown stumps of the former were abundant and their skeletal fragments dominated rubble piles. Some of this destruction was due to hurricanes, particularly Gilbert. South of Booby Cay most of a previously magnificent staghorn reef framework was intact, but almost no live coral was seen. Local observers noted this had been the most heavily used snorkeling reef, causing severe anchor and diver touching damage to corals. They also reported that large groups had been encouraged to stand on top of living coral to pose for photographs. Protected shallow sites had been dominated by large head corals but these were largely dead.
Completely bleached, partially bleached, or unusually coloured coral colonies were seen in almost every species (Appendix 2). Although only few corals were bleached fully white, about a quarter to one third of all corals seen were partially bleached to abnormal colours. Most corals were in recovery phases seen towards the end of a bleaching event. The fire corals, which are among the first to bleach completely white, and the first to recover, showed only minor bleaching in recovery phases. One of the major reef building corals, Montastrea annularis, showed only relatively minor bleaching (10 to 20%), almost entirely partial, and also appearing to be in recovery. Normally this species is one of the slowest to recover. Other species which usually suffer prolonged and complete bleaching, such as the Agaricias, showed partial bleaching in parallel white stripes, especially in deeper water species. Orange-red coloured colonies of Montastrea annuularis, Agaricia agaricites, Mycetophyllia, and other species with similar colours did not appear to be affected by bleaching. The unusual colour could come from pigments in a unique symbiotic algal variety with high resistance to bleaching stresses, which may be locally-adapted to absorb light wavelengths which are not blocked by peat-stained surface waters.
Severe bleaching was noted in three of the most common coral species in the Negril area. About 80% of the Montastrea cavemosa were bleached, but largely to ashy-grey colours from normal brown, green, and reddish colours. Some colonies were bleached white between polyps, with a ring of pigment only around the oral tentacular disk. Around 90% of the Siderastrea siderea were affected: around 80% were bleached uniformly indigo blue, characteristic of moderate bleaching in this species. around 10% were normal dark brown, and around 10% (mostly on South Side and deeper reefs) had measles-like polka-dots on a dark background typical of early bleaching and late recovery. Montastrea annularis was known to have markedly slowed growth rates in Negril even before 1986, the last year of normal growth before Caribbean mass bleaching began. Bleaching has probably stopped growth of many, perhaps most, colonies of this and many other corals for long periods in the last five years.
Bleached coral colonies were most abundant in the least disturbed sites to the South (around 40%), and declined from deeper offshore reef towards sites near freshwater sources of turbidity and nutrients (around 10 - 20%). Corals normally subject to higher pollution stress being less affected by bleaching than those from more pristine areas rules out local pollution as a major cause of bleaching. It is in accord with temperature as a cause of bleaching, since light and temperature stress are reduced in polluted sites due to light absorption in reddish-brown brackish surface waters.
Much bleaching might not be easily recognized since the most obvious white phase of bleaching was relatively uncommon. Dive operators nevertheless identified abnormally coloured coral as "looking sick". The same bleaching patterns were reported in Grand Cayman during 1991 by Phillipe Bush of the Cayman Government Natural Resources Unit and by Raymond L. Hayes of Howard University, in Belize, and in Bermuda by Clayton Cook of the Bermuda Biological Station, but bleaching was more severe in Florida. Satellite-derived sea surface temperature data shows notably warmer waters in the Florida Keys than in Jamaica and Cayman in 1991. Satellite maps indicate that the mid November storm rains and cloudiness cooled Negril waters below 30 degrees celsius. A large fraction of the corals in the Negril area are likely to have largely stopped growth during four of the last fve years. This includes at least a third of all corals seen in November. This underestimates total bleaching since many corals may have already recovered by then or bleached during more intense episodes in 1987, 1989, and 1990. In their starved condition, such corals are barely surviving, and lack the energy needed to keep surfaces dean of suspended sediment or maintain their space from being overgrown.
Coral species diversity was still very high at the better sites and included almost all Caribbean species. Wave adapted and sediment adapted species dominated shallow areas near rocky shores and rivers respectively. There were unusually high abundances of Montastrea cavernosa and of many less common species such as Mycetophyllia. A key indicator of coral reef health is the ability of corals to occupy space and grow upwards. Coral cover declined inshore and towards the river, from 80% or higher coral cover at deeper sites to 10% or less of hard bottom areas in shallow inshore areas. At all sites algae, sponge, and soft coral diversity and biomass was very high. Although some corals appeared healthy and holding their own, a very large proportion were dead or being overgrown by other organisms, especially at shallower sites. Healthy reefs have 90% or more of hard surface covered by living stony corals, but unhealthy reefs are dominated by algae, sponges, or soft corals. By these standards South Side reefs were healthy, deeper reefs were mildly ill, and inshore reefs were sick, terminal, or dead.
Algal biomass was extremely high at all sites, especially inshore. Algae were extensively overgrowing coral heads at all sites except the South Side. Because extremely severe storm and heavy rains caused record floods the week before, algal biomass had only recently been much higher than was seen in this survey. The most abundant living fleshy algae were identified at all sites (Appendix 3).
Local divers reported that algae were extensively overgrowing corals in the Negril area. In shallow water on the South side of Bloody Bay, near the UDC sewage plant, mats of a fine, red, "mossy" algae were reported spreading. Unfortunately I did not have time to dive there personally, but a diver familiar with the area confirmed its identity as a Ceramium species from specimens collected earlier on the South Side of Negril Bay. Ceramium is an indicator of high nutrient levels, and is overgrowing shallow areas and reef crest sites on the North Coast of Jamaica near elevated nitrogen inputs.
