coastal conservation project an assessment of the coral reefs of

CCOOAASSTTAALL CCOONNSSEERRVVAATTIIOONN PPRROOJJEECCTT

AANN AASSSSEESSSSMMEENNTT OOFF TTHHEE

CCOORRAALL RREEEEFFSS OOFF

TTOOBBAAGGOO

Prepared by

Kahlil Hassanali

September 2009

i

INSTITUTE OF MARINE AFFAIRS

RESEARCH REPORT

CCOOAASSTTAALL CCOONNSSEERRVVAATTIIOONN PPRROOJJEECCTT

AANN AASSSSEESSSSMMEENNTT OOFF TTHHEE

CCOORRAALL RREEEEFFSS OOFF

TTOOBBAAGGOO

Prepared by

Kahlil Hassanali

Jr. Research Officer

September 2009

Institute of Marine Affairs

Hilltop Lane, Chaguaramas

P.O. Box 3160, Carenage Post Office

Trinidad and Tobago, W.I.

Tel. (868) 634-4291-4

Fax. (868) 634-4433

Email: [email protected]

ii

ABSTRACT

In 2004, while preparing the State of the Marine Environment Report, it became apparent

that there was a lack of recent information on the state of reefs in Tobago except for

Buccoo Reef which has been monitored under the CARICOMP project. In addition,

there were many requests from the public for information on the state of reefs. This

project sought to bridge this knowledge gap and entailed a re-surveying of reefs at

Buccoo, Culloden, Arnos Vale and Speyside. Surveys were also conducted at La Guira

Bay.

Results from the assessment of the various areas around Tobago showed the fates of reefs

to be mixed in terms of comparative extent of hard coral cover over the last two and a

half decades. Mean hard coral declines were seen in some areas e.g. Buccoo and

Culloden, whereas improvements were observed in others e.g. Arnos Vale and Speyside

Reefs. Reefs in Man-o-War Bay, showed no overall trend with slight improvements in

mean hard coral cover at the mid and lower fore reef zones while there was a large

reduction in cover at the upper fore reefs. Surveys carried out at the reefs in La Guira Bay

were novel and thus comparisons could not be made. However, macroalgae cover is of

major concern especially at the Cove Ledge dive site.

Coral bleaching was not observed during surveys which was not surprising as they were

not conducted during an ‘El Niño’ period. However, high incidence of yellow band

disease was observed especially on the Buccoo and Culloden Reefs. Also, the rapid

assessment method revealed that Diadema antillarum numbers remain low compared to

pre die off density estimates.

iii

TABLE OF CONTENT

Page

ABSTRACT i

1.0 INTRODUCTION 1

2.0 METHODOLOGY 8

3.0 RESULTS 19

3.1 ARNOS VALE 19

3.2 BUCCOO 23

3.3 CULLODEN 33

3.4 MAN-O-WAR BAY 43

3.5 SPEYSIDE 50

3.6 LA GUIRA BAY 60

3.7 RAPID ASSESSMENT 68

4.0 DISCUSSION 69

5.0 CONCLUSIONS 78

6.0 REFERENCES 80

7.0 ACKNOWLEDGEMENTS 83

iv

LIST OF FIGURES

Page

Figure 2.1: Map of Tobago 8

Figure 2.2: Map of Arnos Vale Reef showing general location of

survey sites

10

Figure 2.3: Map of Buccoo Reef showing general location of survey

sites

11

Figure 2.4: Map of Culloden Bay showing general location of survey

sites

12

Figure 2.5: Map of Man-o-War Reef showing general location of

survey sites

13

Figure 2.6: Map of La Guira Bay showing general location of survey

sites

14

Figure 2.7: Map of Speyside Reef showing general location of survey

sites

15

Figure 2.8:

Species recovery curves for eight photoquadrats at 9m

Culloden West (Site A) and 16m Buccoo Western Reef

(Site B).

17

Figure 4.1: Hard coral cover comparisons between the Laydoo (1985

a-e) surveys and the current assessments

70

v

LIST OF TABLES

Page

Table 1.1:

Table 1.2:

Table 2.1:

Table 3.1.1:

Table 3.1.2:

Table 3.1.3:

Table 3.1.4:

Table 3.2.1:

Table 3.2.2:

Table 3.2.3:

Table 3.2.4:

Table 3.2.5:

Table 3.2.6:

Table 3.2.7:

Number of species found in varying benthic categories in

surveys of Tobago’s reefs (excluding those done by Laydoo,

1985 a-e).

Number of species found in varying benthic categories in

Laydoo (1985 a-e) surveys of Tobago’s reefs.

Depths surveyed at the various sampling sites located in the

different reef areas studied around Tobago. Number of survey

depths was dependent upon the fore-reef slope structure at the

sites.

Species cover on reef crest (3m) at Arnos Vale survey site.

Species mean density and frequency on reef crest (3m) at

Arnos Vale survey site.

Species cover on the upper fore reef (9m) at Arnos Vale

survey sites.

Species mean density and frequency on the upper fore reef

(9m) at Arnos Vale survey sites.

Species cover on the reef crest (3m) at Buccoo survey sites.

Species mean density and frequency on the reef crest (3m) at

Buccoo survey sites.

Species cover on the upper fore reef (9m) at Buccoo survey

sites.


(9m) at Buccoo survey sites.

Species cover on the mid fore reef (16m) at Buccoo survey

sites

Species mean density and frequency on the mid fore reef

(16m) at Buccoo survey sites

Species cover on the lower fore reef (21m) at Buccoo survey

sites

3

4

9

19

20

21

22

24

25

26

28

29

30

31

vi

LIST OF TABLES

Page

Table 3.2.8:

Table 3.3.1:

Table 3.3.2:

Table 3.3.3:

Table 3.3.4:

Table 3.3.5:

Table 3.3.6:

Table 3.3.7:

Table 3.3.8:

Table 3.4.1:

Table 3.4.2:

Table 3.4.3:

Species mean density and frequency on the lower fore reef

(21m) at Buccoo survey sites.

Species cover on the reef crest (3m) at Culloden survey site


Culloden survey site.

Species cover on the upper fore reef (9m) at Culloden survey

sites.


(9m) at Culloden survey sites

Species cover on the mid fore reef (16m) at Culloden survey

sites.


(16m) at Culloden survey sites

Species cover on the lower fore reef (21m) at Culloden

survey site.


(21m) at Culloden survey site

Species cover on the reef crest (3m) at the Man-o-War Bay

survey sites


Man-o-War Bay survey sites

Species cover on the upper fore reef (9m) at Man-o-War Bay

survey sites

32

34

35

36

37

39

40

41

42

44

45

47

vii

LIST OF TABLES

Page

Table 3.4.4:

Table 3.4.5:

Table 3.4.6:

Table 3.4.7:

Table 3.4.8:

Table 3.5.1:

Table 3.5.2:

Table 3.5.3:

Table 3.5.4:

Table 3.5.5:

Table 3.5.6:

Table 3.5.7:


(9m) at Man-o-War Bay survey sites

Species cover on the mid fore reef (16m) at Man-o-War Bay

survey site


(16m) at Man-o-War Bay survey site

Species cover on the lower fore reef (21m) at Man-o-War

Bay survey site


(21m) at Man-o-War Bay survey site

Species cover on the reef crest (3m) at Angel Reef, Speyside

survey sites


Angel Reef, Speyside survey sites

Species cover on the upper fore reef (9m) at Angel Reef,

Speyside survey sites


(9m) at Angel Reef, Speyside survey sites

Species cover on the mid fore reef (16m) at Angel Reef,




Species cover on the lower fore reef (21m) at Angel Reef,


48

49

49

50

50

52

53

54

55

57

58

59

viii

LIST OF TABLES

Page

Table 3.5.8:

Table 3.6.1:

Table 3.6.2:

Table 3.6.3:

Table 3.6.4:

Table 3.6.5:

Table 4.1:



Species cover on the upper fore reef (9m) at Cove Ledge, La

Guira survey sites

Species cover on the upper fore reef (9m) at Majeston, La

Guira survey sites


(9m) at the Cove Ledge and Majeston, La Guira Bay

Species cover on the upper fore reef (9m) at Flying Reef, La

Guira Bay


(9m) at Flying Reef, La Guira survey

Siderastrea siderea percentage cover change between the

Laydoo (1985 a,b,c,e) surveys and the present surveys

60

62

63

64

66

67

76

LIST OF PLATES

Page

Plate 2.1: Photoquadrat from 16m depth at Western Reef, Buccoo 16

1

1.0 INTRODUCTION

Coral reefs are an integral part of the Caribbean environment, providing over 100 million

people in more than 25 countries and territories with food, coastal protection, and

revenue from tourism (Population Reference Bureau 1996). Tourism is the region’s most

important economic sector, and reefs provide much of the sand for the region’s beaches,

where most tourists spend their time (Burke & Maidens 2004). Tourist arrivals to the

Caribbean were estimated at nearly 28 million in 1999 (Caribbean Tourism Organisation

2001) and tourism revenue alone brings in over US $25 billion a year to the region

(Burke & Maidens 2004).

In spite of their importance, Caribbean coral reefs are under severe pressure mainly from

human activities such as coastal development, land clearance and intensive agriculture

among others (Mora 2008). In addition to this, there are impacts from global phenomena

such as climate change (Johannes 1970; Rogers 1985, 1990; Woodley 1992; Glynn 1997;

UNEP 2002). Thus, seventy-five percent of Caribbean reefs are in serious decline or

under threat (Hinrichsen 1996). There has been a major loss of coral cover and diversity

(Hoegh-Guldberg 1999; Wilkinson 2000; Gardner et al. 2003), coupled in many areas

with an increase in algal biomass and shift in algal community structure (Littler et al.

1992; Lapointe 1997; Hughes 1994).

Tobago reefs are not exempt from these deleterious impacts and changes. However, the

extent to which they are being affected is not quite known. Most studies on coral reefs

here have concentrated on the Buccoo and Speyside Reef ecosystems. Reefs at Man-O-

War Bay, Culloden and Arnos Vale have also received some attention.

Buccoo Reef, designated a marine protected area (MPA) in 1973 and with its long history

of being one of Tobago’s major tourist attractions, is the most studied reef system in

Tobago. Goreau (1967) carried out largely qualitative work within the context of

potential impacts on the reef from planned tourist development on the south-western

2

coast of the island. Aerial and underwater observations were made of the reef, the latter

being done on the eastern side of the ecosystem noting general floral and faunal zonation

and species diversity patterns. Bon Accord Lagoon and associated mangrove habitat was

also similarly observed. Goreau commented and speculated upon the trophic relationships

amongst and between the Buccoo Reef-Bon Accord Lagoon complex noting the close

interdependence between the reef and mangrove lagoon, and actually considering them as

a single ecological unit. Recommendations were also made for the preservation of the

entire wetland ecosystem in the face of what was then proposed development. These

included the creation of a marine park and initiating long term study on the ecology of the

Buccoo area with particular reference to the mangroves and reefs.

Several years later Kenny (1976) conducted a preliminary study of the Buccoo Reef/Bon

Accord complex in response to the general question of development and management of

coastal resources. This preliminary study entailed a hydrographic survey with a view to

producing an up-dated map of the reef and lagoon, a faunal assessment so that a

comprehensive species list could be made and lastly, a delineation of the extent of

damage to the reef.

In the purely qualitative damage assessment it was found that, generally speaking, the

seaward slopes and the reef crest, excluding the tourist area, were only slightly damaged.

Damages observed in these areas were attributed to natural causes and more specifically

storm surge. The tourist area (the reef crest and back reef of Outer Reef) however, was

extensively damaged although, due to compounding historical data, it was uncertain

whether the damage here was caused by human activity. Kenny did suggest that storms

initially damaged the tourist area and subsequent activity on the reef prevented the

natural regeneration that might be expected.

With regards to the flora and fauna of the reef and lagoon Kenny (1976) concluded that

compared with other reefs in Tobago, Buccoo Reef is somewhat impoverished. He cited

sub-oceanic conditions e.g. low salinity during the rainy season, high turbidity and high

3

turbulence, as being possible reasons for this. His investigations yielded 1 sponge

species, 2 hydrozoans, 21 hexacoral species and 8 octocoral species (Table 1.1).

Table 1.1: Number of species found in varying benthic categories in surveys of Tobago’s

reefs (excluding those done by Laydoo, 1985a-e)

Reef Location

BENTHIC CATEGORY

Study Author Sponges Hydrozoans Octocorals Zoanthids Hexacorals Black Corals

Man-O-War Bay

Ramsaroop (1981)

4* (4)

3

21

2

24

2* (2)

Buccoo Reef

Kenny (1976)

1* (1)

2

8

2

21

undetermined

Speyside

IMA (2002)

3* (2)

2

9* (6)

1

26

undetermined

* (#) - no. undetermined at species level

Laydoo’s (1985a) benthic species number counts on Buccoo Reef (Table 1.2) differ

slightly from those reported by Kenny (1976). The former surveys at different reef sites

throughout Tobago however, did validate claims by the latter that benthic species number

in Buccoo is less than most other reef areas in Tobago. The mapping and benthic

sampling studies done by Laydoo remain today the most comprehensive quantitative

survey ever carried out on the Buccoo Reef ecosystem, gathering information on mean

species cover (% m-2

), mean density (col. m-2

) and frequency of occurrence (%) of the

benthic community.

In the aforementioned ecological surveys, the authors all highlighted the need to monitor,

assess and mitigate the impacts of sewage pollution and land run-off. Little quantitative

work was done until Lapointe et al. (2003) who did provide actual evidence of sewage

driven eutrophication in the Buccoo Reef Complex (BRC). Laydoo and Heileman (1987)

in a study of the environmental impacts of sewage treatment plants in Buccoo and Bon

Accord did find that improperly functioning plants exhibited direct impacts on the quality

of the downstream and receiving marine environment, including the BRC.

4

Table 1.2: Number of species found in varying benthic categories in Laydoo (1985a-e)

surveys of Tobago’s reefs

BENTHIC CATEGORY

Reef Location Algae Sponges Hydrozoans Octocorals Zoanthids Hexacorals Black Corals

Buccoo Reef 3* (3) 2* (2) 3 14 1 21 1

Speyside 4* (4) 1** 4* (1) 16 2 24 2* (1)

Man-O-War Bay 3* (3) 4* (4) 5* (2) 19 3 27 3

Culloden 1* (1) (undet.) 4 16 1 27 0

Arnos Vale 2* (2) 3* (3) 3* (1) 13 1 20 0

* (#) - no. undetermined at species level

** - distinguished up to level of class

Several years later Lapointe et al. (2003) reported that dissolved inorganic nitrogen

(DIN) and chlorophyll-a within the BRC were high compared to other fringing reefs

around Tobago and were indicative of eutrophication. In addition, values for δ15

N ratios

(= 15

N/14

N) for macroalgae in the BRC were within the range reported for those growing

on sewage nitrogen. They alluded to the fact that relatively low cover of hermatypic

corals and high cover of macroalgae, turf algae, octocorals and Palythoa was evidence of

cumulative impacts of land based nutrient enrichment.

