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FAU Institutional Repository FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: ©1986 Elsevier B.V. The final published version of this manuscript is available at http://www.sciencedirect.com/science/journal/00220981 and may be cited as: Kilar, J., & McLachlan, J. (1986). Ecological studies of the alga Acanthophora specifera (Vahl) Børg. (Ceramiales, Rhodophyta): vegetative fragmentation. Journal of Experimental Marine Biology and Ecology, 104(1-3), 1-21. doi:10.1016/0022-0981(86)90094-8 J. Exp. Mar. Bioi. £Col., 1986, Vol. 104, pp. 1-21 Elsevier JEM 00786 Ecological studies of the alga, Acanthophora spicifera (Vahl) Berg, (Ceramiales: Rhodophyta): vegetative fragmentation* .. I 1 i John A. Kilar ,** and J. Mcl.achlan? IThe Smithsonian Tropical Research Institution. P.O. Box 2072. Balboa. Panama. and the Department of \. Biology. Dalhousie University. Halifax . Nova Scotia B3H 4J1. Canada ; 2National Research Council of Canada . Halifax. Nova Scotia B3H 321. Canada (Received 4 June 1986; revision received II August 1986; accepted 22 August 1986) Abstract: The propagation of Acanthophora spicifera (Vahl) Borg. was studied on a fringing-reef platform at Galet a Point, Caribbean Panama. Manipulative experiments and detailed descriptive data indicated that fragmentation accounted for this alga's standing crop and distribution. Fragments were broken off by turbulen ce in the fore reef, transported by currents across a seagrass meadow, snagged, and attached or entangled in the back reef. Accumulations of ~66 g (dry wt.) · m - 2 occurred. A . spicifera was a major contributor to drift biomass, and, depending on prevailing current velocities, fragments entering the back reef had a 49 to 93 % chance of recruiting. Higher current velocities ( ~0 .24 m . s - I) decreased the ability of free-floating fragment s to snag by decreasing the frequency of fragment-substratum contacts. 25% of the snagged fragments remained > 3 days, and fragments required < 2 days to attach to Laurencia papil/osa (Forsk.) Grev. or to another frond of Acanthophora spicifera. Fragments were unable to recruit into plots of Thalassia testudinum Bank ex Konig and Sims. Tetrasporic plant s were common, comprising as much as 96% of fore-reef and 80% of the back-reef populat ions. In October , 3% of the fore-reef population had cystocarps; otherwise, no other gametophytes were found. The percentage of spore-bearing tetrasporophytes was significantly greater in the Laur encia zone than in the fragment-derived Acanthophora zone. Vegetative fragmentati on was demonstrated as an effective means of propagati on, while the ecological significance of tetraspores was unclear. Key Words: Acanthophora spicifera; Drift algae; Fragmentation; Asexual propagation; Laurencia papillosa INTRODUCTION Acanthophora spicifera is a rhodophycean alga widely distributed in subtropical and tropical seas (Doty, 1961; Taylor, 1967) and occurs in many intertidal (Tabb & Manning, 1961; Rao & Steeramalu, 1974) and subtidal habitats (Earle, 1972a). l t , Extensive stands are reported from shallow reef flats (Doty, 1969; Meyer & Birkeland, 1974; Conner & Adey, 1977; Santelices, 1977), and at depths to 17 m in Puerto Rico (Dahl, 1973)and 22 m in the Virgin Islands (Mathieson et al., 1975). It occurs on hard substrata (Taylor & Bernatowicz, 1969; Almodovar & Pagan, 1971), as an epiphyte on other algae (Mathieson et al., 1975; Russell, 1981), or as stable, free-living population * Contribution No. 527 from the Harbor Branch Oceano graph ic Institution, Fort Pierce, Florida. ** Present address:Harbor Branch Oceanographic Institution Inc., 5600 Old Dixie Highway, Fort Pierce, FL 33450 U.S.A. 0022-0981/86/$03.50 © 1986 Elsevier Science Publishers B.V.(Biomedical Division) 2 JOHN A. KILAR AND J. McLACHLAN (Russell, 1974; 1981; Cowper, 1978; Benz etal., 1979). In Panama, Hay (l981b) and Kilar & Lou (1984) report A. spicijera in the diets of herbivorous fishes, sea urchins, and crabs, and Hay (1981a) has shown that fish can exclude the species from some habitats. Acanthophora spicifera (Vahl) Berg. is attached to hard substratum by a large, irregularly lobed disc, from which many erect fronds arise that are sparingly branched to bushy (Fig. 1). Main branches are beset with short, determinate branchlets that are markedly spinose and arranged spirally with a 1/4 divergency. A triphasic, alternation of generations has been reported for A. spicijera; tetrasporophytic and gametophytic generations are isomorphic and the gametophyte dioecious (Bergesen, 1918; Taylor, 1967). a . I b. '/ Fig. I. Branching morphology of Acanthophora spicifera: (a) general anatomy, vertical bar = I mm ; (b) determinate branches, horizontal bar = 25 mm ;(c) determinate branch with reatt achment bran ches, horizont al bar = 25 mm. The introduction of A . spicifera to Hawaii was reported by Doty (1961) who suggested that the alga arrived attached to a hull of a fuel-oil barge towed from Guam. Subsequent studies by Mshigeni (l978) and Russell (1981) have