The worst algal overgrowth was in shallow waters all along Negril beach, over seagrasses beds in the Bay, and all over the shallow reefs. Huge masses of a fine, green, "mossy" algae smothered large areas for months until swept away by the storm. So completely was this removed that no live specimens were observed during a week of diving or during extensive glass bottom boat observations in all parts of the Bay. Fragments found in sand channels and washed up on the beach confirm the alga was Chaetomorpha linum, a major cause of overgrowth of back reef areas on the North Coast of Jamaica. It has been shown experimentally to respond strongly to elevated nutrient levels and grows in nitrate-enriched Jamaican coastal waters at the fastest rates measured. It is an indicator species for eutrophication, in particular for phosphorus pollution, since it takes up exceptional amounts of this element. It has severe impacts because masses of loose strands, up to 15 feet long, wrap around coral heads, sea grasses, and algae, blocking water circulation and light.
Despite near complete removal of the worst algal overgrowth by the storm, reefs at all sites were still observed to be undergoing severe overgrowth by surviving algae. Algal biomasses were high at all sites, although problem species were not abundant at the most pristine sites. Most overgrowth in deeper reefs was caused by Sargassum hystrix, Lobophora variegata, Dictyota cervicornis, and Dictyota jamaicensis. They are among those most responsible for algal overgrowth of offshore reefs along the North Coast. These species, especially tightly-attached Lobophora, dominated algal biomass piled in deep reef sand channels after the storm. The large increase of these algae could be due to a combination of reduced herbivory and increased nutrients. Because fish avoid eating these species due to their distasteful compounds, disease-induced die off of black sea urchins (Diadema antillarum) in 1983 has allowed greatly increased biomass, as it was their only common consumer. Reduced herbivory is likely to be responsible for algal abundance on reefs free of freshwater, sediment, or nutrient inputs.
Heavy overfishing and urchin die off had their major impacts by 1983, so proliferation of algae in the Bay since 1986 is not likely to be caused primarily by herbivore reduction. Chemical measurements on North Coast algae by Lapointe et al. show growth is stimulated by land derived nitrate even at the concentrations below 2 micromoles per litre which were found in Negril coastal waters by Greenaway. Although Negril has all characteristics of coral reefs in medium stages of overgrowth by algae stimulated by land derived nutrients, many species typical of late stages of algal overgrowth of reefs were found in only small amounts or were not present (appendix 3). This suggests algal overgrowth is still fairly moderate, and more easily reversible, than reefs near Kingston, Montego Bay, or Ocho Rios with much larger sewage inputs. However, current trends should cause most Negril Bay reefs to be killed by algal overgrowth within a few years even if there is no further boat damage or bleaching. Rapid reduction of external nutrients released into the Bay is essential to protect them.
Seagrass areas were examined by several complete north-south and east-west transects of Negril Bay by glass bottom boat. Seagrasses did not appear healthy. Plants were surprisingly small, pale, and sparsely distributed for shallow and clear waters. They appeared to have been considerably set back by shading from dense green algal mat coverage during the summer and by chronic river-borne turbidity.
Sponge biomass and diversity was exceptionally high, accounting for much of the beauty of the reefs. Most sponges described in Caribbean field guides were present. Large encrusting sponges were overgrowing dead reef rock and sediment-impacted areas near the river mouth, and were overgrowing living corals in the reefs. Corals were losing space to sponges wherever they contacted, except for a single species of sponge symbiotic with corals. Most sponges appeared extremely healthy. Sponges are strongly favored by high levels of organic material floating in the water, which promotes growth of the bacteria on which they feed, making many species excellent indicators of pollution.
5. Soft corals
No effort was made to tabulate soft coral species, such as gorgonians, antipatharians, zoanthids, anemones, hydrozoans, and coralliomorphs. Their biomass and diversity were high, and they were also responsible for significant coral overgrowth. Most of the Caribbean species of these groups were present.
6. Other invertebrates
No effort was made to document invertebrates other than stony corals. High biomass of starfish, sand dollars, sea urchins, and sea cucumbers was noted. This indicates that collection of marine organisms has been successfully controlled in Negril by existing diver education programs, as these animals are easily collected and are only slowly replaced. In this regard the ecosystem was far more healthy than many other parts of Jamaica, such as Discovery Bay where collection of such organisms for visiting lab courses has severely depleted them. Diadema were present in small numbers in only a few shallow sites. Their near absence has probably been a cause of extremely high abundances of seaweeds which contain anti-fish chemicals, such as Lobophora, Sargassum, and Dictyota. These algae were the major coral over growers on the outer reefs.
No effort was made to document fish species. Fish biomass and diversity were generally very low. Low herbivory, caused by overfishing, may account for very high levels of certain algae which are normally strongly controlled by fish grazing, such as Halimeda and calcareous species normally consumed by Parrotfish, or Chaetomorpha, Acanthophora, Bryothamnium, and other species normally consumed by Doctorfish.
1. Ecological training of Negril Reef community
The mooring buoys installed in November 1991 by REEF RELIEF and the Negril Coral Reef Preservation Society will serve as the centre of diving activity in the Negril area. Dive groups using these mooring sites should document changes in the local reef community in order to determine if reef health improves or continues to deteriorate. Corals in those areas which have been severely damaged by anchors and chains should now gradually recover over coming decades, but they will face severe problems due to algae, sponge, and soft coral overgrowth, and from deteriorating conditions due to turbid water, sedimentation, and excessive nutrients. These will get worse unless sewage is tertiary treated and watershed erosion halted.