Two extensive surveys were conducted on the reefs in Speyside, the first being Laydoo

(1985e) and the second by IMA (2002). Laydoo’s study – a preliminary investigation of

the nature and distribution of the reefs at Speyside – found that species distribution and

abundance appeared to be related to the structural zonation of the reefs. Hydrozoans and

zoanthids were common on the shallow reef flats, octocorals and stony corals dominant

on the fore reef slopes and black corals and sponges abundant on the lower fore reef

areas. In total 48 species of hydrozoans, octocorals, zoanthids, stony corals and black

corals were recorded in his surveys (Table 1.2).

IMA (2002) also found scleractinians, gorgonians and sponges to be the dominant

organisms observed on the reefs. This study, which also incorporated physical

oceanography, water and sediment quality and microbiological analysis with the

biological/ecological components found that cover of live substrate did not change

5

significantly since the Laydoo 1985 survey. However, many diseased corals were

observed, especially the reef building corals – Montastrea annularis, Siderastrea siderea

and Diploria strigosa. Infection in corals was observed at all depths. In addition to this

the population of Diadema was still low following mass mortality that occurred region

wide (Lessios et al. 1984, Laydoo 1985 f). Algal mats observed covering dead corals at

Big Reef and Lucy Vale Reef in the Speyside area were most likely a sign of the strained

populations.

Along with Buccoo and Speyside, the nature and distribution of reefs at Man-O-War Bay,

Culloden Bay and Arnos Vale was also determined by Laydoo (1985b-d). The coral reefs

in Man-O-War Bay were previously surveyed by Ramsaroop (1981). He carried out

observations and identified benthic inhabitants at Booby Island Reef, Rest House Reef

and Pirate’s Bay Reef. No quantitative investigations were carried out, but an idea of

species number in varying benthic classes and sub classes was ascertained (Table 1.1).

Although, reefs at Man-O-War Bay lack lagoons - which suggest that these reefs are

younger than the Buccoo Reef - Ramsaroop postulated that being further removed from

the effects of the Orinoco River effluent, especially lower salinity and higher turbidity

levels, allowed reefs at Man-O-War to be more species rich than the Buccoo Reef. Both

he and Laydoo (1985b) endorsed the huge potential, especially of Rest House and

Turpin’s Reef, for marine tourism, due to their accessibility. They did stress however,

that this characteristic could also prove disadvantageous if proper conservation

precautions were not taken.

Man-O-War Bay reefs were impacted by the mortality of Diadema antillarum which was

first officially documented on Tobago’s reefs in March 1984 at Man-O-War’s Booby reef

(Laydoo 1985f). Pre-mortality estimates here were approximately five (5) urchins.m-2

.

As alluded to earlier, this mass mortality was a region wide phenomenon in the

Caribbean (Lessios et al. 1984) with all reefs in Tobago eventually being affected.

Surveys at Culloden Bay, on Tobago’s leeward coast, indicated that the reef system was

in a youthful and vigorous stage of development (Laydoo 1985c). This was inferred

6

through the presence of a diverse benthic fauna (Table 1.2) and more so, a ‘spur and

groove’ fore-reef morphology. The reef comprised of numerous sediment channels, at

right angles to the coastline, separated by buttresses made of mainly large, hermatypic

stony corals. At the time, anthropogenic impacts were negligible apart from some fishing

activity in the marine area adjacent to the reef. However evidence of storm damage and

white band disease was observed in the shallow water Acropora palmata. Like Diadema

die off, high mortality of A. plamata due to white band disease was also reported in

Tobago (Laydoo 1985g), and by extension, a region wide phenomenon. Nonetheless, the

diverse marine life and relatively undisturbed environment at Culloden Bay led Laydoo

(1985h) to suggest that the reef system can represent a control site in monitoring

environmental impacts at other reef localities in Tobago.

Laydoo (1985d) provides the only literature to date on the nature and distribution of reefs

in Arnos Vale. It was found that reefs here had lower species diversity compared to other

reef localities around Tobago (Table 1.2). The smaller horizontal and vertical extent of

the reef areas in the bay was proposed as a reason for this. Nonetheless, it was a popular

area for snorkellers although it was found that the reefs remained generally undisturbed

by their activities. In addition to this, environmental factors that could be detrimental to

marine life, such as sedimentation and pollution, were minimal.

In an executive summary of the ecological survey of reefs around Tobago (Laydoo

1985h) several conclusions were arrived at. These included the fact that extensive

distribution of coral rubble, observed in many shallow reef areas, could be attributed to

the effects of storms and hurricanes. This rubble framework in turn contributed to an

enhanced habitat that led to high fish diversity and abundance on the reefs. Laydoo

(1985h) also concluded that conflicts existed in space utilization at many of the localities

investigated, especially between artisanal fishing activities and marine tourism and

recreation activities. Similar conflicts still remain today.

In 2004, while preparing the State of the Marine Environment Report, it became apparent

that there was a lack of recent information on the state of reefs in Tobago except for

7

Buccoo Reef which has been monitored under the CARICOMP project. In addition,

there have been many requests from the public for information on the state of reefs.

Thus, commencing in late 2007, through the IMA’s Coral Reef Monitoring Project,

which is a component of the larger Coastal Conservation Project, a re-surveying of the

major reefs in Tobago took place.

This research project was designed to address the lack of information on the health of the

major reef systems. Reefs at Buccoo, Culloden, Arnos Vale, Man-o-War Bay and

Speyside, which were qualitatively and quantitatively surveyed in the 1980s, were re-

surveyed. The aim was to identify changes/impacts to these areas. In addition, the

fringing reefs of La Guira Bay were also surveyed. Little qualitative and more so,

quantitative coral reef research has been carried out here (IMA 1990) although it is well

known that diverse reefs, increasing in popularity as dive sites, can be found in this area.

8

2.0 METHODOLOGY

Surveys of reefs in six areas of Tobago (Figure 2.1) were conducted using a belt-quadrat

method (Dodge et al. 1982) to quantitatively describe the non-living and living benthos in

terms of species frequency, density and cover. In five of the reef areas, their associated

sampling sites were determined using maps from Laydoo (1985a-e). A sixth area with

reefs, La Guira Bay – located on the Atlantic side of southwest Tobago – was not

previously surveyed by Laydoo but the lack of quantitative data of reefs in this region

and its increasing importance in dive tourism warranted its inclusion in this study. Here

major dive spots in the area were selected as locations for sample sites. Survey depths at

each of the sampling localities throughout the reef areas in Tobago were 3m, 9m, 16m

and 21m, fore-reef slope structure permitting (Table 2.1).

Figure 2.1: Map of Tobago

9

Table 2.1: Depths surveyed at the various sampling sites located in the different reef areas

studied around Tobago. Number of survey depths was dependent upon the

fore-reef slope structure at the sites

Area Sampling Site Surveying Depths

Arnos Vale Bay (Figure 2.2) Eastern side of Arnos Vale Bay 3m, 9m

Western side of Arnos Vale Bay 9m

Buccoo (Figure 2.3) Eastern Reef, Buccoo 3m, 9m

Outer Reef, Buccoo 3m, 9m, 16m

Northern Reef, Buccoo 3m, 9m, 16m, 21m

Western Reef, Buccoo 3m, 9m, 16m, 21m

Culloden Bay (Figure 2.4) Eastern side of Culloden Bay 3m, 9m, 16m, 21m

Western side of Culloden Bay 9m, 16m

Man-o-War Bay (Figure 2.5) Eastern side of Booby Island 3m, 9m, 16m, 21m

Western side of Booby Island 3m, 9m

Rest House Reef 3m

Pirate's Reef 3m, 9m

La Guira Bay (Figure 2.6) Dive site known as Cove Ledge

(2 sites)

9m

Dive site known as Majeston (2 sites) 9m

Flying Reef (5 sites) 9m

Speyside (Figure 2.7) In line with concrete Jetty adjacent to white

house on Goat Island

3m, 9m, 16m, 21m

Approximately 100m South of jetty 3m, 9m, 16m, 21m

Approximately 100m North of jetty 3m, 9m, 16m, 21m

A methodology similar to that adopted by Laydoo (1985a-e) was applied to allow for

easier comparison of the two studies. At each sampling depth a sequence of eight 1.0 m2

quadrats were photographed. A sample area of 8.0m2 was previously determined to be

optimum by Laydoo (1985a-e) through the use of a species area curve. At the respective

depths at each sampling locality the starting point for photographs i.e. photoquadrat 1,

was selected at random with successive photoquadrats taken thereafter at 2m intervals

along the contours.

10

Figure 2.2: Map of Arnos Vale Reef showing general location of survey sites

11

Figure 2.3: Map of Buccoo Reef showing general location of survey sites

12

Figure 2.4: Map of Culloden Bay showing general location of survey sites

13

Figure 2.5: Map of Man-o-War Reef showing general location of survey sites

14

Figure 2.6: Map of La Guira Bay showing general location of survey sites

15

Figure 2.7: Map of Speyside Reef showing general location of survey sites

The photoquadrats were taken using a Canon S3-IS digital camera with an IKELITE W-

20 underwater wide angle conversion lens (magnification 0.56x). The camera was

enclosed in an IKELITE underwater housing. The entire setup was attached to a 1.3m rod

which maintained a fixed distance between camera and reef to ensure a minimum area of

1.0m2 was captured in each photoquadrat (Plate 2.1).

16

Plate 2.1: Photoquadrat from 16m depth at Western Reef, Buccoo

The photoquadrats were analysed using the National Coral Reef Institute (NCRI)/Nova

South-eastern University Oceanographic Centre’s Coral Point Count with Excel

extensions program (CPCe V 3.4). Through CPCe seventy five (75) random points were

applied to each photoquadrat. Laydoo in his 1985 studies used 450 points however use of

this many in these assessments was determined to be impractical. Seventy five random

points was found to be sufficient through the use of species recovery curves (Figure 2.8).

17

Figure 2.8: Species recovery curves for eight photoquadrats at 9m Culloden West (Site A)

and 16m Buccoo Western Reef (Site B). Species recovery curves were used to

determine the appropriate number of random points to be used for analysis of

photoquadrats in this study

Sixteen photoquadrats, eight each from what were deemed, through preliminary

investigations, to be the two most species diverse sampling sites, were overlaid using

initially fifty (50) random points. The number of points was increased by increments of

five until all coral species within photoquadrat fell under at least one random point over

the series of overlays. The maximum number of points needed to do this during the

testing of the sixteen photoquadrats was then taken as the number to be used in

photoquadrat analysis in the study.

In the study, the non-living/living bottom underlying each random point was identified.

Living bottom was identified to species where possible, especially in the case of hard

corals, and the results expressed as mean species cover (% m-2

) and frequency (%). The

number of colonies, irregardless of whether or not they underlay a random point, of each

hard coral species, gorgonian (family level primarily and genus level in the cases of

18

Pseudopterogorgia, Erythropodium and Briareum) and sponges (family level) in the

photoquadrat was also counted to obtain mean densities (col. m-2).

The AGRRA (2005) rapid assessment method was also applied to obtain information on

additional parameters such as recruitment, presence/absence of Diadema antillarum and

disease and bleaching occurrence. A twenty (20) metre transect line was laid just above

the reef surface at each sample depth. In disease/bleaching assessment, any stony coral

for which any part of its skeleton underlay the transect line was observed. Affected coral

species and percentage of coral surface impacted was recorded. The type of coral

disease/severity of discolouration (for bleaching) was also noted. Recruitment was

analysed by recording any hard coral species smaller than 2cm in diameter within a half

metre belt on either side of the transect line. Any Diadema antillarum seen within these

half metre zones were also recorded.

19

3.0 RESULTS

3.1 ARNOS VALE

Arnos Vale’s reefs extended to a maximum depth of about 10m allowing for surveys to

be conducted at only two depths. At an eastern locality, the reef flat at 3m and upper fore

reef at 9m was surveyed while only a depth of 9m was surveyed at a western locality

(Figure 2.2).

At 3m the total live bottom cover was very high (97.8%) with encrusting coralline algae

(41.2% cover) and the zoanthid Palythoa caribaeorum (39.3% cover) dominating the

species assemblage (Table 3.1.1). Hard coral made up only 11.2% of the benthos with

Millepora complanata (6% cover, 1.375 col.m-2

) and, to a lesser extent, the knobby brain

coral Diploria clivosa (2.1% cover, 1.5 col.m-2

) dominating among the 7 hard coral

species recorded (Table 3.1.2). In comparison to the other major groups recorded,

sponges (4.8% cover) and gorgonians (1.3% cover) covered a very limited amount of the

benthos.

Table 3.1.1: Species cover on reef crest (3m) at Arnos Vale survey site

Arnos Vale 3m

MAJOR CATEGORY (% of transect) Species (% of transect) Arnos Vale East MEAN

HARD CORAL 11.17 11.17

Diploria clivosa 2.17 2.17

Diploria strigosa 0.83 0.83

Favia fragum 0.33 0.33

Meandrina meandrites 0.50 0.50

Millipora alcicornis 0.33 0.33

Millipora complanata 6.00 6.00

Porites astreoides 1.00 1.00

GORGONIANS 1.33 1.33

Erythropodium 0.33 0.33

Gorgonian (general) 1.00 1.00

SPONGES 4.83 4.83

ZOANTHIDS 39.33 39.33

Palythoa sp. 39.17 39.17

Zoanthid (general) 0.17 0.17

ENCRUSTING CORALLINE ALGAE 41.17 41.17

SAND, PAVEMENT, RUBBLE 2.17 2.17

20

Table 3.1.2: Species mean density and frequency on reef crest (3m) at Arnos Vale survey

site

Arnos Vale 3m Mean Density (col. m

-2) (n=8) Frequency (%)

Species Arnos Vale East Arnos Vale East



Favia fragum 0.75 25






Gorgonian (general) 0.5 25

Sponge 3.5 87.5

At 9m the total mean live cover dropped to about 60%. Here a red macroalgae (unknown

species: tuft like, growing on rubble) dominated with an average cover of 22.2±0.2%

(Table 3.1.3). Mean hard coral cover was 14±7.5% although cover at the western survey

site was more than twice that at the eastern locality (19.3% and 8.7% cover respectively).

On average however, no one hard coral species dominated out of the 14 recorded.

Diploria strigosa (4.5±0.7%) and Montastrea cavernosa (3.7±4%) had the highest

coverages. Encrusting coralline algae also had a fair mean percentage cover

(12.9±17.3%) although this was largely because it made up 25.2% of the cover at the

western survey site with only 0.7% cover recorded at the eastern site. All the other major

category groupings – sponges, gorgonians and zoanthids each had approximately 5% or

less mean cover although sponge cover was relatively high (7.8%, 4.25col.m-2

) at the

western locality (Table 3.1.4).