yielded conflicting results as how the alga rapidly spreads. Mshigeni (l978) concluded that A. spicifera was propagated THE VEGETATIVE FRAGMENTATION OF ACANTHOPHORA SPICIFERA 3 by vegetative fragmentation, whereas Russell suggested that tetraspores, produced year-round, were the principal agent of dispersal. The objective of our stud~ was to evaluate the roles of vegetative fragments and spores of A . spicifera as means of propagation. At Galeta Point, Panama, Birkeland et al. (1973) and Kilar et al. (in press) report A . spicifera in the fore reef or Laurencia zone, and in the back reef or Acanthophora zone (see maps within these papers). Our working hypotheses are that: (i) the fore-reef A. spicifera gives rise to fragments that colonize the back reef and (ii) spores account for the reef-flat distribution of A. spicifera. By fragment, we mean an unspecialized, detached piece of thallus. METHODS STUDY SITE Research was done on the reef flat adjacent to the Galeta Marine Laboratory of the Smithsonian Tropical Research Institute (S.T.R.I., located at Galeta Point, Caribbean Panama (9°24.3'N : 79°51.8'W; Fig. 2). On the reef flat luxuriant growths of algae occur, mainly Halimeda opuntia (L.) Lamour., Laurencia papillosa (Forsk.) Grev., and Acanthophoraspicifera,and the seagrass, Thalassia testudinum Bank ex Konig and Sims. STUDY SITE Fig. 2. Study site at Galeta Point, Panama. The seaward side of the reef extends down to ~ 13-m depth, while on the landward side the reef is bordered by a lagoon or mangrove swamp. The reef flat is usually covered by 0.1 to 0.4 m of water, with tidal fluctuations of 0.7 m (Macintyre & Glynn, 1976). The topography of the reef flat is such that the highest elevation occurs in the fore reef 4 JOHN A. KILAR AND J. McLACHLAN and decreases landward, resulting in an almost unidirectional flow of water over the reef surface. Sea-water temperature on the reef flat is generally between 26 and 29 0 C, and the salinity between 32 and 35%0 (Hendler, 1977). The reefis typical of the fringingreefs of the Caribbean coast of Panama (Glynn, 1972; Macintyre & Glynn, 1976). During Panama's dry season, mid-November to March, northern and northeasterly trade winds blow at a mean velocity of 24 to 27 km h - 1 (Hendler, 1976), ~ 3 times the mean velocityoccurring in the wet season, April to mid-November. Waves generated by these strong winds cause considerable turbidity and remove weakly attached organisms on the reef platform. During the wet-season, when calm seas coincide with a low pressure system or with low spring tides, the reef surface may be exposed to air (Hendler, 1977). Exposure periods of several days to weeks kill most non-swimming herbivores and greatly reduce algal cover (Hendler, 1976; 1977; Hay, 1981a; Kilar, 1984; Kilar & Lou, 1984). More detailed information about the geology, species composition, and hydrodynamics at Galeta Point is available in Earle (1972b), Birkeland et al. (1973), Macintyre & Glynn (1976), Cubit & Williams (1983) and Kilar et al. (in press). STANDING CROP The standing crop of Acanthophora spicifera was obtained using the methods previ­ ously reported by Kilar et at. (in press). About 100 samples in the Acanthophora zone and 50 in the Laurencia zone were taken each month from February 1979 to March 1980. FROND SURVIVORSHIP Frond survivorship was determined at three stations: exposed (exposed station) and sheltered (sheltered station) Laurencia zone and the Acanthophora zone (back-reef station). At each station, two plots (0.3 x 0.6 m) were haphazardly selected in stands of A. spicifera, and in each plot 20 fronds were tagged with 4-mm wide x 50-mm long, plastic coated "twist ties" that were secured between two determinate branches. To keep the size of the tag to a minimum, excessive portions (10-20 mm) were removed. Frond losses were monitored twice weekly for a 4-month period from 19 September to 18 December 1981, under both wet- and dry-season conditions (for station locations, see Fig. 3). The percent cover of A. spicijera was measured along with survivorship from four permanent quadrats (0.3 x 0.5-m), located adjacent to each survivorship station. As wave-zone turbulence made sampling difficult, each quadrat was divided into 20, 0.05 x 0.15-m areas that were visuallyestimated for coverage. The mean of 80 estimates of cover defined each station 's coverage. THE VEGETATIVE FRAGMENTATION OF ACANTHOPHORA SPICIFERA 5 SUBTIDAL REEF E ~ Ph. C-2 r.r. .--- c-s LAGOON t------l 30m Fig. 3. Station locations on the reeffl at at Galeta Point , Panama : N-(no.) = drift nets; C-(no.) = recruitm ent plots in Thalassia beds ; survivorship study : E = exposed fore-reef stati on; S = sheltered fore-reef station; BR = back -reef station; Ph = phenology tr ansects; snagging experiment: A = Acanthophora zone station; Tr = Thalassia-rubble station; Th = Thalassia station; attachment study (*). DRI FT SAMPLING To illustrate differences in exported biomass between a fragmenting and non­ fragmenting species, estimates of driftA .
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