A workshop should be held to provide the Negril diving community instruction on the ecology of major local species, so divers can recognize normal and unhealthy appearances and identify species which indicate stressed environmental conditions. A visual species key should be prepared, including the range of bleaching characteristics and environmental preferences, to allow dive operators to identify and document coral health. Most dive instructors already know more kinds of fish than the number of major indicator coral and algae species they might need to learn. Training should be held as soon as possible to establish a locally-based monitoring program. The best time for such a workshop would be during the tourist off-season. Because of the importance of comparing bleached to normal appearances of corals it should be held during the warmest time of year, September or October.
Changes can be documented using dated photographs containing suitable scales, and by simple monitoring methods. The entire Negril diving community should be interviewed in order to record observations of past changes, especially persons swimming regularly at the same sites for many years. This should be done right after the training workshop on local species. A network should be established under the Negril Coral Reef Preservation Society to photographically document changes of species abundances in a form allowing compilation, comparison, and identification of species from photographs or samples.
2. Coral monitoring
Fundamental measures of coral reef health include coral cover, coral growth rate, degree of coral overgrowth, and extent of bleaching. These need to be followed because Negril corals were already slowing down growth markedly between 1970 and 1986, probably because of sediment stress, before mass bleaching even began, and before algae had become a widespread problem. Repeated photographic monitoring of selected sites could allow changes in bottom cover, growth rates, and bleaching to be determined from dated photographs with a length scale. A measuring rod with a colour chart should be used so changes can be evaluated under different light and turbidity conditions. Corals near to, as well as away from, buoy mooring lines should be selected for long term growth monitoring by photographic methods. Monitoring should also be done at selected undisturbed sites to the south and to the north of Negril.
At these selected sites all coral, algae, and other major species should, and overall species abundance patterns in surrounding areas recorded on long transects using underwater writing slates with species lists analogous to those of Appendices 2 and 3. A few drill cores should be taken from coral heads at sites near land inputs to document changes in growth rates since 1986 from coral growth patterns. These can be obtained using much thinner cores than the mooring installations, and can be extracted without permanent damage to the coral using the latest methods.
The collection of bottom drill cores taken during mooring installation should be catalogued and located on a detailed map. These cores can be used to document past evolution of the reef framework. Initial inspection of the first 15 cores taken in November 1991 showed a wide range of hard bottom types, including a) poorly cemented coral reef sands, (perhaps only decades old) b) lithified seagrass beds (possibly centuries old) c) recently killed coral (probably only a few years old), d) fossil Holocene coral (younger than 5,000 years), e) fossil Pleistocene coral (about 130,000 years old), f) Ice Age soil deposits (between 100,000 to 5,000 years old), and g) recent influxes of red sediment (probably Morass-derived and postdating 1960). Some areas were rapidly growing upwards, but others had been strongly eroded. These cores should be described scientifically, and if possible fossil material should be radiocarbon dated, so that the overall past growth of the Negril reef framework can be determined. It is clear from initial inspection of these samples that the history of Negril reefs has been extremely dynamic and varied, and further study of these ancient historical records could help evaluate how much current conditions have changed from those in the past.
3. Algae monitoring
Algae and algal fragments were virtually absent except on wave-beaten shores in 1960. In 1986 they were highest near river and canal mouths. By 1991 they were omnipresent all over the Bay. The rapid and recent expansion of algae requires careful watching. Algal cover should be monitored photographically at selected sites, and a watch kept for species which are indicators of nutrient pollution or herbivores. Some can spread very rapidly under conditions of nutrient or temperature stress. Because local algae are often quite distinctive in appearance from those recorded in scientific guides, a photographic key should be prepared.
Algae which were collected in November 1991 should be analyzed for carbon, nitrogen, and phosphorus contents. High ratios of nitrogen and phosporus to carbon in algal tissue indicates eutrophication, the stimulation of algae by excessive nutrients. Higher values in certain areas can be used to identify source regions of excess nutrients. Nitrogen to phosphorus ratios indicate which of the two nutrients is in greatest excess. This is important for designing control strategies since reduction of the limiting element does most to abate eutrophication. Monitoring should include sampling of indicator species for analysis of nitrogen and phosphorus contents. Nutrients can be rapidly removed from the water column by proliferating algal growth under clear sunny conditions, and nutrient concentrations in water are highly variable depending on waves, tides, and winds. Nitrogen and phosporus contents of seaweeds are measures of long term nutrient exposure at each site, and a more efficient way to document ecosystem impacts of eutrophication than rapidly fluctuating values in the water that require much larger numbers of measurements to characterize.
4. Sponge monitoring
Sponge diversity and biomass at Negril are exceptional. Sponges are actively overgrowing corals and appear to be stimulated by high levels of bacteria and organic particulate materials which they filter out of the water. A photographic sponge key should be made, potential pollution indicator species identified, and sponge cover changes documented at certain sites. Overgrowth by fire coral, soft corals, anemones, and other groups is a real, but lesser, problem, and should also be monitored.
5. Environmental monitoring
An effort should be made to establish Negril as a site for long term physical monitoring, including water temperature, salinity, turbidity, and meteorological variables, so environmental effects of episodic weather events and global changes can be distinguished from human stresses. Nutrient and pollution inputs to the water need to be documented and their sources characterized for effective coastal zone management. The incidence of mass bleaching in recent years following extended periods of sunny, calm, hot weather indicates that the Negril area is undergoing temperature stress.