21

Table 3.1.3: Species cover on the upper fore reef (9m) at Arnos Vale survey sites

Arnos Vale 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Arnos Vale East Arnos Vale West MEAN STD. DEV. STD. ERROR

HARD CORAL 8.67 19.33 14.00 7.54 5.33

Agaricia agaricites 0.00 0.33 0.17 0.24 0.17

Dichocoenia stokesi 0.00 0.50 0.25 0.35 0.25

Diploria labyrinthiformis 0.17 1.17 0.67 0.71 0.50

Diploria strigosa 5.00 4.00 4.50 0.71 0.50

Meandrina meandrites 0.17 0.83 0.50 0.47 0.33

Millipora alcicornis 1.17 0.83 1.00 0.24 0.17

Millipora complanata 0.00 1.50 0.75 1.06 0.75

Montastraea cavernosa 0.83 6.50 3.67 4.01 2.83

Montastrea faveolata 0.00 2.50 1.25 1.77 1.25

Mycetophyllia ferox 0.17 0.00 0.08 0.12 0.08

Porites astreoides 0.17 0.50 0.33 0.24 0.17

Scolymia cubensis 0.00 0.17 0.08 0.12 0.08

Siderastrea radians 0.50 0.17 0.33 0.24 0.17

Siderastrea siderea 0.50 0.33 0.42 0.12 0.08

GORGONIANS 6.00 2.00 4.00 2.83 2.00

Briareum 0.83 0.00 0.42 0.59 0.42

Erythropodium 1.33 0.33 0.83 0.71 0.50

Gorgonian (general) 0.83 0.50 0.67 0.24 0.17

Pseudopterogorgia 3.00 1.17 2.08 1.30 0.92

SPONGES 3.00 7.83 5.42 3.42 2.42

ZOANTHIDS 0.83 2.67 1.75 1.30 0.92

Palythoa sp. 0.83 2.67 1.75 1.30 0.92

MACROALGAE 22.00 22.33 22.17 0.24 0.17

Macroalgae (general) 22.00 22.33 22.17 0.24 0.17

ENCRUSTING CORALLINE ALGAE 0.67 25.17 12.92 17.32 12.25

SAND, PAVEMENT, RUBBLE 58.83 20.67 39.75 26.99 19.08

22

Table 3.1.4: Species mean density and frequency on the upper fore reef (9m) at Arnos Vale survey

sites

Arnos Vale 9m Mean Density (col. m


Species Arnos Vale East Arnos Vale West Arnos Vale East Arnos Vale West

Agaricia agaricites 0 0.625 0 12.5

Dichocoenia stokesi 0 0.25 0 12.5

Diploria labyrinthiformis 0.125 0.25 12.5 25

Diploria strigosa 1.375 2.875 87.5 62.5

Meandrina meandrites 0.125 0.5 12.5 25

Millipora alcicornis 1.25 1 62.5 37.5

Millipora complanata 0.125 0.375 0 25

Montastraea cavernosa 0.375 2.5 25 75

Montastrea faveolata 0 0.5 0 25

Mycetophyllia ferox 0.125 0 12.5 0

Porites astreoides 0.125 0.25 12.5 25

Scolymia cubensis 0 0.125 0 12.5

Siderastrea radians 0.625 0.5 37.5 12.5

Siderastrea siderea 0.125 0.125 12.5 12.5

Briareum 0.625 0 37.5 0

Erythropodium 0.75 0.25 50 25

Gorgonian (general) 1.25 0.75 50 12.5

Pseudopterogorgia 1.5 0.625 37.5 37.5

Sponge 3 4.25 87.5 100

Diploria clivosa 0.125 0 n/a n/a

23

3.2 BUCCOO

The fore reef slope structure on the Buccoo Reef allowed for surveys to be conducted at all four

depths at two sites (Northern reef and Western reef), 3m through 16m depths at Outer reef and

only the two shallowest depths at Eastern reef (Figure 2.3).

The total mean live cover at 3m was just under 80% with Palythoa caribaeorum, a zooanthid,

making up on average, 32±40.9% out of that live cover (Table 3.2.1). There was great variability

in Palythoa cover throughout the survey localities however, with 86.3% and 40.7% cover at

Outer and Northern reefs respectively, whereas there was less than 1% cover at Western reef. At

Eastern reef no Palythoa was recorded.

Encrusting coralline alga was also a dominant component of the benthos at 3m (22.8±19.5%

mean cover) but like Palythoa was also highly variable among survey sites. Mean macroalgal

cover was 14.3±16.9% at this depth. However, macroalgae and Palythoa coverage varied

inversely - macroalgal cover was high (35% at Western reef and 21.3% at Eastern reef) where

Palythoa coverage was low and vice versa. There was very little to no macroalgal cover at the

Outer and Northern reef sites. Mean hard coral cover was only 5.6±4.3% with Millipora sp.

making up over half of this figure. Millipora complanata coverage (9.3%) (Table 3.2.1) and

density (6.25 col.m-2

) (Table 3.2.2) was particularly high at Eastern reef. Gorgonians (2.6±4.7%)

and sponges (1.4±1.3%) showed very low coverage at 3m.

Encrusting coralline algae dominated the living benthos at 9m accounting for 38.8±38.7% of the

total mean living cover, which was about 71% (Table 3.2.3). Variability in cover among sites

was high, ranging from 0% at Eastern reef to 82.3% at Northern reef. At this depth, mean hard

coral and gorgonian cover were both about 14% each with Montastrea faveolata dominating

(6.3±5.1%) the cover for the 12 hard coral species recorded, and Erythropodium caribaeorum

(5.8±2.9%) doing the same for the gorgonian category. Interestingly, hard coral cover decreased

steadily from site to site as one moved in an east to west direction around the arc of reef

platforms that make up part of the Buccoo Reef complex. It was also observed that on average,

23.2±17.3% of the coral was affected by yellow band disease. The highest incidences took place

24

Table 3.2.1: Species cover on the reef crest (3m) at Buccoo survey sites

Buccoo 3m

MAJOR CATEGORY (% of transect) Species (% of transect) Buccoo Eastern Reef Buccoo Outer Reef Buccoo Northern Reef Buccoo Western Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 11.50 2.33 2.50 6.17 5.63 4.30 2.15

Acropora palmata 0.00 0.00 0.00 2.33 0.58 1.17 0.58

Diploria clivosa 0.00 0.00 0.67 3.67 1.08 1.75 0.88

Diploria strigosa 0.00 0.00 0.17 0.00 0.04 0.08 0.04

Favia fragum 0.00 0.00 0.17 0.17 0.08 0.10 0.05

Millipora alcicornis 1.33 1.83 0.67 0.00 0.96 0.80 0.40

Millipora complanata 9.33 0.50 0.50 0.00 2.58 4.51 2.25

Montastraea annularis 0.33 0.00 0.00 0.00 0.08 0.17 0.08

Porites astreoides 0.50 0.00 0.33 0.00 0.21 0.25 0.13

GORGONIANS 0.83 0.00 9.67 0.00 2.63 4.71 2.36

Erythropodium 0.83 0.00 0.67 0.00 0.38 0.44 0.22

Gorgonian (general) 0.00 0.00 9.00 0.00 2.25 4.50 2.25

SPONGES 1.50 0.33 3.17 0.67 1.42 1.27 0.63

ZOANTHIDS 0.00 86.33 40.67 0.83 31.96 40.92 20.46

Palythoa sp. 0.00 86.33 40.67 0.83 31.96 40.92 20.46

MACROALGAE 21.33 0.67 0.33 35.00 14.33 16.92 8.46

Halimeda 0.00 0.00 0.33 0.17 0.13 0.16 0.08

Macroalgae (general) 21.33 0.67 0.00 34.83 14.21 16.95 8.47

ENCRUSTING CORALLINE ALGAE 2.17 10.00 40.17 38.67 22.75 19.52 9.76

SAND, PAVEMENT, RUBBLE 62.67 0.33 3.50 18.67 21.29 28.72 14.36

25

Table 3.2.2: Species mean density and frequency on the reef crest (3m) at Buccoo survey sites

Buccoo 3m Mean Density (col. m


Species Buccoo Eastern

Reef

Buccoo Outer

Reef

Buccoo Northern

Reef

Buccoo Western

Reef

Buccoo Eastern

Reef

Buccoo Outer

Reef

Buccoo Northern

Reef

Buccoo Western

Reef

Acropora palmata 0 0 0 0.125 0 0 0 12.5

Diploria clivosa 0 0 0.375 2 0 0 37.5 100

Diploria strigosa 0.125 0 0.125 0 0 0 12.5 0

Favia fragum 0 0 0.25 0.125 0 0 12.5 12.5

Millipora alcicornis 1.625 1.375 0.75 0 50 75 37.5 0

Millipora complanata 6.25 0.375 0.25 0 75 12.5 25 0

Montastraea annularis 0.25 0 0 0 12.5 0 0 0

Porites astreoides 0.25 0 0.125 0 12.5 0 12.5 0

Erythropodium 0.5 0 0.5 0 37.5 0 25 0

Gorgonian (general) 0 0 4 0 0 0 75 0

Sponge 0.875 0.125 1.125 0.625 50 12.5 37.5 37.5

26

Table 3.2.3: Species cover on the upper fore reef (9m) at Buccoo survey sites

Buccoo 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Buccoo Eastern Reef Buccoo Outer Reef Buccoo Northern Reef Buccoo Western Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 20.50 17.67 10.00 7.00 13.79 6.34 3.17

Agaricia agaricites 0.00 0.00 0.50 0.00 0.13 0.25 0.13

Colpophyllia natans 0.00 1.33 0.00 0.83 0.54 0.66 0.33

Diploria labyrinthiformis 0.00 0.00 0.33 0.50 0.21 0.25 0.13


Eusmilia fastigiata 0.00 0.00 0.00 0.33 0.08 0.17 0.08



Montastraea annularis 1.00 0.00 0.00 0.00 0.25 0.50 0.25

Montastrea faveolata 2.33 13.50 6.17 3.00 6.25 5.11 2.56


Porites porites 0.00 0.00 0.00 0.67 0.17 0.33 0.17

Siderastrea siderea 0.00 1.83 0.00 0.00 0.46 0.92 0.46

GORGONIANS 4.17 12.33 5.67 32.83 13.75 13.21 6.60

Briareum 0.00 0.00 0.00 18.00 4.50 9.00 4.50

Erythropodium 4.17 7.50 2.50 8.83 5.75 2.92 1.46


Pseudopterogorgia 0.00 2.67 0.00 1.33 1.00 1.28 0.64

SPONGES 4.50 0.33 0.00 0.00 1.21 2.20 1.10

ZOANTHIDS 0.00 0.50 1.67 0.00 0.54 0.79 0.39

Palythoa sp. 0.00 0.50 1.67 0.00 0.54 0.79 0.39

MACROALGAE 6.67 0.00 0.00 3.17 2.46 3.18 1.59

Halimeda 0.00 0.00 0.00 2.67 0.67 1.33 0.67


ENCRUSTING CORALLINE ALGAE 0.00 59.67 82.33 13.17 38.79 38.70 19.35


27

at the Outer and Northern reefs sampling locations with 38.7% and 36.7% of corals

affected respectively. Yellow band disease primarily affected Monatstrea faveolata.

Mean macroalgal (2.5±3.2%) and sponge (1.2±2.2%) cover at this depth were very low.

However, at Eastern reef in particular, macroalgal cover (6.67%) and sponge cover

(4.5%, 3.375 col.m-2

) (Table 3.2.4) was especially above the average.

At 16m depth mean hard coral cover was highest (30.4±1.4%) (Table 3.2.5) when

compared with all other survey depths on the Buccoo reef. Hard coral cover was

consistently high at all survey sites with Montastrea faveolata dominating at Outer reef

(16.2%, 2.5 col.m-2

) and Northern reef (9.7%, 3.125 col.m-2

) and Colpophyllia natans at

Western reef (11.3%, 1 col.m-2

) (Table 3.2.6). Fourteen hard coral species were recorded

and this was the highest number recorded at any depth in Buccoo. Encrusting coralline

algae also had substantial, consistent coverage throughout with an average cover of

27.3±7%. Gorgonians were the only other major component of the living biota

(14.5±8.5% mean cover) with Erythropodium caribaeorum making up the vast majority

(8.9±2.7%) of this grouping. Gorgonian cover was more variable than that of hard corals

and coralline algae though, with 7.2%, 12.5% and 23.8% cover at Outer reef, Northern

reef and Western reef respectively. As alluded to earlier, out of the total mean living

cover of 74.2%, zoanthids (1.4±1.3%), sponge (0.4±0.5%) and macroalgal (0.2±0.3%)

cover were all extremely low.

Total mean live cover at 21m on the Buccoo reef was just under 80% with mean

encrusting coralline algal cover accounting for about half of this (40.4±3.7%) (Table

3.2.7). Although not as high as at 16m, mean hard coral cover was still good (21.5±7.3%)

with Western reef (26.7%) having 10% higher coverage than Northern reef (16.3%).

Montastrea faveolata (14.3%, 2.25 col.m-2

and 7.2%, 2 col.m-2

respectively) (Table 3.2.8)

once again dominated out of the 12 species identified at this depth. Although variable

(23.2% at Northern reef and 9% at Western reef) mean gorgonian cover was 16.1±10%

with Erythropodium caribaeorum largely dominating at both survey sites. Once again,

sponges (1.2±0.5% cover), macroalgae (0.3±0.5% cover) and zoanthids (0.2±0.2% cover)

were minorities in the benthic cover makeup.

