Ecological aspects of the distribution of

reef corals in the Netherlands Antilles

by

Rolf P. M. Bak

Caribbean Marine Biological Institute, Curaçao, Netherlands Antilles

Abstract INTRODUCTION

The vertical and horizontal of the distribution patterns The Netherlands Antilles consist of two groups of

of corals and coral reefs (to a depth of 90 m) are dis- islands: the Leeward Group of , Bonaire and cussed in relation the environmental factors: to geomor- Curaçao and the Windward Group of St. Martin, phology of the bottom, available substrate, light, tur- St. Eustatius and Saba inset in fig. 1). Most bidity, sedimentation, water movement and temperature. (see of the research coral has been done in There is a general pattern which is comparable to other on reefs well-developed Caribbean reefs. However, as in other the Curaçao, with some attention being paid to areas variations are found, e.g. the depth and growth reefs of Aruba and Bonaire. The marine habitats form of Acropora palmata will depend on the degree of of the Windward are unstudied. There correla- Group relatively exposure to water movement. are strong show Leeward tions between the environmental variables and the oc- The few explorations made, the currence of coral species and their growth form, the Group of islands to be much richer in reef develop- species composition of coral communities and the charac- ment (and in speoies diversity) than St. Martin, St. ter of the coral reef. In some cases the relationship is not Eustatius and Saba. that obvious. The absence of Agaricia species at certain The factors coral distribution are points along the coast of Aruba and the dominance of governing

Sargassum on the deep bottom at some places along the well known and include: light, sedimentation, wa- windward coast of Curaçao is not yet explained. ter movement, substrate, bottom morphology and

The relative importance of the different factors in an temperature. I will attempt to show how these environmental setting is shown by a comparison of reef factors influence the actual distribution of reef cor- communities and reef habitats with a coral community of als and coral reefs, and a muddy, shallow inland bay. The community of the bay vertically horizontally,

from consists, apart the hardier coral species, of corals in the Netherlands Antilles. which are characteristic of the deep reef: Scolymia lacera,

S. cubensis and Helioseris cucullata. These corals are adapted to sedimentation and low light intensities and I. THE REEFS OF THE LEEWARD GROUP to are apparently able withstand a temperature and salin-

much broader than that of their ity range deep reef habitat. Vertical distribution

The paucity of corals and coral reef development Goreau & Wells noted that the around the islands of the Windward Group (deeper habi- (1967) geomorphol- of influences the tat included) can generally be explained by the morphol- ogy a coast strongly morphology

of the the lack of suitable substrate and the ogy sea floor, of the coral reef. Although the profiles of the coast effect of hurricanes. The exposed coasts of Saba and St. there are variable, are a number of general fea- Eustatius, being virtually unexplored, may have richer tures of the S.W. coasts of Curaçao and Bonaire: coral growth.

a cliff or a A new list of species of the Scleractinia of the Leeward steep shingle beach, gradually sloping and Windward groups, consisting of 57 species, is in- terrace in front, a drop-off at 7 to 12 m with a cluded. seaward slope varying from 45° to vertical. This

either slope may be continuous or interrupted by

an inclined terrace at a depth of 50 to 60 m.

There is generally a second vertical drop-off at !) Paper presented at the Seventh Caribbean Geological 70 to 80 m coming to an end at a sandy plain at Conference, Section 5: Caribbean Reef Systems, Gua- 80 90 this deloupe, 1-5 July 1974. to m (fig. 1). Over profile there is a

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Fig. 1. General profile of the S.W. coasts of Curaçao and Bonaire. The inset shows the geographical position of the

Netherlands Antilles in the Caribbean.

The reef at the first agaricites Fig. 2. drop-off (12 m) on the N.W. coast of Bonaire. Dominant corals are Agaricia

and Montastrea annularis. The latter species occurs in several different growth forms in the same place.