Turbidity at Negril also requires monitoring. While exacerbated at the time of the study by the recent storm, it appears to be a chronic problem. High abundances of sponges and soft corals are characteristic of reefs undergoing sediment stress. Levels of turbidity were extremely high throughout the Bay although sufficient time had transpired for most resuspended limestone or mineral sediment particles to settle. Turbidity is largely caused by suspended peat particles and humus-stained swamp water which covers large parts of the bay at different times depending on currents and winds. Offshore reefs had high levels of flocculent suspended particulate matter which settled very slowly, if at all. This organic material is decomposed by bacteria and fungi, providing the food supply for the detritus-ingesting sponge and invertebrate community which is replacing corals. Changing turbidity has strong consequences for the future of the reef community since most coral species cannot tolerate excessive levels. As reef building corals are replaced by species which are unable to build wave resistant frameworks, shore line protection from hurricane waves will deteriorate irreversibly. More intense hurricanes are predicted if global warming continues.
It was noted that waters offshore from obviously brown-stained surface waters were always markedly green in colour, rather than the normal blue colour of Jamaican reef and ocean waters. Large phytoplankton blooms were probably taking place, probably caused by nutrient inputs from fresh water. Particulate matter, dissolved turbidity, and phytoplankton should therefore be monitored along with temperature, salinity, and nutrients.
6. Tertiary sewage treatment
Abatement of anchor and diver damage by proper use of moorings will not be enough to save the reefs unless excessive algal overgrowth of corals is curbed by reducing nutrient inputs to the reefs. This requires tertiary wastewater treatment to remove nutrients from sewage. Otherwise the reef will be eventually be killed by algal overgrowth, as has happened to reefs near Kingston, Montego Bay, Ocho Rios, St. Anns Bay, and other major sewage disposal sites around Jamaica There are many simple methods of biological tertiary treatment which could readily be applied in the Negril area to strip excess nutrients from waste waters. Some examples include use of artificial wetlands, algal mariculture, and fish or shellfish ponds. Pilot scale tests of several of the most viable technologies should be implemented in Negril in order to determine which are most cost effective under local conditions.
Funding for the installation of the new sewage treatment plant in the next few years should include development and implementation of tertiary wastewater treatment. Current plans apparently include secondary treatment, plus potential use of chemical phosphorus removal if loading from phosphate detergents is found too high. Halting use of phosphate detergents would help immediately, but high levels of phosphorus and nitrogen in garbage and excrement would still require removal. Most methods of chemical phosphorus removal generate solid and liquid sludge wastes to be disposed and remove only very little nitrogen. Bioremediation, growing organisms in waste water ponds to absorb nutrients, should remove both elements more cheaply than chemical treatment and possibly generate economic products from biomass and bioenergy. Nutrient monitoring should evaluate performance of sewage control measures and identify possible non-sewage nutrient inputs from the Morass or surrounding watershed areas.
7. Reef restoration and sustainable management
If external environmental threats to the Negril reef, such as sewage, sediment, and global warming, can be abated, these reefs may be prime candidates for reef restoration. One potential method for restoring damaged areas is artificial reefs made by coral transplantation onto limestone structures accreted underwater using electricity from solar or wind sources. They could greatly increase fish biomass, diversity, and recruitment while protecting the shore. Successful pilot work has already been done in Jamaica by Hilbertz, indicating these methods could be useful in restoring shallow reefs in Negril, particularly in the now-vanished shallow wave-breaking reefs of staghorn and elkhorn reefs.
Altematives to overfishing are needed to maintain reef species diversity. Pilot work in Jamaica by Macfarlane and Goreau has shown many Jamaican coastal waters can be used for mariculture of local seaweeds and fish at record levels of productivity. Development of these techniques to pilot scale is recommended through programs with the Negril Fisherman's Cooperative and the NCRPS. Education of local fishing communities about local species using photograph slide shows should also be initiated. Dr. Peter Goreau, who has done this on the North Coast, found most fishermen eager to learn about the organisms they have seen. Public education of fishermen is essential to manage Jamaica's reef resources sustainable by selectively propagating desired species instead of over harvesting them.
Reduction of turbidity requires effective erosion control measures in the surrounding watershed by reforesting water courses draining hilly areas above the Morass. Morass peat is being eroded because the water table was lowered by drainage canals to permit resort development on the seaward side and for slash and burn agriculture in the interior. Peat soils which are drained will either be eroded onto the reef or consumed by decomposing microorganisms right down to the water table. Florida Everglades peat soils, which are made by the same plant, Cladium jamaicense, have subsided up to 20 feet following drainage for agriculture. Altemative land management strategies are clearly needed in the Morass to build and conserve the rich organic soil instead of destroying it. The best alternatives for soil conservation are either lack of natural vegetation disturbance or adoption of techniques developed in similar swamps thousands of years ago by Indians in Central America. They used swamps for extremely productive farming on raised fields, built up from sediment and algae dredged from canals, where they grew fish and shrimp. These methods could increase production of fresh vegetables, stabilize watercourses and drainage patterns, provide alternatives to destructive practices, restore soil fertility, and reduce erosion of peat which could otherwise harm the reefs. They could be developed in parallel with efforts to test and improve bioremediation alternatives for wastewater, using upriver sites.