28

Table 3.2.4: Species mean density and frequency on the upper fore reef (9m) at Buccoo survey sites

Buccoo 9m Mean Density (col. m-2

) (n=8) Frequency (%)

Species Buccoo Eastern

Reef

Buccoo Outer

Reef

Buccoo Northern

Reef

Buccoo Western

Reef

Buccoo Eastern

Reef

Buccoo Outer

Reef

Buccoo Northern

Reef

Buccoo Western

Reef

Agaricia agaricites 0 0.125 0.375 0 0 0 25 0

Colpophyllia natans 0 0.5 0 0.5 0 37.5 0 25

Diploria labyrinthiformis 0 0 0.125 0.125 0 0 12.5 12.5

Diploria strigosa 0 0 0.625 0.875 0 0 25 37.5

Eusmilia fastigiata 0 0 0 0.25 0 0 0 25

Millipora alcicornis 5.375 0.375 1 0 100 12.5 25 0

Millipora complanata 2.75 0.375 0.125 0 62.5 12.5 12.5 0

Montastraea annularis 0.625 0 0 0 12.5 0 0 0

Montastrea faveolata 1.375 2.625 1.375 1.125 37.5 50 50 75

Porites astreoides 0.25 0.125 0.375 0 12.5 12.5 25 0

Porites porites 0 0 0 0.125 0 0 0 12.5

Siderastrea siderea 0 0.5 0 0 0 12.5 0 0

Briareum 0 0 0 3.25 0 0 0 62.5

Erythropodium 0.875 2.75 2.875 1.625 37.5 62.5 75 37.5

Gorgonian (general) 0 0.75 1.125 4.125 0 37.5 37.5 87.5

Pseudopterogorgia 0 0.5 0.125 0.375 0 37.5 0 25

Sponge 3.375 0.375 0 0.125 100 25 0 0

Scolymia cubensis 0 0 0 0.125 n/a n/a n/a n/a

29

Table 3.2.5: Species cover on the mid fore reef (16m) at Buccoo survey sites

Buccoo 16m

MAJOR CATEGORY (% of transect) Species (% of transect) Buccoo Outer Reef Buccoo Northern Reef Buccoo Western Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 31.00 31.33 28.83 30.39 1.36 0.78

Agaricia agaricites 0.00 0.50 0.50 0.33 0.29 0.17

Colpophyllia natans 2.00 8.83 11.33 7.39 4.83 2.79

Diploria labyrinthiformis 0.33 0.67 0.33 0.44 0.19 0.11

Diploria strigosa 1.00 2.50 5.00 2.83 2.02 1.17

Madracis decactis 0.33 0.17 0.33 0.28 0.10 0.06

Meandrina meandrites 0.17 0.83 0.17 0.39 0.38 0.22

Millipora alcicornis 1.00 1.50 0.83 1.11 0.35 0.20

Millipora complanata 1.67 1.17 0.33 1.06 0.67 0.39

Montastraea cavernosa 0.00 2.50 1.33 1.28 1.25 0.72

Montastrea faveolata 16.17 9.67 6.67 10.83 4.86 2.80

Porites astreoides 2.83 0.33 0.50 1.22 1.40 0.81

Porites porites 0.00 0.00 0.33 0.11 0.19 0.11

Siderastrea radians 0.33 0.33 0.00 0.22 0.19 0.11

Siderastrea siderea 5.17 2.33 1.17 2.89 2.06 1.19

GORGONIANS 7.17 12.50 23.83 14.50 8.51 4.91

Briareum 0.00 2.00 0.00 0.67 1.15 0.67

Erythropodium 6.83 7.83 12.00 8.89 2.74 1.58

Gorgonian (general) 0.00 1.83 0.83 0.89 0.92 0.53

Pseudopterogorgia 0.33 0.83 11.00 4.06 6.02 3.48

SPONGES 0.00 0.17 1.00 0.39 0.54 0.31

ZOANTHIDS 0.00 2.67 1.50 1.39 1.34 0.77

Palythoa sp. 0.00 2.67 1.50 1.39 1.34 0.77

MACROALGAE 0.00 0.00 0.50 0.17 0.29 0.17

Halimeda 0.00 0.00 0.50 0.17 0.29 0.17

ENCRUSTING CORALLINE ALGAE 25.00 35.17 21.83 27.33 6.97 4.02

SAND, PAVEMENT, RUBBLE 36.83 18.17 22.50 25.83 9.77 5.64

30

Table 3.2.6: Species mean density and frequency on the mid fore reef (16m) at Buccoo survey sites



Species Buccoo Outer Reef

Buccoo Northern Reef Buccoo Western Reef Buccoo Outer Reef Buccoo Northern Reef Buccoo Western Reef

Agaricia agaricites 0.125 1.125 0.625 0 25 37.5

Colpophyllia natans 0.125 0.625 1 12.5 50 62.5

Diploria labyrinthiformis 0.125 0.25 0.125 12.5 25 12.5

Diploria strigosa 0.25 0.75 0.75 25 50 50

Madracis decactis 0.125 0.125 0.25 12.5 12.5 25


Millipora alcicornis 0.875 1.5 0.625 25 50 25

Millipora complanata 0.5 0.25 0.25 25 25 12.5

Montastraea cavernosa 0 0.5 0.625 0 25 62.5

Montastrea faveolata 2.5 3.125 2.25 75 100 87.5

Porites astreoides 1.5 0.125 1.125 75 12.5 25

Porites porites 0 0 0.125 0 0 12.5

Siderastrea radians 0.375 0.125 0 25 12.5 0

Siderastrea siderea 1.125 1.375 0.5 62.5 62.5 50

Briareum 0 0.5 0 0 25 0

Erythropodium 1.25 2.25 3.125 87.5 100 100

Gorgonian (general) 0 1 0.75 0 62.5 37.5


Sponge 0.125 0.25 0.5 0 12.5 37.5

Scolymia cubensis 0.125 0 0.125 n/a n/a n/a

31

Table 3.2.7: Species cover on the lower fore reef (21m) at Buccoo survey sites

Buccoo 21m

MAJOR CATEGORY (% of transect) Species (% of transect Buccoo Northern Reef Buccoo Western Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 16.33 26.67 21.50 7.31 5.17


Colpophyllia natans 4.83 0.00 2.42 3.42 2.42



Eusmilia fastigiata 0.17 0.67 0.42 0.35 0.25

Madracis decactis 0.17 1.00 0.58 0.59 0.42







GORGONIANS 23.17 9.00 16.08 10.02 7.08

Erythropodium 20.33 7.33 13.83 9.19 6.50



SPONGES 0.83 1.50 1.17 0.47 0.33

ZOANTHIDS 0.33 0.00 0.17 0.24 0.17

Palythoa sp. 0.33 0.00 0.17 0.24 0.17

MACROALGAE 0.67 0.00 0.33 0.47 0.33


DEAD CORAL WITH ALGAE 0.33 0.00 0.17 0.24 0.17



32

Table 3.2.8: Species mean density and frequency on the lower fore reef (21m) at Buccoo survey sites



Species Buccoo Northern Reef Buccoo Western Reef Buccoo Northern Reef Buccoo Western Reef

Agaricia agaricites 0.875 0.5 25 12.5

Colpophyllia natans 0.5 0 37.5 0

Diploria labyrinthiformis 0.125 0.125 12.5 12.5

Diploria strigosa 0 0.5 0 37.5

Eusmilia fastigiata 0.125 0.125 12.5 12.5

Madracis decactis 0.125 0.375 12.5 25

Millipora alcicornis 0.875 0 37.5 0

Millipora complanata 0.25 0.125 25 12.5

Montastraea cavernosa 0.125 0.625 12.5 50

Montastrea faveolata 2.25 2 100 100

Porites astreoides 0.125 0.75 12.5 50

Siderastrea siderea 0.375 0.375 37.5 25

Erythropodium 3.875 2.5 100 87.5

Gorgonian (general) 0.75 0.5 37.5 25

Pseudopterogorgia 0.375 0.75 37.5 62.5

Sponge 0.125 0.75 12.5 37.5

Briareum 0 0.25 n/a n/a

Scolymia cubensis 0 0.125 n/a n/a

Mycetophyllia sp. 0 0.125 n/a n/a

33

3.3 CULLODEN

Reefs at eastern locality of Culloden were quite extensive and allowed for surveys to take

place at all four sampling depths (Figure 2.4). At the western site, however, due to fore

reef slope structure, surveys were conducted at only two depths; 9m and 16m.

Of the approximate 98% live cover observed at 3m in the Culloden area, encrusting

coralline algae made up about 50% and Palythoa caribaeorum made up about 32%

coverage (Table 3.3.1). In comparison, sponge cover at this depth was 6.3% however this

was the highest mean sponge cover for all depths surveyed in Culloden. Similarly, the

sponge density of 4.25 col.m-2

was also the highest observed when compared to all the

other localities surveyed (Table 3.3.4). Nonetheless, in this reef crest zone, like sponge

cover, coverage of hard corals (4.5%) and gorgonians (4.8%) was relatively low. They

were all higher than macroalgal cover (1.3%) though, which for the most part was

identified as Halimeda sp.

Total mean live cover at 9m was lower (71.2%) than at 3m but dominance was more

evenly distributed among the major categories. Encrusting coralline alga was once again

most dominant with just under 31±12.2% mean cover (Table 3.3.3). The 16 species of

hard corals recorded had an average cover of 16.3±4.7% with Montastrea faveolata

accounting for about a third of this (5.7±7.5%). This species was especially abundant at

the western locality – 11% (0.75 col.m-2

density) out of 20% hard coral cover compared

to 0.3% (0.25 col.m-2

density) out of 13% hard coral cover at the eastern site (Table

3.3.4). Consequently, so was the noted occurrence of yellow band disease which has an

affinity to this species in particular. It affected nearly 13% of the hard corals captured in

the photoquadrats taken at this western site while none were affected in the Eastern site’s

photoquadrats.

Gorgonians at 9m had a mean cover of about 13±10.8% but coverage at the eastern

survey site was much higher (20.8%) compared to that of the western site (5.5%) (Table

3.3.3). This was also the case with sponge cover where, although the mean cover was low

(3.8±4.5%), 7% coverage was recorded at the eastern locality compared with 0.7% at the

34

western site. About five percent (4.9±2.5%) mean zoanthid cover and just under 2%

mean macroalgal cover made up the remainder of the total mean live cover.

Table 3.3.1: Species cover on the reef crest (3m) at Culloden survey site

Culloden 3m

MAJOR CATEGORY (% of transect) Species (% of transect) Culloden East MEAN



Diploria labyrinthiformis 0.33 0.33






Montastraea cavernosa 0.17 0.17

Montastrea faveolata 0.33 0.33



Briareum 0.17 0.17



Pseudopterogorgia 0.67 0.67

SPONGES 6.33 6.33

ZOANTHIDS 31.67 31.67

Palythoa sp. 31.67 31.67

MACROALGAE 1.33 1.33

Halimeda 1.33 1.33



35

Table 3.3.2: Species mean density and frequency on the reef crest (3m) at Culloden survey

site

Culloden 3m Mean Density (col. m


Species Culloden East Culloden East

Diploria clivosa 0.25 25

Diploria labyrinthiformis 0.125 12.5








Porites astreoides 0.625 25

Briareum 0.125 12.5




Sponge 4.25 75

36

Table 3.3.3: Species cover on the upper fore reef (9m) at Culloden survey sites

Culloden 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Culloden East Culloden West MEAN STD. DEV. STD. ERROR

HARD CORAL 13.00 19.67 16.33 4.71 3.33


Colpophyllia natans 1.33 0.67 1.00 0.47 0.33



Eusmilia fastigiata 0.50 0.00 0.25 0.35 0.25

Isophyllia rigida 0.00 0.17 0.08 0.12 0.08











GORGONIANS 20.83 5.50 13.17 10.84 7.67

Erythropodium 2.33 3.67 3.00 0.94 0.67



SPONGES 7.00 0.67 3.83 4.48 3.17

ZOANTHIDS 3.17 6.67 4.92 2.47 1.75

Palythoa sp. 3.00 6.67 4.83 2.59 1.83

Zoanthid (general) 0.17 0.00 0.08 0.12 0.08

MACROALGAE 1.83 1.83 1.83 0.00 0.00

Halimeda 1.83 1.50 1.67 0.24 0.17





37

Table 3.3.4: Species mean density and frequency on the upper fore reef (9m) at Culloden

survey sites



Species Culloden East Culloden West Culloden East Culloden West

Agaricia agaricites 1.125 0.125 75 12.5

Colpophyllia natans 0.125 0.125 12.5 12.5

Diploria labyrinthiformis 0.125 0.125 12.5 12.5

Diploria strigosa 0.375 2 25 50

Eusmilia fastigiata 0.125 0 12.5 0

Isophyllia rigida 0 0.125 0 12.5

Madracis decactis 0.875 0.125 37.5 12.5

Meandrina meandrites 0.25 0.25 12.5 12.5

Millipora alcicornis 0.75 0.375 37.5 12.5

Millipora complanata 0.5 0.125 12.5 12.5

Montastraea cavernosa 1.125 0.5 62.5 37.5

Montastrea faveolata 0.25 0.75 25 50

Porites astreoides 0 0.25 0 12.5

Scolymia cubensis 0.25 0 12.5 0

Siderastrea radians 0.125 0.125 12.5 12.5

Siderastrea siderea 0.25 0.125 25 12.5

Erythropodium 1.25 2.375 50 75

Gorgonian (general) 3 0.125 100 0

Pseudopterogorgia 2.5 0.125 75 12.5

Sponge 2.625 1 87.5 37.5

Favia fragum 0 0.125 n/a n/a

38

Total mean live cover at 16m was just over 88%. Here, mean encrusting coralline algae cover

continued to dominate the benthos. Its mean cover was more than three times higher than that

of the next highest major category, hard corals (55±18.2% compared to 17.9±8.1%) (Table

3.3.5). Like at 9m though, out of the 15 species of hard corals documented Montastrea

faveolata was once again most abundant with an average cover and density of 9.9±6% and

1.75 col.m-2

respectively (Table 3.3.6). Gorgonians had a mean cover of just under

11.8±5.4% with Pseudeopterogorgia sp. (5.6±4.1% cover) making up about half that

number.

Interestingly, the survey locality on the eastern side of Culloden had almost twice the hard

coral and gorgonian cover than the western side. However, Yellow Band disease occurrence

was also much higher here with 30.3% of the hard corals affected compared to 6.9% at the

western site. The remaining categories – sponges (1.3±0.2%), macroalgae (1.2±0.7%) and

zoanthids (0.9±1.3%) – had very low mean coverage at this depth.

High coverage of encrusting coralline algae was again observed at 21m making up just over

61% of the benthic cover (Table 3.3.7). Hard corals had similar coverage to that observed at

9m and 16m about 16%. At this depth however, Montastrea faveolata (4.5% cover) shared

its hard coral dominance with Meandrina meandrites (3.8% cover). The former was clearly

encountered more frequently however (Table 3.3.8). Gorgonians had about 10% cover with

Erythropodium caribaeorum (5.3%) accounting for just over half of this. Sponges made up

4% out of the approximate 92% live cover documented with almost negligible contribution

made from macroalgae (0.5%) and none from zoanthids.

39

Table 3.3.5: Species cover on the mid fore reef (16m) at Culloden survey sites

Culloden 16m

MAJOR CATEGORY (% of transect) Species (% of transect) Culloden East Culloden West MEAN STD. DEV. STD. ERROR

HARD CORAL 23.67 12.17 17.92 8.13 5.75


Dichocoenia stokesi 0.00 0.33 0.17 0.24 0.17









Mycetophyllia aliciae 0.00 0.17 0.08 0.12 0.08





GORGONIANS 15.67 8.00 11.83 5.42 3.83

Briareum 0.00 0.33 0.17 0.24 0.17

Erythropodium 2.83 3.83 3.33 0.71 0.50



SPONGES 1.50 1.17 1.33 0.24 0.17

ZOANTHIDS 1.83 0.00 0.92 1.30 0.92

Palythoa sp. 1.83 0.00 0.92 1.30 0.92

MACROALGAE 1.67 0.67 1.17 0.71 0.50

Halimeda 1.67 0.50 1.08 0.82 0.58





40

Table 3.3.6: Species mean density and frequency on the mid fore reef (16m) at Culloden survey

sites



Species Culloden East Culloden West Culloden East Culloden West

Agaricia agaricites 0.125 0.125 12.5 12.5

Dichocoenia stokesi 0 0.125 0 12.5

Diploria labyrinthiformis 0.125 0 12.5 0

Diploria strigosa 0.25 0.125 25 0

Madracis decactis 1.125 0.875 37.5 75

Meandrina meandrites 0.125 0.125 12.5 12.5

Millipora alcicornis 0.25 0 25 0

Millipora complanata 0.375 0 25 0

Montastraea cavernosa 0.375 0.5 37.5 50

Montastrea faveolata 2.125 1.375 100 50

Mycetophyllia aliciae 0 0.125 0 12.5

Porites astreoides 0 0.25 0 12.5

Scolymia cubensis 0 0.25 0 12.5

Siderastrea radians 0 0.375 0 25

Siderastrea siderea 1.375 0.5 75 37.5

Briareum 0 0.25 0 12.5

Erythropodium 1.5 2.375 62.5 75

Gorgonian (general) 1.875 0.375 100 12.5

Pseudopterogorgia 2.375 0.75 75 37.5

Sponge 1 1.125 62.5 50

41

Table 3.3.7: Species cover on the lower fore reef (21m) at Culloden survey site.

