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Caribbean Staghorn Coral Populations: Pre-Hurricane Allen

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Caribbean Staghorn Coral Populations: Pre-Hurricane Allen III 'LLETIN OF MARINE SCIENCE. 33( I): 132-151. 1983 CORAL REEF PAPER CARIBBEAN STAG HORN CORAL POPULATIONS: PRE-HURRICANE ALLEN CONDITIONS IN DISCOVERY BAY, JAMAICA Verena TunniclifJe ABSTRACT In August 1980, Hurricane Allen caused damage of catastrophic proportions on the coral reefs of Discovery Bay, Jamaica. The extensive staghom coral (Acropora cervicornis) beds were flattened and rapid recovery is not evident. This study describes the staghom populations as they were before the hurricane; work on the present state of the reefs is in progress. Three areas of Discovery Bay were studied. A. cervicornis was most abundant on the haystacks of the west fore reef between 5 and 20 m depth where up to II corals per m2 produced a maximum of 10 m of branches. Here the individual corals were significantly larger than those of the east fore reef or the back reef. Although a significant negative cor- relation of growth rate with depth (cm 'yr 1 = 14.46 - 0.19 [depth, m]) was found on the west fore reef, the shallow corals were not taller but rather showed a higher level of branching; there is an indication that shallow corals branched more frequently. Contacts between adjacent corals were common and showed varying grades of tissue acceptance. The num ber of between- coral grafts did not correlate significantly with population density but did with water depth. An examination of branch orientation across the fore reef revealed a shift coincident with wave refraction but scatter was very high; frequent breakage disrupts the preferred growth directions. Most of the staghoms at all sites had been broken at some time but the predominant method of reattachment differed at each site. The numbers of broken branches in corals on the fore reef was a function of coral size; in the back reef they were directly proportional to depth which suggests a lagoonward transport of the corals. An average of one-third of each coral is bare of tissue. The major predators are a sea urchin, damselfish, a snail and a polychaete. Clionid sponges are abundant in the coral skeleton. There is a large contribution by A. cervicornis to the skeletal calcium carbonate of the reef, and thus to vertical reef accumulation; this study estimates 1.4 kg·m-2·yr-1 in the cervicornis zone. There appeared to be a slight down-reef shift of material before it was cemented in place. A. cervicornis dominated the west fore reef community; abundances of massive corals fell where those of the staghom rose. The success of A. cervicornis derived primarily from its ability to regenerate from fragments. It exerts a competitive dominance over other space-occupiers by virtue of its high growth rate, a pronounced shading effect, mechanical damage to underlying corals through its break- age, and aseasonal recruitment to available spaces by fragmentation. There does appear to be a limit on its tolerance to disruption; present questions center around the fate of these Jamaican reefs. Many reefs of the Caribbean are dominated by monotypic stands of the staghorn coral Acropora cervicornis (Lamarck). Dense "bramble-patches" of this branching stony coral can be found in the protected waters of the Caribbean: Jamaica (Go- reau, 1959), Florida (Shinn, 1976), Bahamas (Bottjer, 1980), Puerto Rico (Almy and Carrion-Torres, 1963) and Cuba (Kuhlman, 1974) among others, and it is present as scattered stands in more exposed areas such as those of the Dutch Antilles (Roos, 1971), St. Croix (Rogers, 1979) and Barbados (Lewis, 1960). As one of three species of Acropora in the Caribbean, it occupies the most extensive depth ranges from 0 to 30 m. In August 1980, Hurricane Allen passed very close to the reefs of the northern shore of Jamaica. The resulting damage to the reef was catastrophic in proportions 132 TUNNICLIFFE: CARIBBEAN STAG HORN CORALS 133 Figure I. Map of Discovery Bay, Jamaica. The three study areas are marked with dashed lines. (Woodley, 1980; Porter et aI., 1981; Woodley et aI., 1981) and the fields of A. cervicornis were flattened; mortality of this coral during and after the hurricane was over 95% (Knowlton et aI., 1981). An extensive research effort to determine the patterns of recovery of these reefs is presently centered at the Discovery Bay Marine Laboratory in Jamaica. The study reported here was conducted during the 3 years prior to the hurricane. It represents an intensive investigation of A. cervicornis beds that no longer exist. These results may be useful, however, (1) as an example of the dynamics of similar assemblages in other areas, and (2) to compare with the recovery sequence now taking place on