Preservation of the reefs at Negril and other parts of Jamaica would be advanced if they were placed within a graded framework of protected areas, as first recommended to the Jamaica Government in draft regulations for marine parks drawn up by the late Prof. T. F. Goreau in the mid 1950s. Effective protection will require more than inadequately enforced "paper parks" declared to protect commercially used tourist reefs after they have been damaged. Protected areas must include reefs that especially need conservation because of unusual species diversity, growth rates, role as fish nurseries, or beauty. Many reefs in the Negril area still fall under one or more of these categories, and should be prime candidates for protected status.
Negril reefs are subject to stress on a wide variety of scales:
1) Local stresses, like diver, boat, and anchor damage, overfishing, reef destruction by dredging, or curio collection, can only be managed through education of reef users and by policing activities in the water,
2) Regional stresses, like sewage from resorts and populated areas, solid wastes from land and passing boats, or erosion, are caused by human activities far from the reef. These require larger scale policies for their abatement. Unless these external stresses to reef health are dealt with in an integrated fashion reefs will continue to be degraded even if all deleterious activities in reef waters were stopped. International agreements to halt global warming may also be needed to abate risk of further bleaching damage.
Protected areas should be experimentally managed as fish sanctuaries. Artificial reefs could provide shelter for recruitment of juvenile fish in areas where the shallow reef framework has been severely damaged. Recent studies in the Phillipines showed establishment of a fish sanctuary increased fish catches in the surrounding reef zones to record levels. These benefits promptly ceased when poachers invaded the sanctuary, indicating the need for long term management.
Areas declared for protection should also include biologically significant areas on land, including those specified in Appendix 1, caves, parts of the Morass, and fossil reefs. In South Negril ancient fossil reefs, 130,000 years old, are beautifully preserved with their wave cut notch. Jamaica shows this ancient sea level, around 20 feet above today's, better than anywhere. Nowhere else can one see so clearly how large a rise in sea level took place the last time in the past when temperatures were around a degree celsius above today's levels, values predicted during coming decades due to global warming. As carbon dioxide in the atmosphere was then 27% below today's level, these sites provide a minimum estimate of the sea level rise that should ultimately result from global warming. They are visible evidence that potential sea level rise could become much greater than policy makers now anticipate. These sites are unexcelled for demonstrating the potential impacts of global climate change and should be protected and used for environmental education of Jamaicans, visitors, and the international community. Some of the best such locations in Jamaica are found near Negril, Discovery Bay, Rio Bueno, and other locations along the North and South Coasts. They urgently merit protection as UNESCO World Heritage Sites since many of the best preserved sites have been destroyed to provide living and commercial space.
The Negril Coral Reef Preservation Society's diver education programs have been remarkably successfull in preventing the destruction of marine species for sale as curios which has caused considerable damage near Montego Bay. However, the fact that many Jamaican reef species can be readily bought near tourist areas will continue to be a potential threat which could quickly expand to Negril unless the trade as a whole is banned. The author observed how quickly all flowering orchids were exterminated near the Discovery Bay Marine Laboratory as soon as a market for them developed. These orchids had been unaffected for decades, indeed during construction of the Laboratory all orchids in the construction area had been carefully removed from trees which had to be cut down and retied onto trees in neighbouring undisturbed areas. Within months most vanished, although located in an official wildlife preserve where signs warmed that collection of any animal or plant was banned, and where several plant species occur which are known nowhere else on earth.
Legislation to ban the curio trade, except for sustainably managed species, needs to be enforced: at present it is not taken seriously. Although Jamaica is signatory to international treaties banning the sale of endangered marine species, many of these are openly sold in duty free tourist shops in Jamaicaís international airports. Persons buying these items and bringing them home are acting in violation of international law. Immediately after this study was done, the author observed turtle shell and other endangered species (including exotic species imported from the Philippines) for sale in the largest curio store in the Sangster International Airport. The salesperson claimed selling turtle shell jewelry was legal because they were only small pieces of shell, not the whole turtle, and since they hadn't killed it themselves. Contempt for unenforced wildlife laws will remain a perpetual threat to economically attractive species in Negril until it is seriously controlled nation-wide.
Negril coral reefs still maintain much of their beauty and diversity although they are under serious stress. They can still be saved for our children's children if prompt action is taken to reduce the major stresses to the reef.
This study was only possible thanks to the kind assistance of all the members of REEF RELIEF and the Negril Coral Reef Preservation Society, who participated in the mooring installations, and because of support for travel, room, and board from Air Jamaica and Singles. Special thanks go to Katy Thacker, Craig Quirolo, DeeVon Quirolo, and Karen McCarthy for mankind the workshop so successful.
Bernasconi, S., 1990, Eidgenossische Technische Hochschule Zurich report on reef conditions in Negril, pp. 2, Negril Coral Reef Preservation Society.
Chapman, V. J., 1961, The marine algae of Jamaica, Part 1. Myxophyceae and Chlorophyceae, Institute of Jamaica, Kingston.
Chapman, V. J., 1963, The marine algae of Jamaica, Part 2. Phaeophyceae and Rhodophyceae, Institute of Jamaica, Kingston.
Colin, P., 1988, Marine Invertebrates and plants of the living reef, 2nd edition, T.F.H. Publications, Neptune City, New Jersey.
Dallmeyer, D., J. Porter, & J. Smith, 1982, Effects of particulate peat on the behaviour and physiology of the Jamaican reef building coral Montastrea annularis, Marine Biology, 68: 229233.
Enell, M., 1984, Water chemistry of the Negril and Black River Morasses, Jamaica, Petroleum Corporation of Jamaica, Kingston, pp. 166.
Goreau, S., & T. J. Goreau, 1988, Fish mariculture potential of Jamaican back-reef springs, Proc. Assoc. Island Marine Labs. Caribbean, 21: 35.