Culloden 21m

MAJOR CATEGORY (% of transect) Species (% of transect) Culloden East MEAN


Agaricia agaricites 0.33 0.33

Dichocoenia stokesi 0.50 0.50


Leptoseris cucullata 0.17 0.17

Madracis decactis 1.67 1.67





Scolymia cubensis 0.17 0.17

Siderastrea radians 0.17 0.17

Siderastrea siderea 1.83 1.83


Briareum 2.00 2.00




SPONGES 4.00 4.00


Macroalgae (general) 0.50 0.50



42

Table 3.3.8: Species mean density and frequency on the lower fore reef (21m) at Culloden

survey site



Species Culloden East Culloden East

Agaricia agaricites 0.375 25

Dichocoenia stokesi 0.125 12.5


Leptoseris cucullata 0.125 12.5

Madracis decactis 1.125 75





Scolymia cubensis 0.25 12.5

Siderastrea radians 0.125 12.5


Briareum 0.75 50

Erythropodium 2.25 75



Sponge 0.75 25

43

3.4 MAN-O-WAR BAY

The reefs in Man-o-War Bay were not deep with the exception of Booby Island, where the

eastern side was surveyed at 21m. Reefs at the three remaining sites extended to depths of just

over 10m except for Rest House Reef whose reefs had a maximum depth of about 7m (Figure

2.5).

At 3m zoanthids were the dominant group accounting for, on average, 23.6±24.7% of the total

mean live cover of about 61% (Table 3.4.1). Palythoa caribaeorum cover was very high at the

Booby Island west (54.3%) and Pirate’s reef (32.8%) sites leading to this high mean zoanthid

cover. At the two remaining sites very little to no zoanthids were recorded. Mean hard coral

cover was 20.3±15.1% and this figure is the highest recorded at any 3m reef area surveyed

during this study. This mean cover figure is deceptive though, as the variability among the four

reef crest sites surveyed in Man-O-War bay was high. While the hard coral cover at the Booby

Island east and west sites and Pirate’s reef site were above the Tobago wide norm (about 12%,

14% and 13% cover respectively), it was the 42.8% cover at the Rest House reef locality that

really increased the hard coral cover average.

The Rest House reef site at 3m was dominated almost entirely by the yellow pencil coral

Madracis mirabilis (42.3% cover, 2.375 col.m-2

density) (Table 3.4.2) and with the fragile nature

of this species the high percentage cover of coral rubble with algae here (41.7%) was expected.

Millipora sp. was common at the Pirate’s reef site but especially so at the western side of Booby

Island where all hard coral recorded was Millipora alcicornis. To the east of Booby Island hard

coral species diversity was higher but was dominated by Millipora complanata (3.2% cover, 0.5

col.m-2

density) and Montastrea faveolata (3% cover, 0.625 col.m-2

density) (Table 3.4.2). In

total 11 species of hard coral were recorded at 3m in Man-o-War Bay among the sites surveyed.

Mean gorgonian cover at 3m was 7±3% while encrusting coralline algae and sponges had

average covers of 4.8±4.7% and 3.7±4.7% respectively. Average macroalgal cover was low

making up just 1.3±1.5% of the total benthic cover.

44

Table 3.4.1: Species cover on the reef crest (3m) at the Man-O-War Bay survey sites

Man-o-War Bay 3m

MAJOR CATEGORY (% of transect) Species (% of transect) Booby Is. East Booby Is. West Pirate's Reef Rest House Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 11.67 13.67 13.00 42.83 20.29 15.05 7.53

Agaricia agaricites 0.17 0.00 0.00 0.00 0.04 0.08 0.04

Diploria clivosa 0.67 0.00 0.00 0.00 0.17 0.33 0.17


Madracis mirabilis 0.00 0.00 0.00 42.33 10.58 21.17 10.58

Meandrina meandrites 1.17 0.00 0.00 0.00 0.29 0.58 0.29



Montastraea cavernosa 1.00 0.00 2.00 0.00 0.75 0.96 0.48

Montastrea faveolata 3.00 0.00 0.00 0.00 0.75 1.50 0.75


Siderastrea siderea 0.33 0.00 1.33 0.00 0.42 0.63 0.32

GORGONIANS 7.33 4.33 5.33 11.17 7.04 3.02 1.51

Briareum 0.00 0.00 0.50 3.00 0.88 1.44 0.72

Erythropodium 4.17 3.00 0.50 0.00 1.92 1.99 1.00


Pseudopterogorgia 1.50 0.00 3.50 5.17 2.54 2.26 1.13

SPONGES 10.67 0.50 2.67 0.83 3.67 4.76 2.38

ZOANTHIDS 3.67 54.33 32.83 3.50 23.58 24.71 12.35

Palythoa sp. 3.67 46.33 32.83 0.00 20.71 22.53 11.27

Zoanthid (general) 0.00 8.00 0.00 3.50 2.88 3.79 1.90

MACROALGAE 3.00 0.17 2.00 0.00 1.29 1.46 0.73


DEAD CORAL WITH ALGAE 0.17 0.00 0.33 41.67 10.54 20.75 10.38

ENCRUSITNG CORALLINE ALGAE 11.33 4.00 4.00 0.00 4.83 4.73 2.36


45

Table 3.4.2: Species mean density and frequency on the reef crest (3m) at Man-O-War survey sites

Man-o-War Bay 3m Mean Density (col. m-2

) (n=8) Frequency (%)

Species Booby Is. East Booby Is. West Pirate's Reef Rest House Reef Booby Is. East Booby Is. West Pirate's Reef Rest House Reef

Agaricia agaricites 0.125 0 0 0 12.5 0 0 0

Diploria clivosa 0.5 0 0 0 25 0 0 0

Diploria strigosa 0.625 0 0.875 0 50 0 50 0

Madracis mirabilis 0 0 0 2.375 0 0 0 100

Meandrina meandrites 0.25 0 0 0 25 0 0 0

Millipora alcicornis 0 4.625 2.75 0.375 0 87.5 62.5 25

Millipora complanata 0.5 0 1.625 0 37.5 0 100 0

Montastraea cavernosa 0.625 0 1.25 0 37.5 0 37.5 0

Montastrea faveolata 0.625 0 0 0 25 0 0 0

Porites astreoides 0.125 0 1 0 12.5 0 50 0

Siderastrea siderea 0.125 0 0.375 0 12.5 0 37.5 0

Briareum 0 0 0.25 1.875 0 0 25 62.5

Erythropodium 1.625 0.625 0.5 0 62.5 37.5 25 0

Gorgonian (general) 1.125 0.75 1.125 2.25 75 37.5 25 37.5

Pseudopterogorgia 0.5 0.125 1 1.75 25 0 62.5 62.5

Sponge 6.375 0.625 3.5 1.75 87.5 25 62.5 37.5

Favia fragum 0.125 0 0 0 n/a n/a n/a n/a

46

At 9m, gorgonians dominated the live benthos with an average of 13.5±8.9% cover (Table

3.4.3). Cover varied among sites though – 23% cover at Pirate’s reef, 12.6% at Booby Island

east and only 5.5% at Booby Island west. However, the 12.9±13.3% mean hard coral cover at

9m in Man-o-War Bay was largely attributable to the high hard coral percentage cover

(28.2%) recorded at Booby Island West. Montastrea faveolata dominated here – 13.3% cover

and a density of 1.125 col.m-2

(Table 3.4.4). In comparison, the other two 9m sites had low

hard coral cover, 6.5% at Pirate’s reef and 4% at Booby Island east. Overall, the total mean

live cover at 9m in Man-o-War Bay was low (about 34%) with zoanthids (2.3±2.3% mean

cover), macroalgae (2.1±2.4% mean cover), sponges (1.9±1.2% mean cover) and encrusting

coralline algae (1.4±1.3% mean cover) making little contributions.

Gorgonians again dominated at 16m with 14% cover (Table 3.4.5). The one locality surveyed

at this depth, the eastern side of Booby Island, was made up of mainly stony rubble and this

substrate type did not support a diversity of hard coral species. Only 4 species were recorded

totalling just over 4% in cover. Of the four species observed though, Millipora alcicornis had

the highest density (1.75 col.m-2

) (Table 3.3.6). Macroalgal cover increased to 6.5% at this

depth as did sponge cover (3%) and encrusting coralline algae cover (1.5%) but once again

the total live cover remained at around 34%.

Live cover dropped even lower at 21m – just over 18% - with gorgonian cover making up

more than half of this (10.8% gorgonian cover) (Table 3.4.7). At this depth only 3% hard

coral cover was evident with even fewer species (3) recorded. Once again however,

Millipora alcicornis density was highest (Table 3.4.8). Sponges also occupied about 3% of

the rubbly substrate with very little colonisation recorded from encrusting coralline algae

(1% cover), zoanthids (0.3% cover) and macroalgae (0.3% cover).

47

Table 3.4.3: Species cover on the upper fore reef (9m) at Man-O-War Bay survey sites

Man-o-War Bay 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Booby Is. East Booby Is. West Pirate's Reef MEAN STD. DEV. STD. ERROR

HARD CORAL 4.00 28.17 6.50 12.89 13.29 7.67










Porites porites 0.00 0.33 0.00 0.11 0.19 0.11


GORGONIANS 12.00 5.50 23.00 13.50 8.85 5.11

Briareum 0.00 0.00 0.50 0.17 0.29 0.17

Erythropodium 4.50 2.17 0.50 2.39 2.01 1.16



SPONGES 2.17 0.67 3.00 1.94 1.18 0.68

ZOANTHIDS 4.83 1.67 0.33 2.28 2.31 1.33

Palythoa sp. 0.00 1.50 0.17 0.56 0.82 0.47

Zoanthid (general) 4.83 0.17 0.17 1.72 2.69 1.56

MACROALGAE 0.00 4.67 1.50 2.06 2.38 1.38

Macroalgae (general) 0.00 4.67 1.50 2.06 2.38 1.38

DEAD CORAL WITH ALGAE 0.17 0.33 1.17 0.56 0.54 0.31



48

Table 3.4.4: Species mean density and frequency on the upper fore reef (9m) at Man-O-War Bay survey sites

Man-o-War Bay 9m Mean Density (col. m


Species Booby Is. East Booby Is. West Pirate's Reef Booby Is. East Booby Is. West Pirate's Reef

Agaricia agaricites 0 0 0.125 0 0 12.5

Colpophyllia natans 0 0.625 0 0 50 0

Diploria labyrinthiformis 0.125 0 0.25 12.5 0 25

Diploria strigosa 0.375 1 0.25 37.5 25 25


Millipora alcicornis 0.5 0.25 0.125 25 0 0


Montastrea faveolata 0 1.125 0 0 37.5 0

Porites astreoides 0 0.25 0 0 12.5 0

Porites porites 0 0.25 0 0 12.5 0

Siderastrea siderea 0 1.25 0 0 37.5 12.5

Briareum 0 0 0.125 0 0 12.5

Erythropodium 0.375 0.75 0.125 12.5 50 25

Gorgonian (general) 3.875 2.5 4.75 75 50 100

Pseudopterogorgia 1 0.125 1.5 62.5 12.5 87.5

Sponge 3.75 0.625 2.5 62.5 37.5 75

49

Table 3.4.5: Species cover on the mid fore reef (16m) at Man-O-War Bay survey site

Man-o-War Bay 16m

MAJOR CATEGORY (% of transect) Species (% of transect) Booby Is. East MEAN










SPONGES 3.00 3.00

ZOANTHIDS 4.33 4.33




DEAD CORAL WITH ALGAE 0.83 0.83



Table 3.4.6: Species mean density and frequency on the mid fore reef (16m) at Man-O-War

Bay survey site



Species Booby Is. East Booby Is. East


Millipora alcicornis 1.75 75

Montastraea cavernosa 0.5 50





Sponge 3.625 87.5

50

Table 3.4.7: Species cover on the lower fore reef (21m) at Man-o-War Bay survey site

Man-o-War 21m

MAJOR CATEGORY (% of transect) Species (% of transect) Booby Is. East MEAN


Agaricis lamarcki 0.33 0.33







SPONGES 2.83 2.83

ZOANTHIDS 0.33 0.33




DEAD CORAL WITH ALGAE 0.33 0.33



Table 3.4.8: Species mean density and frequency on the lower fore reef (21m) at Man-o-War

Bay survey site



Species Booby Is. East Booby Is. East

Agaricis lamarcki 0.125 12.5

Meandrina meandrites 0.25 25

Millipora alcicornis 0.75 50



Pseudopterogorgia 0.25 25

Sponge 3.875 50

51

3.5 SPEYSIDE

In Speyside the reef fringing the western coast of Goat Island and Angel Reef, was surveyed

at three separate localities, each at all four depths (Figure 2.7). At 3m, the vast majority of

the benthic structure was composed of just bare rock with a total mean live cover of only

17%. Generally low frequencies and densities of hard corals, gorgonians and sponges is a

reflection of this (Table 3.5.2). Hard corals had a low mean cover of 7.6±5.6% with 10

different species recorded (Table 3.5.1). The northern most site on Angel reef however had

14% hard coral cover, more than twice that of the other two sites i.e. in line with the jetty

(about 4% cover) and to the south of the reef (5% cover). Zoanthids, namely Palythoa

caribaeorum, occupied another 7±5.4%, on average, of the 3m zone but seemed to be more

concentrated towards the two northernmost sites with there being only 0.8% coverage at the

southern locality. A mean of about 2±1.1% represented gorgonian cover with very minimal

cover (mean 0.3±0.5%) being attributable to encrusting coralline algae.

Mean hard coral cover at 9m was relatively high (34.5±13%) and 18 different species were

recorded (Table 3.5.3). The reefs at this depth appeared healthy and diverse. The hard coral

cover at the southern site was especially high with nearly 50% of the substrate being covered

by hard corals. However, the assemblage at this site was dominated by Madracis mirabilis

(33.5% cover, 2.875 col.m-2

density) (Table 3.5.4). At the jetty site the 25% hard coral cover

was dominated by Montastrea faveolata (10.2% cover, 3.25 col.m-2

density) and at the

northern site (28.7% hard coral cover), Montastarea annularis (11% cover, 2.25 col.m-2

density) dominated. On this reef, in the upper fore reef zone about fifteen percent (15±7.7%)

of the benthos was occupied by gorgonians. Again there were variations from site to site with

the highest cover being found at the jetty site (23.8% cover). The southern and northern

surveying localities had 11.7% and 9.5% gorgonian cover respectively. The remainder of the

approximately 55% total mean live cover was shared among species in the encrusting

coralline algae (2.8±3.8% mean cover), sponge (1.9±1.7% mean cover) and macroalgae

(0.3±0.5% mean cover) categories.

