Abstract
Overgrowth interactions between encrusting cyclostome (eight species) and cheilostome (over 50 species) colonies were examined in subtidal communities in the northern Adriatic Sea off Rovinj, Croatia, in May–June 1988 and September–October 1990. Cheilostome colonies occupied ca. 80% of the available substrate space, whereas the cyclostome colonies occupied <5%. Out of 210 recorded interactions, cheilostomes overlapped cyclostomes in 164, while cyclostomes overlapped cheilostomes in only 16; 30 encounters resulted in mutual overlap or at least temporary growth termination along the line of contact. Most of the recorded interactions were for the cyclostomes Diplosolen obelia (Johnston) and Plagioecia patina (Lamarck). Both species elevated their colony margins on contact with a cheilostome in some instances, but D. obelia only overgrew competing cheilostomes in 12/120 encounters, and P. patina never prevailed. I propose that the cheilostomes have a key evolutionary innovation, lacking in cyclostomes, consisting of more rapid ontogenetic development of zooids along colony margins so that encrusting cheilostomes have substantially higher colony margins. This results in an outflow of filtered water along the edge of encrusting cheilostome colonies, while cyclostomes take water in at colony margins. These contrasting feeding currents apparently give a competitive advantage to the cheilostomes where colony margins approach and make contact with those of cyclostomes.
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Literature cited
Buss, L. W. (1979). Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. In: Larwood, G. P., Rosen, B. R. (eds.) Biology and systematics of colonial organisms. Academic Press, London, p. 459–497
Buss, L. W. (1980a). Competitive intrasivity and size-frequency distributions of interaction populations. Proc. natn. Acad. Sci. U.S.A 77: 5355–5359
Buss, L. W. (1980b). Bryozoan overgrowth interactions — the interdependence of competition for space and food. Nature, Lond. 281: 475–477
Buss, L. W. (1981). Mechanisms of competition between Onychocella alula (Hastings) and Antropora tincta (Hastings) on an eastern Pacific rocky shoreline. In: Larwood, G. P., Nielsen, C. (eds.) Recent and fossil Bryozoa. Olsen & Olsen, Fredensborg, p. 39–49
Buss, L. W., Jackson, J. B. C. (1979). Competitive networks: non-transitive competitive relationships in cryptic coral reef environments. Am. Nat. 113: 223–234
Carle, K. J., Ruppert, E. E. (1983). Comparative ultrastructure of the bryozoan funiculus: a blood vessel homologue. Z. zool. Syst. Evol. Forsch. 21: 181–193
Chamberlain, J. A. Jr., Graus, R. R. (1975). Water flow and hydromechanical adaptations of branched reef corals. Bull. mar. Sci. 25: 112–125
Cook, P. L. (1977). Colony-wide water currents in living Bryozoa. Cah. Biol. mar. 17:31–47
Day, R. W., Osman, R. W. (1981). Predation by Patiria miniata (Asteroidea) on bryozoans: prey diversity may depend on the mechanism of succession. Oecologia 51: 300–309
Dayton, P. K. (1971). Competition, disturbance, and community organization: the provision and subsequent utilization of space in a rocky intertidal community. Ecol. Monogr. 41: 351–389
Dick, M. H. (1987). A proposed mechanism for chimney formation in encrusting cheilostomes. In: Ross, J. R. P. (ed.) Bryozoa: present and past. Western Washington University Press, Bellingham, p. 73–80
Gordon, D. P. (1972). Biological relationships of an intertidal bryozoan population. J. nat. Hist. 6: 503–514
Harmelin, J.-G. (1976). Le sous-ordre de Tubuliporina (Bryozoaires Cyclostomes) en Méditerranée, écologie et systématique. Mém. Inst. océanogr. Monaco 10: 1–326
Harmelin, J.-G. (1977). Bryozoaires des Iles dltyères: cryptofaune bryozoologique des valves vides de Pinna nobilis rencontrées dans les herbiers des Posidonies. Trav. sci. Parc nation. Port-Cros 3: 143–157
Harvell, C. D. (1984). Predator-induced defense in a marine bryozoan. Science, N. Y. 224: 1357–1359
Jackson, J. B. C. (1977). Competition on marine hard substrata: the adaptive significance of solitary and colonial strategies. Am. Nat. 111: 743–767
Jackson, J. B. C. (1979). Overgrowth competition between encrusting cheilostome ectoprocts in a Jamaican cryptic reef environment. J. Anim. Ecol. 48: 805–823
Jackson, J. B. C. (1985). Distribution and ecology of clonal and aclonal benthic invertebrates. In: Jackson, J. B. C., Buss, L. W., Cook, R. E. (eds.) Yale University Press, New Haven, p. 297–355
Jackson, J. B. C., Hughes, T. P. (1985). Adaptive strategies of coral-reef invertebrates. Am. Scient. 73: 265–274
Jackson, J. B. C., McKinney, F. K. (1990). Ecological processes and progressive macroevolution of marine clonal benthos. In: Ross, R. M., Allmon, W. D. (eds.) Causes of evolution. University of Chicago Press, Chicago, p. 173–209
Jackson, J. B. C., Winston, J. E. (1982). Ecology of cryptic coral reef communities. I. Distribution and abundance of major groups of encrusting organisms. J. exp. mar. Biol. Ecol. 57: 135–147