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definite series of coral communities, each one rep- waves. The relatively rapid growth of the A. palm-

above-men- increase in to 10 resenting a different response to the ata branches, mean length 6 cm a

tioned environmental factors. Of these, tempera- year in situ (Bak, unpublished observations), ac-

ture is probably not of major influence. Tempera- comodates greatly the establishment of broken-

off branches Whole ture measurements in situ from 10 to 50 m have as new colonies. colonies, being

never shown a difference over the reef in excess thrown over, cement themselves to the substrate

branches. In of 2°C. As the mean monthly temperature at 18 and start forming new upward this

in varies from of will be m depth Curaçao waters 26° to zone, where settlement young colonies

28 °C throughout the year, temperature never dif- hindered by the scouring of sand over the rocky fers significantly from optimum values. substrate and the abundance of the grazing sea

The reef be A urchin Diadema antillarum Bak & Van following zones may recognized: (cf. Eys,

shore zone, either in front of a shingle beach or in 1975), this mode of "vegetaltive" reproduction is of

Also in the front of a cliff. In the first case, the most con- particular adaptive value. barren zone,

and the of sand carried spicuous organisms are algae echinoderms, scouring the course by oscil-

few corals will be In will be while a encrusting present. lating water movement over the bottom, of the main factors of the latter case, the rock directly under the water one checking settlement

surface will be covered with algae, the zoanthid sessile organisms. In more exposed areas the bot-

Palythoa mammillosa and encrusting Diploria tom consists of coral rock and no loose debris is

In this the of corals and clivosa. These are areas of heavy stress, subject to present. case, density strong water movement and occasionally, to par- gorgonians is higher.

At 4to 5 the number of coral colonies in- tial emergence during low tides. Diploria clivosa m

and in is a typical coral of these high energy environ- creases, reaching a great density diversity ments. the area before the first drop-off (the blue edge:

6 in shal- From 1 to 4 m Acropora palmata is by far the Roos, 1964) at to 12 m. The bottom the most abundant coral and the most important cal- lower part is very sandy with isolated coral col- cium carbonate producer. The growth form of the onies and gorgonians. Here Acropora cervicornis colonies varies in relation to water movement sometimes forms extensive fields. Towards the

decreases and (Shinn, 1963; Roos, 1964, 1971) from a variety drop-off the sandy surface most of of branched forms to encrusting colonies. In this the substrate is living coral and coral rock, covered

of reef zone and in the barren zone (the next one sea- with a variety organisms {fig. 2). Among

the Montastrea ici wards) encrusting calcareous red algae (Corallina- corals, annularis, Agar a agarici- ceae), especially Porolithon pachydermum (cf. Van tes and Madracis mirabilis are most abundant.

Calcareous red become den Hoek et al., in press) are of consequence. algae more prominent

dead is in is Their mode of growing over coral rubble again. Except very rough weather, this area

influenced important in stabilizing the substrate. In some lo- not by the destructive force of the calities, these waves. algae or Millepora complanata may dominate reef formers. Over the the as drop-off, on steep slope, high coral

and The barren zone (3 to 4.5 m) is devoid of great density diversity continues, only to decrease builders. substrate consists below 35-40 quantities of reef The rapidly m. Calcareous red algae are of sand and coral fragments, the latter being abundant, often preferring the darker crevices of mostly dead Acropora palmata branches, partly the reef, e.g. the important species Hydrolithon cemented together by the crustose corallines. On- boergesenii occurs on the undersides of flattened

coral ly a few living corals and gorgonians are present. colonies. They are important reef frame

and often coral surfaces The Acropora palmata zone and the barren zone cementers protect against

both far above the normal are 1/2 wave length. penetration by boring sponges (Clionidae) (Bak,

The Acropora zone in particular takes the full unpublished observations). On the deeper part of

of the After of the force waves. a stormy period (maxi- the slope, with diminuation light intensity, Corallinaceae mum winds up to 36 knots, wind direction paral- become less cryptic and grow on

the examined broken branch- more Other sessile lel to coast) I many exposed outcrops. organisms

