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<I>Sinularia Flexibilis</I> BULLETIN OF MARINE SCIENCE. 56(1): 303-311. 1995 CORAL REEF PAPER PRELIMINARY EVIDENCE FOR DIRECTIONAL ALLELOPATHIC EFFECTS OF THE SOFT CORAL SINULARIA FLEXIBILIS (ALCYONACEA: OCTOCORALLIA) ON SCLERACTINIAN CORAL RECRUITMENT Mauro Maida, Paul W. Sammarco and John C. Coil ABSTRACT Certain species of alcyonacean soft corals have been shown to release toxins which inhibit growth and cause tissue necrosis in selected adult scleractinian corals in their vicinity. We have now demonstrated experimentally that soft corals with such allelopathic capabilities can affect recruitment of juvenile scleractinian corals. A large colony of Sinularia flexibilis (Oc- tocorallia, Alcyonacea) was placed in the center of each of two steel grids implanted on a shallow reef slope on Orpheus Island, Great Barrier Reef, Australia. Settlement plates were attached to the grids radiating outward from each soft coral in a uniform pattern. Two other grids with settlement plates arranged similarly, but without soft coral colonies, comprised the controls. Scleractinian coral recruitment was assessed against presence or absence of S. flex- ibilis. and as a function of recruitment position with respect to the soft coral colony. Prevailing current direction was measured using a recently developed small current meter, permitting current effects on recruitment to be evaluated. The presence of S. flexibilis was found to significantly depress coral recruitment. Settlement plates positioned down-current from the soft corals exhibited the lowest densities of coral recruitment, and there was a significant negative correlation between recruitment density in each directional quadrant and the pro- portion of time during which current flowed towards that quadrant. There was no correlation between density of juvenile corals and distance from the soft coral, probably due to directional variance. Coral spat included Acropora sp., Pocillopora sp. Seriatopora. Merulina sp., and a Mussidae, but there was no significant difference between the controls and treatments in relation to the taxa found on each grid. The overall mortality rates of coral spat were not significantly different between treatments and controls. These results show that the soft coral S. flexibilis. known to release allelopathic toxins, can depress recruitment of scleractinian corals, and that this depression is exercised in a directional manner, depending upon the prevailing currents. Many plants and animals in terrestrial, freshwater, and marine environments produce secondary metabolites which can be used as chemical signals between interacting species. When released into the environment, these secondary metab- olites can exert either inhibitory or stimulatory influences on other organisms sharing the same habitat. The inhibitory influences are termed allelopathy (Rice, 1984; Gauthier and Aubert, 1981). Certain species of alcyonaceans are known to release into the environment terpenoid toxins (ColI et aI., 1982a), which can act as allelopathic agents in com- petitive interactions with nearby scleractinian corals (Sammarco et aI., 1983). ColI and Sammarco (1983) demonstrated that certain secondary metabolates cause tis- sue necrosis and mortality in scleractinian corals in the laboratory (Sammarco et aI., 1983). To date, all the studies related to the interactions between alcyonacean and scleractinian corals have focused on those between adult colonies (Sammarco and ColI, 1988). It is not known whether soft coral derived toxins can also affect the recruitment of juvenile scleractinian corals in the vicinity of an allelopathic soft coral. A variety of aspects of the settlement of scleractinian corals have been studied on the Great Barrier Reef. These include spawning processes (Harrison et aI., 303 304 BULLETIN OF MARINE SCIENCE, VOL. 56, NO. I, 1995 ,, ,, ,, ,, ~rus Island ,, ,, ~ . '. ,.. .,, 0\\ P;"."t •N ,, ,, ; ,, }'" ,, , Phant~'i\sland ,, "<:l ,, ,, o S. Km Figure 1. Map of the study site, indicating location of the Palm Island group off the central coast of Queensland, Australia; the location of Orpheus Island within that group, and the study site on Orpheus Island. 