Goreau, T. F., 1959, The ecology of Jamaican reefs. I. Species composition and zonation, Ecology, 40: 67-90.
Goreau, T. F., 1960, Report on a biological survey of the offshore regions bordering upon the Negril Green Island area, Negril Area Land Authority, pp. 7.
Goreau T. F., & N. I. Goreau, 1973, The ecology of Jamaican coral reefs II. Geomorphology, zonation, and sedimentary phases. Bulletin of Marine Science, 23: 399-464.
Goreau, T. J., in press, Bleaching and reef community change in Jamaica: 1951-1991, Symposium on Long Term Dynamics of Coral Reefs, American Society of Zoologists, Atlanta, Georgia.
Goreau, T. J., & A. H. Macfarlane, 1990, Reduced growth rate of Montastrea annularis following the 1987-1988 bleaching event, Coral Reefs, 8: 211-216.
Goreau, T. J., R. E. Dodge, & P. Goreau, 1988, Decline of coral growth rates at Negril, Jamaica, Proc. Assoc. Island Marine Labs. Caribbean, 21: 43.
Goreau, T. J., R. Hayes, J. Clark, D. Basta, & C. Robertson, in press, Elevated sea surface temperatures correlate with Caribbean coral reef bleaching, in R. A. Geyer (Ed.), Geophysical and Geochemical Aspects of Global Warming, CRC Press, Boca Raton, Florida.
Goreau, T. J., B. Lapointe, J. O'Connell, P. Goreau, & A. Macfarlane, 1988, Groundwater nutrient inputs to Jamaican reefs, Proc. Assoc. Island Marine Labs. Caribbean, 21: 40.
Greenaway, A., 1991, Coastal water monitoring programme, Negril, final report, Caribbean Environmental Consulting Services, Kingston, pp. 53.
Hendry, M., J. Woodley, C. Holmes, & R. Miller, in press, Coral growth records from Negril, Jamaica: growth band timing, the effect of swamp drainage alteration and a correlation with rainfall, Coral Reefs.
Hilbertz, W., 1991, Solar-generated construction material from sea water to mitigate global warming, Building Research and Information, 19: 242-255.
Humann, P. 1982, Ocean Realm guide to reef creatures; marine invertebrates, Ocean Realm Publishing, Miami, Florida.
Humann, P., 1983, Ocean Realm guide to corals, Ocean Realm Publishing, Miami, Florida.
Kaplan, E., 1982, A field guide to coral reefs of the Caribbean and Florida, Houghton Mifflin Co., Boston, Massachusetts.
Lapointe, B., J. O'Connell, A. Macfarlane, & T. J. Goreau, 1988, Photosynthesis of algae in Jamaican back-reef spring communities, Proc. Assoc. Island Marine Labs. Caribbean, 21: 36.
Littler, D., M. Littler, K. Bucher, & J. Norris, 1989, Marine plants of the Caribbean, Smithsonian Press, Washington D.C.
Macfarlane, A., T. J. Goreau, A. Smith, P. Goreau, S. Goreau, & B. Lapointe, 1988, Algal mariculture in Jamaican back-reef springs, Proc. Assoc. Island Marine Labs. Caribbean, 21: 37.
Sefton, N., & S. Webster, 1986, Caribbean reef invertebrates, Sea Challengers Press, Monterey, California.
BIOLOGICAL SURVEY OF THE OFFSHORE REGIONS BORDERING UPON THE NEGRIL / GREEN ISLAND AREA
REPORT TO THE NEGRIL AREA LAND AUI HORITY
BY THOMAS F. GOREAU, 1960
(SUMMARY BY THOMAS J. GOREAU, 1991)
Survey of marine communities and inshore regions bordering the Negril Area Land Development Area, as defined in the Provisional Development Order of 1958, with "recommendations regarding measures to be taken for the conservation of the marine communities and objects of natural beauty which may be threatened by the current great strides being made in the economic and physical development of this part of Jamaica."
1) Lighthouse to South Negril Point
A) Rocky coast: Caves in cliff many contain "Extremely rich growth" of marine organisms, zoned according to light levels, containing organisms which are normally found only in deep water or deep inside reef caves. "These caves are places of great natural beauty, and everything should be done to preserve them as they are now."
B) Offshore: corals small and scattered, do not go much deeper than 40 feet, no real reef structure, fleshy seaweeds on limestone blocks and eroded fossil reef.
2) Long Bay or Negril Bay
A) Inshore: Beach is "certainly one of the most attractive in the West Indies". Water very clear, bottom nearly always visible from surface. Inshore strip of "barren white sand" next to beach, making first class bathing due to nearly all white coral reef limestone sand with seagrass beds, very little living coral, rocks, sea urchins, or seaweeds. "Everything should be done to preserve this beach as a National Park, and prevent its disfigurement by the erection of hotels and other buildings too close to the sea".
B) Offshore: small bank reefs in north eastern Long Bay, Booby Cay, and at entrance to Long Bay. Much dead coral, scattered broken heads of elkhorn and brain corals. Navigational charts erroneously show barrier reef. "The absence of good reefs in this area may be a drawback from the point of view of tourism. Only a very limited amount of spearfishing will be possible on the bank reefs described above, but the size of this fish reservoir is so small that overfishing is unavoidable within a very short time."
3) Bloody Bay
A) Bloody Bay Scattered coral heads, no reefs as such, no large fish, sand, sea grass, and coral gravel bottom, with clean white sand beaches free of sea grass and urchins. 'One of the most attractive areas" in the region, which "if suitably developed could be made most attractive as a sea-side resort area".