A. Jan Thiel of show similar in Madra- es of palmata at (S.W. coast a change microhabitat, e.g.

cis Their Curaçao). The largest fracture measured 4 by 30 pharensis. place in the holes and crev-

section. of the reef is taken cm in None fractures showed any ices of the, deep over mainly by signs of boring organisms, which shows that the sponges. This area is rarely influenced by strong only agent of destruction was the force of the water movement, light diminishes rapidly down

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Fig. 3. Alternating coral prominences and sediment rivers at a depth of 20 m on the S.W. coast of Curaçao.

the and is influence of sedi- available and sedimentation slope there a strong substrate, turbidity

variations in reef mentation. On this Slope sea slides sometimes are largely responsible for the

form found in the Antilles. On occur with a width of more than 10 m, scouring Netherlands the the whole slope of organisms. But under normal S.W. coasts of Curaçao and Bonaire these varia-

from the conditions colonies also break off and roll into tions are slight. Aside vertical differen-

addition there is decrease in coral around deeper water. In finer sediments constant- ces, some cover

down the The that be and the outlets of inland This be ly move slope. terrace may bays bays. may

present from 40-80 m consists mainly of loose either due to limited setdement of planulae be-

of col- sediments, with some rocky outcrops carrying cause of sedimentation, reduced growth isolated coral colonies. onies because of reduced light intensity, or pol-

Depending on the geomorphology of the bottom, lution effects. structures have developed resembling the descrip- The S.W. coast of Aruba is mainly a sandy flat

with few In tion of the Jamaican fore reef slope (Goreau & populated corals. some places (e.g.

coral the into bot- Goreau, 1973). Large outcrops growing over Arashi) sandy flat changes a shingle the edge of the drop-off are separated by rivers tom about 1 km off-shore at a depth of 20 m. On

this loose sediment small coral colonies oc- of sediments and loose material, 15-30 m wide many

sediments flow In where is (fig. 3). As the down along the cur. other places, a steeper slope pres- slope they fan out, inhibiting extensive growth of ent, dense Montastrea annularis reefs sometimes sessile organisms. Among the corals that penetrate develop (Roos, 1971) (fig. 4). Down the slope as sedimentation coral decreases deepest (80 m) are Montastrea cavernosa and Aga- increases, growth ricia undata. The crustose coralline algae are still until the sandy flat is reached again at a depth

Remarkable abundant on the second drop-off at 70 to 80 m. of 20 to 30 m. and inexplicable is the

absence of Agaricia species at the deeper reef in Horizontal distribution certain areas. Towards the exposed S.E. point of

The difference in the morphology of die coast Aruba coral growth is extensive. Here the bottom

varied zones of together with exposure to water movement, fauna, characteristic of the shallower

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reefs, occurs In the with the observations of that A. deeper. response to strong Glynn (1973) pal-

water mata the surf beaten coast of the movement, Millepora species, Agaricia aga- grows deeper on

ricites and gorgonians are very common at a depth San Bias islands. The sparcity of corals on the

of 10 all colonies oriented in be attributed to the force of the water m, being uniformly terrace may

a plane parallel to the wave front. Millepora spe- movements and the relative absence of herbivores

cies form large ridges extending seawards. Other (Van den Hoek, 1969; Wanders & Wanders-Faber,

coral colonies are encrusting or flattened. in press). The explored slopes over the drop-off

The N.E. coasts of the islands are, with the ex- carry a dense coral cover, the species composition

of few the leeward reefs. the ception a days a year, subjected to strong resembling However, growth

trade winds. Normal velocity is 7.2 m/sec. and form of the colonies differs from that on the S.W.

is corals flattened, persistency higher than 96%. Reef formation on coast. The are encrusting or very

these coasts varies with morphology of the bottom. an exception being the columnar form of Mon-

The morphology generally follows the same pat- tastrea annularis (fig. 6). Gorgonacea are very

tern as the S.W. coasts. The most conspicuous abundant. The slope ends at a sandy terrace at

difference between leeward and windward coasts 35-45 m; at about 60 m there is another drop-off.

is the presence, in the latter, of a submarine plat- Apart from this deep terrace, nearly no sediments

form densely covered with the alga Sargassum are present.