52

Table 3.5.1: Species cover on the reef crest (3m) at Angel Reef, Speyside survey sites

Speyside 3m

MAJOR CATEGORY (% of transect) Species (% of transect) Angel Reef North Angel Reef Jetty Angel Reef South MEAN STD. DEV. STD. ERROR

HARD CORAL 14.00 3.83 5.00 7.61 5.56 3.21


Diploria clivosa 0.00 2.00 1.67 1.22 1.07 0.62



Favia fragum 0.17 0.17 0.17 0.17 0.00 0.00


Millipora complanata 0.67 0.00 0.00 0.22 0.38 0.22




GORGONIANS 1.33 3.33 1.50 2.06 1.11 0.64

Erythropodium 0.33 0.00 0.00 0.11 0.19 0.11


ZOANTHIDS 9.83 10.50 0.83 7.06 5.40 3.12

Palythoa sp. 9.83 10.50 0.83 7.06 5.40 3.12



53

Table 3.5.2: Species mean density and frequency on the reef crest (3m) at Angel Reef, Speyside survey sites

Speyside 3m Mean Density (col. m


Species Angel Reef North Angel Reef Jetty Angel Reef South Angel Reef North Angel Reef Jetty Angel Reef South

Agaricia agaricites 0 0 0.125 0 0 12.5

Diploria clivosa 0 0.125 0.25 0 12.5 12.5

Diploria labyrinthiformis 0 0.125 0 0 12.5 0

Diploria strigosa 0 0.125 0.125 0 12.5 12.5

Favia fragum 0.125 0.125 0.75 12.5 12.5 12.5

Millipora alcicornis 1.125 0 1.75 37.5 0 50

Millipora complanata 0.125 0 0 12.5 0 0

Porites astreoides 0.625 0 0.625 25 0 25

Siderastrea radians 0.75 0.125 0 25 12.5 0


Erythropodium 0.25 0 0 25 0 0

Gorgonian (general) 0.125 0.875 0.625 12.5 50 25

Sponge 0 0 0.25 0 0 0

54

Table 3.5.3: Species cover on the upper fore reef (9m) at Angel Reef, Speyside survey sites

Speyside 9m


HARD CORAL 28.71 25.33 49.33 34.46 12.99 7.50





Leptoseris cucullata 0.17 0.00 0.00 0.06 0.10 0.06


Madracis mirabilis 2.00 0.00 33.50 11.83 18.79 10.85



Montastraea annularis 11.02 5.00 0.33 5.45 5.36 3.09



Mycetophyllia aliciae 0.00 0.17 0.00 0.06 0.10 0.06


Porites porites 0.00 0.00 0.17 0.06 0.10 0.06

Scolymia cubensis 0.17 0.17 0.00 0.11 0.10 0.06



GORGONIANS 9.52 23.83 11.67 15.01 7.72 4.46

Briareum 1.34 6.50 2.67 3.50 2.68 1.55

Erythropodium 6.51 8.67 2.33 5.84 3.22 1.86



SPONGES 0.17 3.50 2.00 1.89 1.67 0.96

ZOANTHIDS 0.00 0.83 0.00 0.28 0.48 0.28

Palythoa sp. 0.00 0.83 0.00 0.28 0.48 0.28

MACROALGAE 0.83 0.00 0.00 0.28 0.48 0.28





55

Table 3.5.4: Species mean density and frequency on the upper fore reef (9m) at Angel Reef, Speyside survey sites




Agaricia agaricites 2.625 1.125 0.875 62.5 37.5 25

Colpophyllia natans 0.125 0 0.125 12.5 0 12.5

Diploria labyrinthiformis 0 0 0.125 0 0 12.5

Diploria strigosa 0.375 0.125 0.25 25 12.5 12.5

Leptoseris cucullata 0.125 0 0 12.5 0 0

Madracis decactis 0.125 0.25 0 12.5 12.5 0

Madracis mirabilis 0.5 0 2.875 37.5 0 100

Meandrina meandrites 0 0.125 0 0 12.5 0

Millipora alcicornis 0 0.125 0.25 0 0 12.5

Montastraea annularis 2.25 2.125 0.125 62.5 50 12.5

Montastraea cavernosa 0 0.25 0.125 0 0 12.5

Montastrea faveolata 0 3.25 0 0 62.5 0

Mycetophyllia aliciae 0 0.125 0 0 12.5 0

Porites astreoides 0.625 1 0.25 50 50 12.5

Porites porites 0 0 0.125 0 0 12.5

Scolymia cubensis 0.375 0.125 0 12.5 12.5 0

Siderastrea radians 0 0.5 0.375 0 25 37.5

Siderastrea siderea 0.25 0.375 0.5 25 25 37.5

Briareum 0.125 1.25 1.875 12.5 37.5 75

Erythropodium 1 2.125 0.75 62.5 75 50

Gorgonian (general) 1.25 2.25 2.875 37.5 87.5 75

Pseudopterogorgia 0 0.375 0.125 0 25 12.5

Sponge 0.25 1.75 3.125 12.5 50 50

Favia fragum 0 0 0.125 n/a n/a n/a

56

At 16m, hard coral cover was the highest category with about 24±11.2% mean cover (Table 3.5.5),

similar to 9m. Hard coral cover decreased southwards; the highest cover (36.7% cover) was on the

northern site. It fell to 18.5% at the jetty surveying site and then to about 16% at the southern

locality. On average, Diploria strigosa (6±8.8% mean cover), Madracis mirabilis (5.2±4.6% mean

cover), Agaricia agaricites (4.4±1.2% mean cover) and Porites astreoides (3.8±3.3% mean cover)

were the species of hard corals that dominated the sites at this depth. A few large colonies of

Diploria strigosa stood out at the northern survey site having 16.2% cover (0.25 col.m-2

density) at

this locality in particular (Table 3.5.6).

Gorgonian cover represented on average about 10±6.3% of the benthic cover at 16m on Angel reef

decreasing from the mean of 15±7.7% recorded at 9m. Mean encrusting coralline algae and mean

sponge cover, on the other hand, showed considerable increases with species in these categories

colonising 8.5±1.6% and 7.2±2.5% of the reef respectively. Unlike hard coral cover, sponge cover

increased steadily from north to south along Angel reef. Macroalgae, although almost negligible

(0.2±0.2% mean cover), made up the remainder of the nearly 50% total mean live cover.

Twenty one metres (21m) depth was at the vertical extent of the reef, lying at the reef/sand interface

especially at the northern and southern sites. Consequently, total mean live cover at this depth was

lower than at 9m and16m (38.6%). The paucity of suitable substrate for colonisation was reflected in

the low mean hard coral cover (Table 3.5.7). About 13.4±8% of the benthic surface was occupied by

the 12 hard coral species. The hard coral coverage at the jetty site – where the reef seemed to extend

a bit deeper – represented the only real fairly substantial amount of coverage (22.2% hard coral

cover). In addition, Agaricia sp. seemed to out-compete other hard corals at the 21m survey sites,

having the highest densities (Table 3.5.8) and representing about half of the total mean hard coral

cover.

Sponges continued their trend of increasing coverage with depth as there was 12.3±2.2% mean

coverage in this lower fore reef zone. Mean gorgonian cover declined once again to about 8±2.9%.

Encrusting coralline algae, which cannot colonise sandy substrates, had 4.7±3% mean cover and, as

was the norm with this reef, mean macroalgal cover was again minimal (0.2±0.2%).

57

Table 3.5.5: Species cover on the mid fore reef (16m) at Angel Reef, Speyside survey sites

Speyside 16m


HARD CORAL 36.67 18.50 16.17 23.78 11.22 6.48


Agaricis lamarcki 0.00 0.00 0.33 0.11 0.19 0.11



Eusmilia fastigiata 0.00 0.33 0.83 0.39 0.42 0.24









Scolymia cubensis 0.00 0.17 0.00 0.06 0.10 0.06



GORGONIANS 4.83 17.17 8.50 10.17 6.33 3.66

Briareum 0.00 2.83 1.83 1.56 1.44 0.83

Erythropodium 1.83 3.17 1.17 2.06 1.02 0.59



SPONGES 4.33 8.00 9.17 7.17 2.52 1.46

MACROALGAE 0.17 0.00 0.33 0.17 0.17 0.10





58

Table 3.5.6: Species mean density and frequency on the mid fore reef (16m) at Angel Reef, Speyside survey sites




Agaricia agaricites 2.5 3.125 2.5 75 87.5 75

Agaricis lamarcki 0 0 0.875 0 0 25

Colpophyllia natans 0.125 0 0 12.5 0 0

Diploria strigosa 0.25 0.125 0.375 25 12.5 37.5

Eusmilia fastigiata 0 0.25 0.125 0 12.5 12.5


Madracis decactis 0.125 0 0 12.5 0 0

Madracis mirabilis 1 0.5 1 62.5 25 50


Millipora alcicornis 0 0 0.125 0 0 12.5

Montastraea cavernosa 0 0.375 0 0 25 0

Montastrea faveolata 0.375 0.375 0 25 12.5 0

Porites astreoides 1 0.875 0.875 50 50 50

Scolymia cubensis 0.25 0.125 0 0 12.5 0

Siderastrea radians 0 0.625 0.25 0 25 25

Siderastrea siderea 0.125 0.375 0.25 25 12.5 25

Briareum 0 0.75 0.25 25 12.5 12.5

Erythropodium 0.75 1.375 0.5 37.5 75 50

Gorgonian (general) 1.375 3.875 1.375 50 87.5 50

Pseudopterogorgia 0.125 0 0.5 0 0 50

Sponge 2.625 4.625 3.375 87.5 100 100

59

Table 3.5.7: Species cover on the lower fore reef (21m) at Angel Reef, Speyside survey sites

Speyside 21m


HARD CORAL 11.50 22.17 6.50 13.39 8.00 4.62


Agaricis lamarcki 1.50 5.17 3.50 3.39 1.84 1.06











GORGONIANS 5.33 11.00 7.67 8.00 2.85 1.64

Briareum 0.00 1.83 0.00 0.61 1.06 0.61

Erythropodium 0.00 2.00 0.00 0.67 1.15 0.67



SPONGES 14.83 10.50 11.67 12.33 2.24 1.29

MACROALGAE 0.17 0.33 0.00 0.17 0.17 0.10





60

Table 3.5.8: Species mean density and frequency on the lower fore reef (21m) at Angel Reef, Speyside survey sites




Agaricia agaricites 2.25 2.375 0.75 75 75 37.5

Agaricis lamarcki 0.375 0.75 0.75 25 37.5 37.5

Colpophyllia natans 0 0.125 0 0 12.5 0


Madracis decactis 0.375 0.5 0.125 25 25 12.5

Madracis mirabilis 0.25 0.125 0 12.5 0 0

Meandrina meandrites 0.375 0.75 0.25 25 50 12.5


Montastrea faveolata 0.25 0.125 0 25 12.5 0

Porites astreoides 1 0.625 0 37.5 25 0

Siderastrea radians 0 0.25 0 0 12.5 0

Siderastrea siderea 0.625 0.375 0 37.5 25 0

Briareum 0 0.375 0 0 37.5 0

Erythropodium 0 0.625 0 0 25 0

Gorgonian (general) 1.875 2.25 1.75 62.5 75 75

Pseudopterogorgia 0.25 0.25 0.625 25 12.5 25

Sponge 4.25 3.875 3.375 100 100 100

Scolymia cubensis 0.125 0 0 n/a n/a n/a

61

3.6 LA GUIRA BAY

This bay is on the Atlantic side of the island and is a high energy environment with a fairly strong

current causing drift parallel to the coastline in a westerly direction. Reefs here were intermittent;

interspersed by patches of coral rubble, Palythoa beds, Thalassia patches and sand. Due to the

structures of the reefs, surveys were only conducted at one depth – 9m – as the reef sites started at

depths ranging from 4-6m and descended to about 12-15m.

Two sets of eight photoquadrats were taken at the dive site known as ‘Cove Ledge’ (Figure 2.6).

Here macroalgae had a mean coverage of 35.6±9.8% (Table 3.6.1) - making up more than half of the

total mean live cover of the area (70.4%). This macroalgae was observed overgrowing and out

competing some of the sponges in the locality. Nonetheless, sponge cover and density were still

fairly high, having averages of 15.1±0.6% cover and 8.06 col.m-2

(Table 3.6.3) respectively.

Zoanthids, namely Palythoa caribaeorum, were the only other really significant component of the

benthos, occupying a mean of 9.2±12%. Hard coral (4.7% cover), gorgonians (4% cover) and

encrusting coralline algae (1.8±0.7%) all contributed to a lesser extent to the total mean live cover.

Two sets of eight photo-transects were also conducted at the ‘Majeston’ dive site (Figure 2.6). At

this locality the gorgonian category dominated with a mean cover of 35.3±5.9% (Table 3.6.2). This

gorgonian cover however was made up almost entirely by Erythropodium caribaeorum which

colonised, on average, 34.8±6% of the benthic surface. Macroalgae also had a high mean coverage

(21.6±11.4%) with Dictyota sp. and, to a lesser extent, an unidentified reddish-gray delicate plant

with a fluffy appearance and soft texture, making up the vast majority of the macroalgal species

assemblage. Mean encrusting coralline algae coverage was much higher here than at ‘Cove Ledge’

with 10.8±2.6% mean cover. Total number of hard coral species recorded also varied between

‘Majeston’ and ‘Cover Ledge’. At ‘Cove Ledge’ 10 species were recorded whereas only half that

number was observed at ‘Majeston’. In addition, while no one species dominated at ‘Cove Ledge’,

of the 7.8±4.1% mean hard coral coverage at ‘Majeston’ 86% of this was Siderastrea siderea

colonies (6.7±3,8% mean cover). It was also noted during analysis that an average of 22.7±32% of

the S. siderea colonies here was affected by Dark Spot Disease. Mean sponge cover at ‘Majeston’

was only 3.2±1.2% making up a very small part of the almost 79% total mean live cover.