Keough, M. J. (1984). Dynamics of the epifauna of the bivalve Pinna bicolor: interactions among recruitment, predation, and competition. Ecology 65: 677–688
Lang, J. (1973). Interspecific aggression by scleractinian corals. 2. Why the race is not only to the swift. Bull. mar. Sci. 23: 260–279
Lidgard, S. (1981). Water flow, feeding, and colony form in an encrusting cheilostome. In: Larwood, G. P., Nielsen, C. (eds.) Recent and fossil Bryozoa. Olsen & Olsen, Fredensborg, p. 135–142
Lidgard, S. (1985). Zooid and colony growth in encrusting cheilostome bryozoans. Palaeontology 28: 255–291
Liem, K. F. (1973). Evolutionary strategies and morphological innovations: eichlid pharyngeal jaws. Syst. Zool. 22: 425–441
Liem, K. F. (1990). Key evolutionary innovations, differential diversity, and symecomorphosis. In: Nitecki, M. H. (ed.) Evolutionary innovations. University of Chicago Press, Chicago, p. 147–170
Lutaud, G. (1961). Contribution a l'étude du bourgeonnement et de la croissance des colonies chez Membranipora membranacea (Linne), Bryozoaire chilostome. Annls Soc. r. zool. Belg. 91: 157–300
McKinney, F. K. (1988). Elevation of lophophores by exposed introverts in Bryozoa: a gymnolaemate character recorded in some stenolaemate species. Bull. mar. Sci. 43: 317–322
McKinney, F. K. (1989). Two patterns of colonial water flow in an erect bilaminate bryozoan the cheilostome Schizotheca serratimargo. Cah. Biol. mar. 30: 35–48
McKinney, F. K. (1990). Feeding and associated colonial morphology in marine bryozoans. Rev. aquat. Sci. 2: 255–280
McKinney, F. K. (1991a). Colonial feeding currents of Exidmonea atlantica (Cyclostomata). Bull. Soc. Sci. nat. Ouest France Mém. H. S. 1: 263–270
McKinney, F. K. (1991b). How phylogeny limits function — the example of Exidmonea. Nat. Geogr. Res. Expl. 7: 432–441
McKinney, F. K., Boardman, R. S. (1985). Zooidal biometry of Stenolaemata. In: Nielsen, C., Larwood, G. P. (eds.) Bryozoa: Ordovician to Recent. Olsen & Olsen, Fredensborg, p. 193–203
McKinney, F. K., Jackson, J. B. C. (1989). Bryozoan evolution. Unwin Hyman, Boston
Okamura, B. (1988). The influence of neighbors on the feeding of an epifaunal bryozoan. J. exp. mar. Biol. Ecol. 120: 105–123
Osborne, S. (1984). Bryozoan interactions: observations on stolonal outgrowths. Aust. J. mar. Freshwat. Res. 35: 453–462
Osman, R. S. (1977). The establishment and development of a marine epifaunal community. Ecol. Monogr. 47: 37–63
Palumbi, S. R., Jackson, J. B. C. (1983). Ageing in modular organisms: ecology of zooid senescence in Steginoporella sp. (Bryozoa: Cheilostomata). Biol. Bull. mar. biol. Lab., Woods Hole 164: 267–278
Rubin, J. A. (1985). Mortality and avoidance of competitive over-growth in encrusting Bryozoa. Mar. Ecol. Progr. Ser. 23: 291–299
Russ, G. R. (1982). Overgrowth in a marine epifaunal community: competitive hierarchies and competitive networks. Oecologia 53: 12–19
Ryland, J. S. (1976). Physiology and ecology of marine bryozoans. Adv. mar. Biol. 14: 285–443
Silén, L., Harmelin, J.-G. (1974). Observations on living Diastoporidae (Bryozoa, Cyclostomata), with special regard to polymorphism. Acta zool., Stockh. 55: 81–96
Simpson, G. G. (1959). The nature and origin of supraspecific taxa. Cold Spring Harb. Symp. quant. Biol. 24: 255–271
Stebbing, A. R. D. (1973). Competition for space between the epiphytes of Fucus serratus L. J. mar. biol. Ass. U. K. 53: 247–261
Todd, C. D., Turner, S. J. (1988). Ecology of intertidal and sublittoral cryptic epifaunal assemblages. II. Non-lethal overgrowth of encrusting bryozoans by colonial ascidians. J. exp. mar. Biol. Ecol. 115: 113–126
Vogel, S. (1981). Life in moving fluids. Willard Grant Press, Boston
Walters, L. J., Wethey, D. S. (1986). Surface topography influences competitive hierarchies on marine hard substrata: a field experiment. Biol. Bull. mar. biol. Lab., Woods Hole 170: 441–449
Ward, M. A., Thorpe, J. P. (1991). Distribution of encrusting bryozoans and other epifauna on the subtidal bivalve Chlamys opercularis. Mar. Biol. 110: 253–259
Winston, J. E. (1977). Feeding in marine bryozoans. In: Woollacott, R. M., Zimmer, R. L. (eds.) Biology of bryozoans. Academic Press, New York, p. 233–271
Winston, J. E. (1981). Feeding behavior of modern bryozoans. Univ. Tennessee Dept. Geol. Sci. Stud. Geol. 5: 1–21
Yoshioka, P. M. (1982). Predator-induced polymorphism in the bryozoan Membranipora membranacea. J. exp. mar. Biol. Ecol. 61: 233–242
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Communicated by J. Grassle, New Brunswick
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McKinney, F.K. Competitive interactions between related clades: evolutionary implications of overgrowth interactions between encrustin cyclostome and cheilostome bryozoans. Marine Biology 114, 645–652 (1992). https://doi.org/10.1007/BF00357261
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DOI: https://doi.org/10.1007/BF00357261