B) Little Bloody Bay: "A spot of very great beauty", "crowned by a grove of great trees overgrown with festoons of vines and lianas". The area has been marred by a building aggregates plant, whose blasting has damaged the cliffs. "It is to be hoped that this will be removed in due course, and the natural beauty of the place restored". "A determined effort should be made to preserve this place through conservation of the trees and shrubbery, and by limiting construction of houses, roads, etc. North Negril Point would make a beautiful public park overlooking the sea. It is suggested that a public park here should include not only the point but also the entire foreshore of the Bay to keep it as a natural unit."
4) Salt Creek and Orange Point
A) Salt Creek: Cliff shore with fossil reef. Building operations (canal construction for diversion of the Orange River and drainage of Negril Morass) muddied the sea water with swamp material for 300 yards from the shore. Water was far too turbid for good observations.
B) Offshore: Rocky bottom with large boulders at base of cliff. Large fish probably hide in crevices between large rocks. Typical Jamaican exposed rocky shore organisms. Thick carpet of fleshy algae (mostly Sargassum) and encrusting organisms. Very little active coral growth. Corals do not go deep, and reef development is poor.
5) Omnge Bay to Green Island
A) Orange Bay Bay water turbid, shallow, muddy. From Ireland Point to Halfmoon Bay a line of well developed patch reefs with deep, narrow, winding channels. Growth of corals is very vigorous, reef rising to near the surface. Many large corals: sides of reef have immense colonies of the massive reef-framework building coral Montastrea, tops have large branching elkhom corals.
B) Samuels Bay: "Best part of the reef. Vigorous coral growth. Richest and most varied proliferation of coral". Great variety of other organisms, including sponges not seen elsewhere in Jamaica. Few large fish, but many small ones. Reefs are protected from waves and accessible from shore. "The great biological richness and variety of plants and animals In this reef would make this area potentially very attractive for guided tours in glass bottom boats as well as for more experienced skin divers. A strong recommendation Is herewith made in favour of declaring the entire reef a Protected Area with a view to the eventual creation of a National Marine Park similar to the one now planned near Ocho Rios".
C) Halfmoon Bay: Shallow inshore regions have large numbers of marine animals. Bathing may be limited by very high densities of the black sea egg (Diadema) which can cause painful injuries to barefoot bathers.
D) Green Island: Some of the best reefs in Jamaica.
NEGRIL CORAL ABUNDANCES AND BLEACHING
CORALSPECES AUG. 1960 NOV 1991 BLEACHING
Millepora complanata* common common 2
Millepora squarrosa* not reported not seen -
Millepora alcicornis* not reported common 2
Stylaster roseus uncommon uncommon -
Stephanocoenia michelinii deeper water uncommon -
Acropora palmata abundant rare 1
Acropora cervicornis common rare 1
Acropora prolifera not reported not seen -
Madracis decactis uncommon not seen -
Madracis mirabilis common common 1
Madracis pharensis not seen not seen -
Madracis formosa not seen not seen -
Agaricia agaricitis* cornrnon common 1
Agaricia tenuifolia not reported not seen -
Agaricia undata not reported common 2
Agaricia fragilis* common common 3
Agaricia lamarcki* not reported common 3
Agaricia grahamae not reported not seen -
Helioseris cucullata common not seen -
Siderastrea siderea* common common 9
Siderastrea radians* common common 6
Porites porites* uncommon uncommon 1
Porites furcata common common 2
Porites divaricata not reported uncommon -
Porites asteroides* abundant abundant 2
Favia fragum* common common 4
Diploria strigosa* common common 3
Diploria clivosa* common common 2
Diploria labyrinthiformis* common common 1
Manicina areolata* common common 3
Colpophyllia natans* common common 2
Montastrea annularis* common common 2
Montastrea cavernosa* common very common 7
Solenastrea hyades not seen not seen -
Astrangia solitaria common not seen -
Phyllangia americana common not seen -
Oculina diffuse not seen not seen -
Oculina valenciennesii not seen not seen -
Meandrina meandrites* common common 5
Dichoecenia stokes) common common 3
Dendrogyra cylindrus* rare rare 1
Mussa angulosa common uncommon -
Isophyllia sinuosa common common 2
Isophyllastrea rigida common common 3
Mycetophyllia lamarckiana common common 2
Mycetophyllia ferox not reported not identified -
Mycetophyllia aliciae not reported not identified -
Mycetophyllia danaana not reported not identified -
Mycetophyllia reesi not reported not identified -
Eusmilia fastigiata* common common 1
Tubastrea aurea not seen not seen -
Scolymia lacera not reported uncommon 1
Scolymia cubensis not reported not seen -
Cladocora arbuscula not reported not seen -
Note: coral species as listed in T. F. Goreau, 1960, generally ammended for later species name changes. There still remains taxonomic uncertainty in certain genera, particularly Agaricia, Montastrea, and Mycetophyllia. Underlined species have either undergone largest reductions in abundance, or are the most abundant species which were highly bleached. Species marked by asterisks have been greatly affected by bleaching in Jamaica during severe episodes, and may have largely recovered at the time of the survey. Relative abundances were noted on long transects by SCUBA or snorkelling. Bleaching frequency was estimated on a 10 point scale, where - indicates the species was not seen or too rare for bleaching frequency to be meaningful, 1 indicates that at least one of those seen were bleached, and 10 indicates that all coral heads showed signs of bleaching over part or all of their surface. Most species not seen bleached were very rare or are small solitary coral species which lack symbiotic algae. Estimates are pooled based on all observations, and are probably accurate to within plus or minus 10%.