62

Table 3.6.1: Species cover on the upper fore reef (9m) at Cove Ledge, La Guira survey sites

Cove Ledge 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Cove Ledge 1 Cove Ledge 2 MEAN STD. DEV. STD. ERROR

HARD CORAL 4.67 4.67 4.67 0.01 0.00


Diploria clivosa 0.33 0.33 0.33 0.00 0.00









GORGONIANS 4.00 4.01 4.00 0.00 0.00

Briareum 0.17 0.67 0.42 0.35 0.25

Erythropodium 1.17 1.50 1.33 0.24 0.17



SPONGES 14.67 15.53 15.10 0.61 0.43

ZOANTHIDS 0.67 17.70 9.18 12.04 8.51

Palythoa sp. 0.67 17.70 9.18 12.04 8.51

MACROALGAE 42.50 28.71 35.61 9.75 6.89





63

Table 3.6.2: Species cover on the upper fore reef (9m) at Majeston, La Guira survey sites

Majeston 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Majeston 1 Majeston 2 MEAN STD. DEV. STD. ERROR

HARD CORAL 4.83 10.67 7.75 4.12 2.92


Agaricia grahamae 0.00 0.17 0.08 0.12 0.08




GORGONIANS 39.50 31.17 35.33 5.89 4.17

Erythropodium 39.00 30.50 34.75 6.01 4.25


SPONGES 4.00 2.33 3.17 1.18 0.83

MACROALGAE 13.50 29.67 21.58 11.43 8.08

Dictyota 10.67 20.50 15.58 6.95 4.92


ENCRUSITNG CORALLINE ALGAE 12.67 9.00 10.83 2.59 1.83


64

Table 3.6.3: Species mean density and frequency on the upper fore reef (9m) at the Cove Ledge and Majeston, La Guira survey sites

La Guira Bay 9m Mean Density (col. m


Species Cove Ledge 1 Cove Ledge 2 Majeston 1 Majeston 2 Cove Ledge 1 Cove Ledge 2 Majeston 1 Majeston 2

Agaricia agaricites 0 0.125 0.125 0.25 0 12.5 12.5 25

Agaricia grahamae 0 0 0 0.125 0 0 0 12.5

Diploria clivosa 0.125 0.125 0 0 12.5 12.5 0 0

Diploria strigosa 0.75 0 0.25 0 50 0 25 0

Madracis decactis 0 0.625 0 0 0 25 0 0

Meandrina meandrites 0 0.125 0 0 0 12.5 0 0

Millipora alcicornis 0.25 0.125 0 0 12.5 12.5 0 0

Millipora complanata 0 0.25 0 0 0 12.5 0 0

Montastraea cavernosa 0 0.25 0 0 0 25 0 0

Porites astreoides 0 0.25 0.125 0.5 0 12.5 12.5 37.5

Siderastrea siderea 0.25 0.375 1.125 1 25 25 50 75

Briareum 0.25 0.375 0 0 12.5 25 0 0

Erythropodium 0.875 1.125 8.875 8.625 50 37.5 100 100

Gorgonian (general) 2 0.375 0.375 0.5 62.5 12.5 25 25

Pseudopterogorgia 0 0.625 0 0 0 25 0 0

Sponge 8.625 7.5 0.875 0.625 87.5 87.5 50 50

65

Five sets of eight photoquadrats were conducted along the length of the ‘Flying Reef’ dive site

(Figure 2.6). This reef, which is about 1.5km long, had a total mean live cover of about 48%,

lower than the other two dive sites surveyed in La Guira Bay. No major group showed

dominance on this reef (Table 3.6.4). Macroalgae had the highest coverage (12.9±5.4% mean

cover) with Dictyota sp. once again being very prolific (10.7±4.6% mean cover). The seventeen

different hard coral species recorded had a mean cover of 11.9±6% of the benthos, with

Sideratrea siderea (4.3±1.7% mean cover) and Montastrea faveolata (2±4.3% mean cover), in

combination, accounting for more than half this figure. Gorgonians also featured in similar

proportions to macroalgae with a mean cover of 10.1±2%. Sponges had an average coverage of

7.6±2.3% and consistently high densities at all the ‘Flying Reef’ surveying localities (Table

3.6.5). Encrusting coralline algae (3.7±2.4% mean cover) and zoanthids (1.9±1.4% mean cover)

however, found it hard to colonise the predominantly sand/rubble bottom.

66

Table 3.6.4: Species cover on the upper fore reef (9m) at Flying Reef, La Guira Bay

Flying Reef 9m

MAJOR CATEGORY (% of transect) Species (% of transect) Flying Reef 1 Flying Reef 2 Flying Reef 3 Flying Reef 4 Flying Reef 5 MEAN STD. DEV. STD. ERROR

HARD CORAL 5.33 9.33 10.00 21.33 13.50 11.90 6.02 2.69

Agaricia agaricites 0.33 0.00 2.00 0.33 0.17 0.57 0.81 0.36

Agaricia grahamae 0.17 0.00 0.00 0.00 0.00 0.03 0.07 0.03

Dichocoenia stokesi 0.00 0.50 0.00 0.00 0.00 0.10 0.22 0.10

Diploria labyrinthiformis 0.00 0.50 0.00 0.00 0.00 0.10 0.22 0.10

Diploria strigosa 1.33 2.83 0.00 2.00 3.33 1.90 1.31 0.59

Eusmilia fastigiata 0.00 0.00 0.17 0.00 0.00 0.03 0.07 0.03

Madracis decactis 0.00 0.17 0.50 0.00 0.00 0.13 0.22 0.10

Meandrina meandrites 0.17 0.00 0.67 0.00 0.17 0.20 0.27 0.12

Millipora alcicornis 0.00 0.17 1.17 1.83 2.33 1.10 1.02 0.46

Montastraea annularis 0.00 0.50 0.00 0.00 0.00 0.10 0.22 0.10

Montastraea cavernosa 1.50 0.00 0.67 1.67 1.00 0.97 0.67 0.30

Montastrea faveolata 0.00 0.00 0.33 9.67 0.00 2.00 4.29 1.92

Porites astreoides 0.00 0.00 0.17 0.00 0.00 0.03 0.07 0.03

Porites porites 0.00 0.00 0.00 0.50 0.17 0.13 0.22 0.10

Scolymia cubensis 0.00 0.00 0.17 0.00 0.00 0.03 0.07 0.03

Siderastrea radians 0.00 0.83 0.00 0.00 0.00 0.17 0.37 0.17

Siderastrea siderea 1.83 3.83 4.17 5.33 6.33 4.30 1.70 0.76

GORGONIANS 8.83 8.83 10.00 13.50 9.33 10.10 1.96 0.88

Briareum 0.00 0.00 1.00 0.17 0.67 0.37 0.45 0.20

Erythropodium 0.00 1.00 2.50 6.67 1.17 2.27 2.62 1.17

Gorgonian (general) 8.83 7.83 6.00 6.33 7.33 7.27 1.15 0.51

Pseudopterogorgia 0.00 0.00 0.50 0.33 0.17 0.20 0.22 0.10

SPONGES 8.50 7.50 9.50 3.67 8.67 7.57 2.29 1.03

ZOANTHIDS 0.50 0.50 3.33 3.00 2.33 1.93 1.36 0.61

Palythoa sp. 0.50 0.50 3.33 3.00 2.33 1.93 1.36 0.61

MACROALGAE 6.50 16.00 17.33 7.50 17.00 12.87 5.39 2.41

Dictyota 5.00 14.83 13.33 6.33 13.83 10.67 4.62 2.07

Macroalgae (general) 1.50 1.17 4.00 1.17 3.17 2.20 1.30 0.58

DEAD CORAL WITH ALGAE 0.00 0.17 0.00 0.00 0.00 0.03 0.07 0.03

ENCRUSTING CORALLINE ALGAE 2.17 3.67 4.33 7.17 1.00 3.67 2.35 1.05

SAND, PAVEMENT, RUBBLE 68.17 54.00 45.50 43.83 48.17 51.93 9.86 4.41

67

Table 3.6.5: Species mean density and frequency on the upper fore reef (9m) at Flying Reef, La Guira survey

Flying Reef 9m Mean Density (col. m


Species Flying Reef 1 Flying Reef 2 Flying Reef 3 Flying Reef 4 Flying Reef 5 Flying Reef 1 Flying Reef 2 Flying Reef 3 Flying Reef 4 Flying Reef 5

Agaricia agaricites 0.625 0 1.875 0.375 0.25 25 0 37.5 25 12.5

Agaricia grahamae 0.125 0 0 0 0 12.5 0 0 25 0

Dichocoenia stokesi 0 0.125 0 0 0 0 12.5 0 0 0

Diploria labyrinthiformis 0 0.375 0 0 0 0 12.5 0 0 0

Diploria strigosa 0.5 1.25 0.125 0.375 0.5 50 62.5 0 25 37.5

Eusmilia fastigiata 0 0 0.125 0 0 0 0 12.5 0 0

Madracis decactis 0 0.125 0.375 0.125 0 0 12.5 25 0 0

Meandrina meandrites 0.125 0 0.125 0 0.125 12.5 0 12.5 0 12.5

Millipora alcicornis 0.125 0.375 1.5 0.75 1.125 0 12.5 50 37.5 62.5

Montastraea annularis 0 0.125 0 0 0 0 12.5 0 0 0

Montastraea cavernosa 0.625 0 0.375 0.625 1 62.5 0 37.5 37.5 37.5

Montastrea faveolata 0 0 0.125 1.75 0 0 0 12.5 25 0

Porites astreoides 0 0 0.125 0 0 0 0 12.5 0 0

Porites porites 0 0 0 0.125 0.125 0 0 0 12.5 12.5

Scolymia cubensis 0 0 0.125 0 0 0 0 12.5 0 0

Siderastrea radians 0 0.125 0 0 0 0 12.5 0 0 0

Siderastrea siderea 0.375 1 0.75 1.375 1.25 37.5 50 37.5 50 62.5

Briareum 0 0 0.125 0.125 0.125 0 0 12.5 12.5 12.5

Erythropodium 0 0.625 1.75 1.875 0.875 0 37.5 50 75 50

Gorgonian (general) 3.625 1.75 2.875 2.875 3.875 100 87.5 87.5 75 87.5

Pseudopterogorgia 0.125 0 0.25 0.25 0.125 0 0 25 25 12.5

Sponge 4.875 5 4.75 3 3.5 87.5 87.5 87.5 75 75

68

3.7 RAPID ASSESSMENT

A rapid assessment was carried out at each survey depth at all reef sites to yield information on

Diadema antillarum presence/absence, coral disease and bleaching occurrence and recruitment.

From the surveys it appeared that D. antillarum densities remain far below the pre-mortality

estimates. This sea urchin was observed at Arnos Vale, Buccoo’s Eastern Reef, Speyside’s Angel

Reef and Flying Reef in La Guira Bay. It must be noted though that Diadema observed at Arnos

Vale were not found along the rapid assessment transects. However, in one area of the western

locality’s reef crest the numbers were very high (> 50). Here, densities were reminiscent of pre-

mortality estimates of 5 urchins.m-2

(Laydoo 1985g). A total of only thirteen (13) D. antillarum

were observed between the other remaining locations.

Rapid assessment also revealed that several coral diseases occurred throughout all the reef areas

in Tobago. Yellow Band disease affecting Montastrea, Dark Spots disease affecting Siderastrea,

Aspergillosis affecting Gorgonia, Black band disease and bleaching were all recorded. However,

these diseases were not prevalent in most locations. The exceptions to this would be Yellow

Band disease in some forereef zones of Buccoo and Culloden, and Dark Spots disease in La

Guira Bay.

The observations with regards to recruitment were also interesting. Generally, hard coral

recruitment was seen in all areas where rapid assessment was carried out except some of the reef

crest sites of Buccoo and Man-o-War Bay. However, sponge recruitment was observed far more

regularly. Of particular note were the species of scleractinian recruits. Overall, it could be said

that the major reef builders on Tobago reefs e.g. Montastrea sp., Siderastrea sp. and Diploria

sp., were observed to be recruiting. However, in comparison, far more abundant in hard coral

recruitment were non-framework builders, especially Agaricia.

69

4.0 DISCUSSION

In this study, reefs fringing Tobago do not appear to strictly adhere to the observations reported

by Gardner et al. (2003) that average hard coral cover across the entire Caribbean basin has seen

massive declines – by 80% - since the late 1970s. Conversely, the various reefs around Tobago

seem to be experiencing mixed fortunes. Mean hard coral declines were seen in some areas e.g.

Buccoo and Culloden, whereas improvements were observed in others e.g. Arnos Vale and

Speyside, when compared with the Laydoo surveys of the 1980s (Figure 4.1). The other re-

surveyed area, Man-o-War Bay, showed no overall trend with slight improvements in mean hard

coral cover at the mid (16m) and lower (21m) fore reef zones while there was large reduction at

the upper (9m) fore reefs. The coupled methodology between the 1985 and current study allowed

for benthic cover differences at the various depths over the last 20 years to be revealed. When

looking at the data from this point of view, the upper reef crest zones (3m) were particularly

interesting showing very significant decreases in hard coral cover in all surveyed areas except

Arnos Vale and Man-o-War Bay.

Several explanations may be proposed for the hard coral decline that took place almost island

wide at 3m. However, it is likely that different areas of Tobago have been affected by varying

synergies of factors. For example, in Culloden, where hard coral cover has seen an almost 85%

reduction at the reef crest - from 29.1% in 1985 to 4.5% in 2009 - the loss of the major reef

framework building coral Acropora palmata can be cited as the chief reason for this. In the

Laydoo (1985c) study 28% of the benthic cover here was made up A. palmata colonies.

However, it seems that inability to recover from the region wide phenomenon of white band

disease that decimated populations of this species (Laydoo 1985g) has left a void in the hard

coral assemblage at the reef crest that has yet to be filled by other hard corals. Palythoa

caribaeorum and coralline algae are now the dominant space occupiers. The situation is unlike

that found in Arnos Vale where, although A. palmata dominated the hard coral fauna in the early

1980s, post die off, Millipora complanata and to a lesser extent Diploria clivosa has filled the

void.

70

Figure 4.1: Hard coral cover comparisons between the Laydoo (1985a-e) surveys and the current

assessments

71

A. palmata die off can also be applicable to Buccoo’s approximate 75% mean hard coral decline

in the 3m zone. Like at Culloden and Arnos Vale, this species was a major constituent of

Buccoo’s reef crests as well. The proliferation of Palythoa caribaeorum and macroalgae

however, also suggests that eutrophication has some part to play (Lapointe 2003) as hard corals

are out competed for space and/or fail to recruit. The losses of grazers e.g. Diadema antillarum -

which have failed to recover to densities anywhere near to pre-mortality estimates (Laydoo

1985f) - may have also exaggerated these competition effects.

Eliciting explanations for the declines at Speyside’s reef crest are not as clear cut however. In the

1980s, unlike at the aforementioned reefs, A. palmata was not a major part of the hard coral

assemblage. Species such as Porites astreoides, Siderastrea siderea and Millipora alcicornis

showed dominance instead (Laydoo 1985e). In addition, no studies in the Speyside area have

pointed to eutrophication of the waters. Diadema antillarum population has remained at low

levels since the die off but this by itself does not elicit a mechanism for scleractinian loss

(Aronson & Precht 2006). This is especially so in light of no eutrophication and low macroalgae

cover as observed in this study. As a matter of fact, it was found that the majority (83%) of the

benthic surface was just bare rock. It is therefore extremely difficult to explain the almost 57%

loss in mean hard coral cover in this zone over the last two decades.