Van den platycarpum (cf. Hoek, 1969) in front of Important exceptions to this general pattern are the steep shore oliff. On the platform, sloping found. Near the S.E. point of Curaçao the reef is

from 2-6 to m a drop-off at 10-15 m, coral growth very much like the exposed S.E. point of Aruba,

is sparee. It consists mainly of encrusting colonies though it is larger and extends much deeper. The

of Diploria clivosa and Porites astreoides. Acro- Sargassum community is not found here and the

in of of coral pora palmata occurs places, but deeper (5-15 shallow terrace consists a system

m) than on the leeward coast This to the shore (fig. 5). agrees grooves and spurs perpendicular (fig.

at Fig. 4. Montastrea annularis reef a depth of 10 m in front of Mangel Altu (Aruba).

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7). There can be little doubt that the influence of The relative importance of environmental factors

eddies and water mixing at the tip of the islands

will favor the formation of a coral reef. A few A curious example of how environmental fac-

the distribution is km farther Northwest along coast, the Sar- tors govern the of corals shown by

bottom ex- coral environmental gassum community covers the again, comparison of population and

of the tending even over the drop-off down to the sandy factors deeper reef slope with a large shal-

plain at 35 m. Corals are only found flattened low inland bay, Spaanse Water in Curaçao. The

corallines between the algae and crustose are very similarity was first noted by Roos (1964, 1971). abundant. Although the bottom is covered with Table I shows the coral community of the bay

Sargassum platycarpum, close inspection of the to be comprised of three components: a shallow- bare substrate shows the algae to be separated by water one, characterized by Favia fragum and Si-

with of and rock, numerous signs the scraping derastrea radians; a deep-water one with Scolymia

biting of herbivores. Still farther to the North- lacera, S. cubensis and Helioseris cucullata; and a

west, the appearance of the bottom (terrace and third group of corals which have a very broad

covered with slope being algae) abruptly changes ecologioal range, e.g. Agaricia agaricites, Porites

On the corals the and Montastrea again. slope, suddenly replace astreoides, Diploria strigosa ca-

algae resulting in a flourishing coral reef. Since vernosa. This corail community lives on a mud

the environmental circumstances are apparently bottom and on mangrove roots at a depth of 1

it is hard to to The is is not changing, give an explanation. 2 m. water very quiet. Turbidity very

Goreau (1969) attributed such variability to ran- high and mainly caused by inorganic matter (De

dom effects connected with the development of Kock & De Wilde, 1964). The light intensity at

reefs youthful communities. However, as these are a depth of 2 m is less than 30% of the surface

inaccessible, detailed studies value. The of the shallow-water very preventing so presence hardy

far, these observations are only preliminary. species is not surprising (Goreau, 1959). The

coast of Fig. 5. N.E. Curaçao at a depth of 12 m. The terrace with the Sargassum community is at the drop-off bordered and dead The dead with by living Acropora palmata colonies. parts are overgrown crustose coralline algae. Herbivores apparently prefer grazing on the crustose corallines to feeding on the Sargassum.

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Table I. List of shallow-water Scleractinia of the Netherlands Antilles. Classification mainly after Wells & Lang

(1973).