the Discovery Bay reefs. Much of this study was conducted with the view to understanding the effects of waves on both coral and community biology. Mechanical and hydrodynamic features of A. cervicornis are described in TunnicIiffe (1979) and (1982) while the effects of wave breakage on reproduction are outlined in Gilmore and Hall (1976) and Tunnicliffe (1981). Study Area Discovery Bay is located on the north shore of Jamaica in the Caribbean (Fig. I). The mouth of the bay is blocked by well-developed reefs but an entrance channel was dredged to a depth of 13 m. The reef structure and communities of this area have been extensively described by Goreau (1959), Goreau and Goreau (1973), Kinzie (1973) and Goreau and Land (1974) among others. The main study areas for this work are indicated on Figure I: Pinnacle Two on the west fore reef, Damsel Gardens in the east back reef and discrete sites on the east fore reef. Supplementary information was added from adjacent areas. Pinnacle Two is a large "haystack" (Kinzie, 1973) extending unbroken from the cresting reef top to 30 m in depth and is about 300 m long and 100 m wide. It is bounded by two deep sand channels on either side and a sandy moat at 30 m separates this haystack from a large seaward mound ("the pinnacle") that rises to 23 m and then drops away steeply to the deep fore reef (Fig. 2). The biological zonation follows that described by Goreau and Goreau (1973): reef crest zone dominated by Acropora palma/a, a mixed zone of many coral species, the cervicornis zone and the deep fore reef where plating corals are common. The coral population lagoonward of the reef crest is sparse and dominated by mound corals. A poorly sorted sediment covers the bottom and frequent resuspension of this sediment results in high turbidity (Dodge et aI., 1974). The bottom slopes down into a deep lagoonal basin 150 m behind the reef crest in which no corals grow. The east fore reef of Discovery Bay is an extensive plateau of low slope. Topographically, the area 134 IlULLETIN OF MARINE SCIENCE. VOL. 3J. NO. I. 19~3 Figure 2. Sketch of Pinnacle Two, west fore reef. This haystack supported prolific A. cervicornis growth on the terrace. It is about 350 m from reef crest to the top of the pinnacle where water depth is 23 m. A. reef crest-palmata zone 2 m; B, mixed zone 7 m; C, cen'icornis zone, 15 111; D, sand channel and moat, 30 m and E, pinnacle, 25 m. has few of the distinctive haystack-sand channel structures of the west side. Organism zonation is indistinct; sponges, gorgon ian corals and mound corals predominate and acroporid corals are less common (Liddell and Ohlhorst, 1981). METHODS On each of the study areas, a line transect was established for study of individual A. cervicornis corals. On Pinnacle Two, this transect extended from the reef crest to 30 m in depth for a total of 280 m downslope. A compass was used to maintain a heading of N30oE, the direction in which the haystack extends from the reef crest. The transect in the back reef traversed 120 m perpendicular to the reef crest from 3 to 13 m in depth, below which the sand bottom was unpopulated. Because of the large distance on the east fore reef from the reef crest to 30 m in depth, 50-m transects with N300E orientations were studied at depths of6 m, 9 m, 12.5 m, 15.3 m, 21.5 m and 26 m. I estimated the population densities of A. cervicornis in two ways: the number of corals per square meter and the total length of live branches of A. cervicornis. Every 10 m from the reef crest, a 4-m' quadrat divided into I-m square areas was placed on the reef; measurements in each I-m' partition were taken with a tape measure and the four measures were averaged. Because of the tendency of A. cervicornis to intergrow and fragment, it is occasionally difficult to identify a single coral. Nearly all structures of this colonial animal could be recognized by a single basal stalk from which all subsequent branches formed. The A. cervicornis that was under the mark at each meter along the transects was examined for a number of variables. For the corals along the Pinnacle Two (n = 187) and back reef(n = 56) transects, these variables were: maximum height above substratum, maximum branch spread (width), highest branching order, basal circumference, estimated amount dead, the number of grafts formed (within the coral and to others), numbers of tip, branch and basal fractures, type of basal attachment and, lastly, the presence and type of predation. The east fore reef corals (n = 128) were examined only for height, width, amount dead, total number offractures, basal attachment and predation. The variable "branching order" was determined by the method of Strahler (1957). The end branches are of order TUNNICLIFFE: CARIBBEAN STAGHORN CORALS 135 A.
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