This is especially so when, from this study, the fore reef zone (9-21m) has seen mean hard coral

cover increase. The fact that these increases have taken place tends to negate the possibility of

recruitment rates being low due to, for example, changing current patterns or inappropriate

settling surfaces. One possible explanation for the reduction though, is that the areas surveyed at

3m are where tourists on glass bottom boats are brought to snorkel. While these boats use fixed

moorings and may not anchor on the reef, other tourist related activities could have caused

sufficient disturbance that may be reflected in a decline of hard coral cover. However, the IMA

(2002) report aimed at formulating a management plan for Speyside did stress that where these

boats presently operate is deep enough to prevent damage to the reef by snorkelling and walking

on corals. This explanation therefore, is tenuous at best.

72

Like decreases in hard coral cover at Speyside’s reef crest, it is also extremely difficult to explain

the opposite - increases in cover at various reef zones surveyed around the island. The reef crest

and fore reefs of Arnos Vale, the entire fore reef of Speyside and the mid and lower fore reefs of

Man-O-War Bay all showed mean hard coral cover increase. In light of the ever increasing

negative pressures on Tobago’s marine environment through a myriad of factors such as

population increases, rampant coastal development, changing watershed and land use that

promotes terrestrial run off, climate change and its associated impacts and increased tourism

volume and artisanal fishing effort (Mora 2008; Burke & Maidens 2004), improvements in

scleractinian cover on reefs is certainly not in keeping with the almost ubiquitous worldwide

trend of reef decline. This is especially so given the fact that little to no safeguarding measures

e.g. establishment of Marine Protected Areas (MPA) or Coastal Zone Management Plans, have

been instituted to protect reefs in those areas where hard coral cover increase has been evident.

On the contrary, Buccoo, where the only MPA in the country exists, has seen declining fortunes

in some areas.

Therefore, at the risk of sounding pessimistic, it is unlikely that the improvements reflected in

the data have occurred in reality. Rather, the data has to be interpreted with caution. It appears

more feasible that better technology e.g. higher resolution coloured photographs and the

application of the user friendly CPCe program, has made the present study more accurate. The

limitations placed on researchers in the 1980s may have led to an underestimation of hard coral

cover and as such, a pseudo improvement in some areas when comparing the both results from

the both time periods. The large data gap that exists in terms of time elapsed between the 1985

study and similar assessments like this one only serves to increase the level of uncertainty when

trying to make comparisons and draw conclusions. If hard coral cover was underestimated in the

Laydoo (1985a-e) surveys however, it would suggest that the declines observed in areas such as

Buccoo and Culloden have been even more drastic.

In South-west Tobago, land use change and coastal development has been freely ongoing for

several decades. Many commentators over the years have offered cautionary words about this

situation as it pertains to Buccoo Reef’s survival and resilience (Goreau 1967; Kenny 1976;

Laydoo 1985a; IMA 1994). Nonetheless, cumulative terrestrial inputs such as fertilizer from golf

73

courses and agriculture, sewage from hotel and domestic wasters and sediments in run off from

impacted catchments and coastal construction, which all negatively affect water quality and reef

health, provide the main, plausible explanations for the observed hard coral decline on Buccoo

(Wilkinson 2008).

To a lesser extent, the influence of fairly novel challenges may also be cited. For example,

Buccoo was severely affected by coral bleaching in 1998 and 2005 (O’Farrell & Day, 2005).

Although the majority of corals recovered1 high incidences of disease e.g. Yellow Band disease

observed in the current study at the upper fore reef in some surveyed areas of Buccoo, may be as

a result of increased susceptibility due to bleaching. Yellow Band disease is known to cause

partial to total colony mortality and is considered to be one the most widespread and damaging

coral diseases (Cervino et al., 2001).

Ironically, deleterious pressures on Culloden’s reefs seem to be less than that of Buccoo’s but

hard coral cover decline from 1985 to now has been far more precipitous. Whereas Buccoo had a

mean scleractinian decrease of 6.2% over the entire reef crest to lower fore reef area, Culloden’s

mean decrease was 18.1% over the same zones. In the 1980s the relatively pristine, undisturbed

coastal and marine environment of Culloden actually led Laydoo to suggest that this area could

have been used as a control site in monitoring environmental impacts at other reef localities in

Tobago (Laydoo 1985h). Today, however, the coast of Culloden and the surrounding region still

remains largely undeveloped so associated anthropogenic impacts should still be minimal –

definitely far less than in Buccoo – yet coral health has deteriorated more rapidly. A thorough

investigation into water circulation in this area needs to be conducted though, to determine if

pollutants may be coming from other areas of the coast.

It is known however, that Culloden was badly affected by bleaching in 2005 with more than 85%

of hard corals being affected (O’Farrell & Day, 2005). However, like at Buccoo, the majority of

corals recovered1. Montastrea faveolata – identified as Montastraea annularis in the Laydoo

1985 studies as several distinct Montastrea species were, in that time, considered just different

growth forms of M. annularis (Weil & Knowlton 1994) – has always been, and remains, the

1 www.buccooreef.org/coral_bleaching.html

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most dominant scleractinian on the fore reef slopes of Culloden (Laydoo 1985c). However, as

mentioned earlier, in the results of the present survey a high incidence of infection of M.

faveolata by Yellow Band disease was noted. It is therefore being postulated that disease has

been responsible for the waning hard coral health of Culloden. If this is indeed the case, declines

might have been relatively recent. The lack of regular surveys over the last two decades

however, precludes any declaration being made with respect to the nature of the decline.

It is interesting to note that Buccoo and Culloden, perhaps the only two real coral reef systems

studies (where aragonite structures have been built up over time) – the other areas can be more

considered as coral assemblages growing on rocks – are the chief areas exhibiting declining hard

coral cover. Perhaps the fates of these two ‘true reefs’ on the leeward coast of Tobago are

intertwined. Conducting paleo-reef studies may be able to shed some light on this theory.

Generally, in all the reef areas surveyed, far more hard coral species were recorded in the present

study’s photoquadrat analyses when compared to those recorded in Laydoo’s (1985a-e) analyses.

This was so even though he applied 450 random points to each 1m2 photoquadrat as opposed to

the 75 that were used in this CPCe assisted analytical method. It is unlikely that these additional

species are a reflection of increasing diversity on Tobago’s reefs as most were identified and

recorded in previous checklists done via visual surveys (Kenny 1976; Ramsaroop 1981; Laydoo

1985a-e; Laydoo 1990; IMA 1990; IMA 2002). It is perhaps, therefore, yet another illustration of

improved technology and fewer limitations yielding more accurate results in this study.

There are a few other noteworthy points to be made. Firstly, it appears that Madracis mirabilis

was misidentified in the Laydoo surveys as Madracis decactis in some areas where its coverage

was high e.g. Angel Reef’s upper and mid fore reef, Speyside and Rest House Reef’s reef crest,

Man-o-War Bay. It was recorded by Ramsaroop (1981) as Madracis asperula when he was

formulating his checklist of the coral fauna found in Man-o-War Bay. Nonetheless, the

specimens in this present surveys’ photoquadrats unmistakably match the examples of the yellow

pencil coral (Madracis mirabilis) found in Humann (1998).

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Secondly, the loss of cover from the reef building coral Siderastrea siderea is very noticeable

between the 1985 surveys and now (Table 4.1) Buccoo’s fore reef slopes went from being S.

siderea dominated to M. faveolata dominated. Here the upper fore reef experienced a 92%

decrease in average cover and both the mid and lower fore reefs an 89% average cover decrease.

At Culloden’s mid fore reef there was an average decrease of 63%. A 93% mean decline was

seen at Man-o-War Bay’s upper fore reefs and a 58% mean decline was seen at the reef crest of

Angel Reef, Speyside. Conversely, no significant S. siderea increases were observed anywhere.

It should be noted however that it is not immediately clear whether Laydoo (1985 a-e)

differentiated between S. siderea and S. radians in his surveys. If he did not, this may over

exaggerate the extent of S. siderea decline. That being said however, in these present surveys S.

radians did not really have high percentage coverage in any of the survey locations and thus the

S. siderea decline is still drastic.

Again, the lack of regular, continuous surveying over the last two decades makes accurately

commenting on the nature of this decline impossible. The same could be said for proposing a

reason for the decline. However, Dark Spots disease which, during rapid assessment, was

observed on Tobago reefs but was not prevalent, bleaching, or some other disease may be

possible factors. Coral Cay Conservation has been monitoring Tobago reefs since 2005, after the

bleaching event. They too have noticed significant loss of S. siderea, at 7m and 12m, since then

and suggest that bleaching (70% of S. siderea were bleached during the event) and subsequent

death of colonies is a major contributor to what is being observed in their surveys (pers. comm.

Jan-Willem van Bochove). S. siderea losses were the only major change in community structure

observed on the surveyed reefs between 1985 and now.

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Table 4.1: Siderastrea siderea percentage cover change between the Laydoo (1985 a,b,c,e) surveys

and the present surveys

Mean Siderastrea siderea % cover

Survey Locations 1985 2009 % Reduction in

cover

3m Speyside 3.87 1.61 58

16m Culloden 7 2.58 63

9m Buccoo 5.7 0.46 92

16m Buccoo 27.33 2.89 89

21m Buccoo 10.75 1.17 89

9m Man-o-War Bay 16.2 1.11 93

Thirdly, the quantitative surveys conducted on the reefs in La Guira Bay are the first of its kind.

The qualitative description given of Flying Reef in IMA (1990) i.e. a fore reef slope community

dominated by gorgonian, with stony corals being ubiquitous and sponges common, does not

seem to have changed much. From the data though, the percentage of bare or exposed substrate

here has increased since 1990. Also, as stated in IMA (1990), and supported by this current

study, the coral species diversity in La Guira Bay seems to be similar to that recorded around the

other reefs of Tobago.

However, further visual surveys would elicit a more detailed species checklist than that found in

the IMA (1990) report. Surveys should be carried out to update the ichthyofaunal species lists as

well. Interestingly, the rapid assessment surveys did reveal a fairly high incidence of Dark Spots

disease affecting S. siderea colonies in La Guira Bay. As mentioned earlier, for reasons

uncertain, this species has seen a drastic fall in cover on some Tobago reefs over the last two

decades. In addition to this, although Flying Reef is by far the most popular dive site in this area,

the exceedingly high macroalgal cover recorded, especially at Cove Ledge, and to a lesser extent,

Majeston, is cause for concern as these areas are also frequented by divers.

Lastly, this study was designed to repeat, where practicable, Laydoo (1985a-e) methodologies so

that one of the main objectives i.e. comparison of datasets, could be more readily carried out.

One limitation of this however, is that for fore reef zones that are extensive and gently sloping -

where average reef depth remains the same over large distances - eight photoquadrats at, for

77

example, 9m or 16m, may not be sufficient to capture the true essence of what exists at that

depth. This holds true for reefs at Culloden and Buccoo for instance. It is a concern that, for

Buccoo especially, although an approximate 6% mean hard coral decline was observed over fore

reef zones here, the aforementioned limitation may have led to an underestimation of this

decrease. Richard Hubbard (pers. comm.), who has been diving in this area since the 1980s, has

painted a grimmer picture of hard coral loss from then to now. In the future therefore, when these

assessments are carried out again, perhaps multiple photo-transects should be carried out at some

of the survey sites e.g. Western, Northern and Outer reefs, Buccoo and Culloden East and West,

to glean more adequate insight as to what can be found at these locations.

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5.0 CONCLUSIONS

Results from the assessment of the various areas around Tobago show the fates of their reefs to

be mixed in terms of comparative extent of hard coral cover over the last two and a half decades.

Mean hard coral declines were seen in some areas e.g. Buccoo and Culloden, whereas

improvements were observed in others e.g. Arnos Vale and Speyside reefs. Reefs in Man-o-War

bay, showed no overall trend with slight improvements in mean hard coral cover at the mid and

lower fore reef zones while there was a large reduction in cover at the upper fore reefs. Surveys

carried out at the reefs in La Guira bay were novel and thus comparisons could not be made.

However, macroalgae cover is of major concern especially at the Cove Ledge dive site.

When making comparisons between the 1985 surveys and those done in this study however, the

data has to be interpreted with caution. It is possible that better technology e.g. higher resolution

coloured photographs and the application of the user friendly CPCe program, has made the

present study more accurate. Limitations placed on researchers in the 1980s may have led to an

underestimation of hard coral cover. This therefore may paint a false picture, one where

improvements in hard coral cover may have been shown when in fact there were none. In

addition, where declines were apparent, the situation may be much worse.

To prevent a similar situation of uncertainty from occurring in the future as well as to further

track changes in the dynamic Tobago reef environments e.g. the decline, or alternatively,

recovery of species such as Sideratrea siderea and Diadema antillarum, it is being suggested

that monitoring be ongoing. Surveys of this kind should be carried out by the IMA on at least, a

3-4 year basis.

In addition to this, a few other projects, pertaining to Tobagonian reefs, are being proposed. The

first is a high resolution examination of the stability of the coral community structure and

demography (recruitment, growth, survivorship etc.) of major coral reef organisms on a reef with

a high degree of anthropogenic impact (Buccoo Reef) and one that is less impacted (Culloden

Reef). It is expected that, through this project, community structure will be assessed by means of

79

surveys of the same permanent transects/quadrats in order to reveal and document changes and

provide insight into the dynamics of coral reef communities.

A second proposed project entails using a Nitrogen loading model to determine the natural load

of nitrogen to the Bon Accord Lagoon and Culloden Bay, to identify the major land uses that

contribute to the nitrogen loads in these areas and to estimate the nitrogen retention provided by

the different components of the watersheds e.g. forest, wetland etc. This research could feed into

management guidelines for an Integrated Coastal Zone Plan for Buccoo Reef and Culloden Reef

especially as it relates to alleviating eutrophication of the coastal waters.

A third project was alluded to in earlier discussion – a paleo-reef study to garner historical

insights from Tobago and Trinidad’s reefs. It is anticipated that cores of old dead reef, recent

dead reef as well as modern corals from locations such as Buccoo and Culloden in Tobago and

Guayaguayare, Salybia and Chaguaramas in Trinidad will be analysed to age reefs and

reconstruct past environmental conditions.

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7.0 ACKNOWLEDGEMENTS

I would like to thank the following persons for their assistance in undertaking this project.

Fellow Researchers Dr. Rahanna Juman, Mr. Richard Hubbard and Dr. George Warner for their

advice and guidance throughout. Technicians Mr. Addison Titus, Mr. Jonathan Gomez and Mr.

Russell Rajnauth in carrying out field surveys. Boat Captain Mr. Allerson Small and his crew,

Launch Engineer Mr. Hayden Branche and Deckhand Mr. Ronald Robinson. Maps in this report

were produced by Mr. Deanesh Ramsewak and Mr. Sean Padmanabhan and Mrs. Charmain

Pontiflette-Douglas who aided in formatting this report.

coastal conservation project an assessment of the coral reefs of

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