Class ANTHOZOA

Order SCLERACTINIA

Suborder ASTROCOENIINA L S W 34. M. cavernosa(Linnaeus, 1766)

L S W 35. Solenastrea bournoni Milne Edwards &

Family ASTROCOENIIDAE Haime, 1850

L S W 1. Stephanocoenia michelinii Milne Edwards

& Haime, 1848 Family RHIZANGIIDAE

L W 36. Astrangia solitaria (Lesueur, 1817)3)

Family POCILLOPORIDAE L 37. Phyllangia americana Milne Edwards &

3 L S W 2. Madracis decactis (Lyman, 1859) Haime, 1850 )

L 3. M. formosa Wells, 1973 L 38. Colangia immersa De Pourtalès, 1871 3)

L W 4. M. mirabilis (Duchassaing & Michelotti,

1861) Family OCULINIDAE

L 5. M. pharensis (Heller, 1868) L 39. Oculina diffusa Lamarck, 1816

L 6. M. myriaster (Milne Edwards & Haime,

1849)2) Family MEANDRINIDAE

L 7. M. senaria Wells, 1973 L W 40. Meandrina meandrites (Linnaeus, 1758)

L W 41. Dichocoenia stokesii Milne Edwards &

Family ACROPORIDAE Haime, 1848

L W 8. Acropora palmata (Lamarck, 1816) L 42. D. stellaris Milne Edwards & Haime, 1849

L S W 9. A. cervicornis (Lamarck, 1816) L W 43. Dendrogyra cylindrus Ehrenberg, 1834 L 10. A. prolifera (Lamarck, 1816) Family MUSSIDAE

L W 44. Mussa angulosa (Pallas, 1766)

Suborder FUNGIINA L S 45. Scolymia lacera (Pallas, 1766)

L S W 46. S. cubensis (Milne Edwards & Haime, 1849)

Family AGARICIIDAE L W 47. Isophyllia sinuosa (Ellis & Solander, 1786)

L S W 11. Agaricia agaricites (Linnaeus, 1758) L W 48. I. multifloraVerrill, 1901

L 12. A. tenuifolia Dana, 1846 L W 49. Isophyllastrea rigida (Dana, 1846)

L 13. A. undata (Ellis & Solander, 1786) L W 50. Mycetophyllia ferox Wells, 1973

L 14. A. lamarcki Milne Edwards & Haime, 1851 L W 51. M. aliciae Wells, 1973

L 15. A. grahamae Wells, 1973 L 52. M. reesi Wells, 1973

L S 16. A. fragilis Dana, 1846

L S W 17. Helioseris cucullata (Ellis & Solander, 1786)

Suborder CARYOPHYLLIINA

Family SIDERASTREIDAE

L S W 18. Siderastrea siderea (Ellis & Solander, 1786) Family CARYOPHYLLIIDAE

L S W 19. S. radians (Pallas, 1766) L 53. Caryophyllia maculata (De Pourtalès, 1874)3)*)

L 54. C. cf. Family PORITIDAE sp. i C. antillarum De Pourtalès,

3 4 L S W 20. Por ites ast reoides Lamarck, 1816 1874 ) )

L 21. P. branneri Rathbun, 1879 L 55. Desmophyllum riisei Duchassaing & Mi-

3 4 L W 22. P. porites (Pallas, 1766) chelotti, 1860 ) )

L 23. P. divaricata Lesueur, 1820 L S W 56. Eusmilia fastigiata (Pallas, 1766)

L S W 24. P. furcata Lamarck, 1816

Suborder DENDROPHYLLIINA

Suborder FAVIINA

Family DENDROPHYLLIIDAE

Family FAVIIDAE L W 57. Tubastrea coccinea Lesson, 1831 3) L S W 25. Favia fragum (Esper, 1795)

L W 26. Diploria clivosa (Ellis & Solander, 1786) L S W 27. D. labyrinthiformis (Linnaeus, 1758)

L S W 28. D. strigosa (Dana, 1846)

L S W 29. Manicina areolata (Linnaeus, 1758)

L S W 30. Colpophyllia natans (Müller, 1775) L Leeward Group

L 31. C. breviserialis Milne Edwards & Haime, S Spaanse Water

1849 W Windward Group

2 L W 32. Cladocora arbuscula (Lesueur, 1820) ) Reported by Roos (1971)

L S W 33. Montastrea annularis (Ellis & Solander, 3 ) Ahermatypic species

4 1786) ) Reported by Lang (1970)