1984; Babcock et a1., 1986), larval dispersal and recruitment (Sammarco and Andrews, 1988; Andrews et a1., 1988), spatial and temporal variability in settle- ment (Sammarco, 1983; Wallace, 1985; Harriot, 1985; Babcock, 1988; Sammarco, 1991), post-settlement mortality and juvenile growth (Babcock, 1988; Fitzhard- inge, 1988), and aspects of microhabitat preferences for settlement by coral larvae (Harriot and Fisk, 1987; Carleton and Sammarco, 1987). Only one study has considered the possibility of chemical mediation in the processes of coral recruit- ment (Morse and Morse, 1993), and that in the northern hemisphere. In this study, we report results which indicate that some soft corals can depress local recruitment of sc1eractinian coral spat in their vicinity. Further, we provide evidence that the recruitment of sc1eractinian corals around a toxic soft coral is dependent upon the direction of prevailing micro-currents. MATERIALS AND METHODS The experiment was performed on the southern reef slope of Pioneer Bay, Orpheus Island, Queens- land (18°40'S, 145°30'E) (Fig. 1). The soft coral studied was Sinularia jiexibilis Quoy and Gaimard, an abundant (Dinesen, 1(83) and toxic (Coli et aI., 1982b) alcyonacean octocoral, known for its allelopathic capabilities (Sammarco et aI., 1983). The experimental design followed that of a three-way analysis of variance (Sokal and Rohlf, 1981). The three fixed factors were: (1) Presence and absence of S. jiexibilis, (2) Directional quadrant in which the settlement units were located around the soft corals, and (3) Distance of the settlement units from the soft coral. Four galvanised steel grids (5 X 5 cm mesh) measuring 180 by 180 cm were used to attach the settlement units to the red. The horizontal grids were attached to metal fencing posts about 40 cm above the sloping bottom at a mean depth of 4 m. A hole (60 X 60 cm) was cut in the middle of MAIDA ET AL.: ALLELOPATHY AND CORAL RECRUITMENT 305 N ~ A '\. "- / / "- "- ~$ B I c '\. "- / "- / ~ "- .L Figure 2. Schematic representation of the experimental design showing: Details of the settlement arrays; A) PVC sheets divided into six settlement units; B) steel grid; C) Whole experiment: two grids with presence of Sinularia jlexibilis (treatments), two grids with absence of S. jlexibilis (controls). each grid. A large single colony of S. jlexibilis was placed in the hole of two of the grids, and secured to expose its polypary (polyp bearing portion of the colony) above the mesh. The other two grids lacked soft coral and served as controls. One or other control grid holes was used for the placement of the current direction meter, which caused no effect on settlement. Settlement substrata consisted of plastic (PVC) sheeting (15 em x 60 em) divided into six equally sized (15 em x 10 em) settlement units. Eight PVC sheets were secured to each grid so that they radiated out from the centre oriented along the magnetic bearings Nand NE (N), E and SE (E), S and SW (S), Wand NW (W); data were grouped into four directional quadrants (N, E, S, W). This afforded a total of 48 settlement units per grid, i.e., six settlement units per PVC sheet, two PVC sheets per quadrant, and four quadrants per grid. The settlement sheets were attached to the grids starting at a distance of 5 cm from the margin of the soft coral or the grid hole (in the case of controls), thus creating sample distances from 5 to 65 cm from the soft coral for each directional quadrant (Fig. 2). The experiment was initiated in October 1989, 2 weeks prior to the coral spawning season (Harrison et aI., 1984; Babcock et aI., 1986), and allowed to run for 9 months. The plastic settlement sheets were then retrieved, and placed in laboratory seawater tanks for in vivo analysis of coral settlement. Loose sediment was removed from the settlement substrata using gentle agitation of the surrounding water. Larger pieces of branching and filamentous algae were removed with fine dissecting forceps to facilitate observations. The settlement substrata were processed visually with the assistance of a dis- secting microscope, using a magnification range of 60-31Ox. Juvenile corals were assessed for survivorship; mortality was evidenced by the absence of tissue, infestation by endolithic green algae, and overgrowth by associated epibenthic organisms. For iden- tification purposes, coral spat were detached from the PVC surface, bleached, and mounted on glass slides. Taxonomy of the coral recruits was based on electron micrograph collections prepared by Sammarco, Carleton, and Mackley at the Australian Institute of Marine Science. Identification was usually possible to genus level, although in several instances, only to the family level. Current direction data were collected during the course of the experiment (23 March through 13 April 1990), using a small current direction meter specifically developed for this experiment. This meter provided data on the number of hours during which local micro-currents flowed in each of the eight directions sampled, and hence for the four sampling quadrants. A detailed description of the meter has been presented elsewhere (Maida et aI., 1993). Settlement density data were analysed using a three-way analysis of variance. Post-hoc test (Thkey's test) was used following routines described by Sokal and Rohlf (1981). Diversity indices were cal- culated using the Shannon-Wiener Index, and comparisons
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