NEGRIL ALGAE, NOVEMBER 1991
Ulvaria not seen
Ulva not seen*
Enteromorpha not seen*
Anadyomene not seen*
Microdictyon boergesenii Long Bay overgrowing sponge
Polyphysa not seen
Acetabularia not seen
Bryopsis not seen*
Chaetomorpha linum serious problem in Bay*
Cladophoropsis not seen*
Derbesia fastigiata S.Booby Cay encrusting hardground
Cladophora not seen*
Caulerpa verticillata South Side
Caulerpa sertularoides abundant, South Side
Caulerpa mexicana abundant, Negril Bay
Caulerpa racemosa abundant, Bay, South Side
Caulerpa cupressoides abundant, Bay
Batophora not seen
Dasycladus not seen
Neomeris annulata South Side
Siphonocladus not seen
Ernodesmis not seen
Valonia ventricosa Bay
Dictyosphaeria cavernosa S. Negril transect*
Codium isthmocladium S. Negril, Bay*
Codium intertextum S. Negril transect
Avranvillea sp. abundant, Bay
Udotea cyathiformis abundant, Bay
Udotea flabellum abundant, Bay
Rhipocephalus not seen
Penicillus sp. abundant
Chamaedoris not seen
Cymopolia not seen
Halimeda goreauii abundant, deeper sites*
Halimeda copiosa abundant, deeper sites*
Halimeda tuna abundant, Bay*
Halimeda opuntia S. Negril transect*
Dictyota divaricata abundant, Bay*
Dictyota bartraysii S. Negril transect*
Dictyota cervicornis Bay, South Side*
Dictyota Jamaicensis abundant. Bay, South Side*
Rosenvingea not seen
Dictyopteris not seen*
Spatoglossum not seen
Dilophus S. Negril
Stypopodium zonale South Side, S. Negril, Bay
Padina sanctae-crucis all sites
Lobophora variegata serious overgrowth of corals in Bay*
Cladosiphon not seen
Hydroclathrus not seen
Colpomenia not seen
Chnoospora not seen
Cytoseira not seen
Sargassum hystrix serious overgrowth of corals in Bay*
Sargassum polyceratium S. Negril transect
Turbinaria tricostata most shallow sites, Bay, South Side
Martensia not seen
Halymenia not seen
Kallymenia not seen
Anotrichum not seen
Griffithsia not seen
Callithamnion not seen
Champia not seen
Catenella not seen
Gigartina not seen
Centroceras not seen*
Sphacelaria not seen
Ceramium sp. serious coral overgrowth in Bloody Bay*
Wrangelia argus South Side
Crouiana not seen
Heterosiphonia not seen
Spyridia not seen
Asparagopsis not seen
Dasya rigidula Bay
Eupogodon not seen
Chondria not seen*
Liagora not seen
Trichogloeopsis not seen
Dictyurus not seen
Haloplegrna not seen
Coelothrix irregularis S. Negril transect
Gelidium pusillum S. Negril
Gelidiella acerosa S. Negril*
Pterocladia arnericana S. Negril
Bostrychia not seen
Ochtodes not seen
Cryptonemia not seen
Grateloupia not seen
Hypnea not seen*
Digenia simplex S. Negril, Bay, South Side*
Laurencia intricata Bay, S. Negril, South Side*
Laurencia poitei S. Negril*
Laurencia obtusa S. Negril
Laurencia papillosa S. Negril
Heterosiphonia not seen
Chrysymenia not seen
Botryocladia not seen
Acanthophora spicifera abundant, Bay*
Bryocladia not seen
Bryothamniurn triquetrum abundant, Bay, S. Negril, S. Side*
Gracilaria not seen
Polycavernosa not seen
Caloglossa not seen
Murayella not seen
Lobosiphonia not seen
Herposiphonia not seen
Polysiphonia not seen
Arnansia not seen
Meristiella not seen
Euchema not seen
Galaxaura rnarginata S. Negril, S. Side, Bay
Scinaia not seen
Jania rubens S. Side
Arnphiroa rigida S. Side
Amphiroa fragilissima Bay
Corallina not seen
Goniolithon not seen
Neogoniolithon S. Side
Lithophyllurn not seen
Mesophyllurn not seen
Titanoderma not seen
Peyssonellia rubra deep reef
Fosliella not seen
Hildenbrandtia not seen
Porolithon not seen
Sporolithon not seenHydrolithon not seen
Note: Species listed according to Chapman and Littler et al., without correction for recent name changes. Algae are listed only to genera if not seen, and generally to species when noted. Only macrophytic (larger) algae were identified, no effort was made to identify small filamentous, turf, or microalgae. Most calcareous reds, especially encrusting species, many common browns, calcareous greens, and some genera with several similar species such as Avranvillea or Penicillus, were identified to genus rather than species. Because of the rapid nature of this survey several more species were certainly present but were not recognized, infrequent, or occur mostly in habitats which were not examined, such as very deep, shallow, or turbid water. Closer examination would probably reveal more Dictyota, Caulerpa, Penicillus, Halimeda, Auranvillea, Ceramium, calcareous reds, and other species. Underlined species were major problem species in Negril during November 1991. Algae marked with an asterisk have been important reef overgrowers in other parts of Jamaica and the Caribbean undergoing severe eutrophication or reduced herbivory .