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6. The of N.E. reef in 20 The bottom is with corals and Fig. slope a coast Curaçao at m. densely populated gorgo-

nians. There is nearly no sediment present. third of group corals can apparently cope with a be surrounded mostly by sandy shallow flats with wide of conditions. The scattered variety group deep-water or no coral growth.

this then it There are the N.E. species clearly requires light intensity, some exceptions, e.g. point with shallow-water can compete successfully spe- of Saba (Cove Bay). Seawards of a rock zone, a

Clean themselves of sedi- oies. They can effectively sloping sandy terrace was populated at 30 m by

the shows fields of These ments. Their occurrence in bay that Acropora cervicornis. peter out at they are able to stand a more varied temperature 35 m leaving the deeper bottom devoid of corals. and salinity regime than that of their normal deep The N.W. point of St. Eustatius (Boobies Point) water environment. has a steep underwater slope, strewn with huge

A of coral is in sand bottom 30 group species that conspicuously rocks, ending abruptly a at m. absent from the Spaanse Waiter consists of those On the slope there is scattered coral growth and

the reefs and reef of the colonies species which flourish on zones most are encrusting. This is one of

and other the habitats exposed to waves strong water move- most exposed to waves and currents ments. Typically, Acropora palmata and Dendro- with good sediment drainage. gyra cylindrus are missing. This example shows Though large areas, especially the exposed that of have been one should not think coral species as being coasts, not explored, the Windward adapted to a special place on reefs but indeed to Group does not seem to have reef development

of a set environmental factors. comparable with the reefs of the Leeward Group. Major factors restricting coral reef development

the character are flat, sandy of extensive parts of II. THE REEFS OF THE WINDWARD GROUP the seafloor and the absence of steep slopes (Go- & reau Wells, 1967). Also, the Windward Group

The stony corals and their distribution in depth is situated in the Caribbean hurricane belt. Studies less than 10 m on St. Martin, St. Eustatius and on the mechanical effect of hurricanes in the Ca-

Saba have been studied by Roos (1971). Scuba ribbean have shown their destructive force to be of diving to depths 35 m, I found the islands to catastrophic to coral reefs (Stoddard, 1963; Glynn

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Fig. 7. Near the base of the cliff (depth 5 m) on the N.E. tip of Curaçao, the abundance of Gorgonia ventalina,

form of demonstrate the Millepora species and the plate-like growth Acropora palmata very rough water con- ditions of the Towards the of and area. drop-off a system grooves spurs develops.

effects to be discussed the et al., 1964). These seem, however, are generally regulating occurrence restricted to rather shallow water. Stoddard (1963) of corals and coral reefs as well as their morphol-

much observed the coral survival to be better even ogy. Also the composition of reef communities at depths as shallow as 7-13 m. He also noted that is strongly dependent on these variables. The fac-

coral. such Acropora cervicornis was the least resistant tors responsible for the exceptions, as the

This is confirmed al. who of reef N.W. of St. Eusta- by Glynn et (1964) re- lack growth at the point port A. cervicornis reefs to be completely destroy- sius, the alternation of coral reef and Sargassum

fields ed. I found healthy A. cervicornis at 30 m community and the absence of Agaricia species on

der Land reefs around remain be around Saba. J. van (pers. comm.) men- some Aruba, to investigat- tioned A. cervicornis at depths of less than 15 m ed. around St. Eustatius and deeper than 17 m at the

Saba Bank. This shows that hurricane effects do not prevent growth of the most sensitive coral at ACKNOWLEDGEMENTS these depths.

friends Caribbean Marine I thank my at the Biological

Institute who shared their diving experience. Thanks are CONCLUSION also due to the authorities of the Royal Navy in the

Netherlands Antilles (in particular to W. B. Donker

It is concluded that the environmental factors Curtius) for their cooperation.

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