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" name="eprints.keywords" /> <meta content="Copyright by the Ecological Society of America." name="eprints.note" /> <meta content="There are two alternate community states in the rocky subtidal of the Atlantic coast of Nova Scotia, an unproductive sea urchin/coralline alga community, and highly productive kelp beds dominated by Laminaria longicruris. Disease-induced mortality of the sea urchins (Strongylocentrotus droebachiensis) triggered a switch from the first state to the second and provided a unique opportunity to study (1) the ability of L. longicruris to recover its former dominant status, and (2) its stability when competing with other seaweeds and when perturbed by storms and grazers other than urchins. Rates of recolo- nization of L. longicruris depended on the proximity of a refugial source of spores. When reproductive plants were nearby, a closed canopy developed within 18 mo of urchin mortality. When a reproductive population was several kilometres away, there was sparse recolonization for 3 yr, then a massive recruitment occurred with closure of the canopy in the 4th yr. Laminaria is clearly the competitive dominant in the seaweed community. Manipulative experiments showed that the kelp limits the abundance of several understory species, but there was no evidence that the abundant annual seaweeds limited kelp recruitment. When sea urchins were rare, the density and growth rates of Laminaria were influenced mostly by intraspecific competition. When the canopy of adult plants was removed there was a dramatic increase in kelp recruitment, but the recruits that grew in dense patches in the clearings were significantly smaller than those of a similar age that grew more sparsely beneath the canopy. Once the kelp recovered from destructive grazing and formed a mature forest, it was able to maintain its dominance, even in habitats subject to severe nutrient stress for 8 mo of the year. For most ofthe year mortality and erosion of laminae outweighed the effects of recruitment and growth, and the canopy declined, especially during winter when storms were frequent. Erosion was exacerbated by grazing of the gastropod Lacuna vincta. However, in late winter and early spring, recruitment and rapid growth restored the canopy. When severe storm damage was simulated by completely removing Laminaria in patches, the kelp rapidly recolonized and soon outgrew other seaweeds. Unlike the competitive dominants in kelp bed systems in the northeast Pacific, L . longicruris in Nova Scotia manifests multiple patterns of adaptation that enable it to dominate early and late stages of succession in a range of habitats of different levels of nutrient stress and of disturbance from storms and grazers. The principal threat to the stability of the kelp beds is destructive grazing by sea urchins. We suggest that the consid- erable differences between the dynamios of kelp beds in Nova Scotia and those of the northeast Pacific, and the high degree of stability of L . longicruris stands in Nova Scotia, is attributable to the low diversity of kelps and therefore low levels of competition in Nova Scotia, and to the multiple adaptations of L . longicruris that enable it to tolerate several stresses and disturbances. We argue that the dynamics of community organization, and therefore the stability properties of this system are determined primarily by biological interactions and not by physical variables. This differs from the kelp communities in the northeast Pacific, in which both biological and physical factors influence dynamics significantly at a primary level. We offer a qualitative model of the dynamics of community structure in Nova Scotia that may be viewed as a set of deterministic "subroutines," in which each subroutine describes the outcome of a particular biological interaction. The subroutine(s) that predominate at one point in time and space are probably determined mostly by physical hydrographic variables that have a large stochastic component. " name="eprints.abstract" /> <meta content="1988-06" name="eprints.date" /> <meta content="published" name="eprints.date_type" /> <meta content="Ecological Monographs" name="eprints.publication" /> <meta content="58" name="eprints.volume" /> <meta content="2" name="eprints.number" /> <meta content="129-154" name="eprints.pagerange" /> <meta content="10.2307/1942464" name="eprints.id_number" /> <meta content="UNSPECIFIED" name="eprints.thesis_type" /> <meta content="TRUE" name="eprints.refereed" /> <meta content="http://dx.doi.org/10.2307/1942464" name="eprints.official_url" /> <meta content="Ambrose, R. F., and B. V. Nelson. 1982. Inhibition of giant kelp recruitment by an introduced brown alga. Botanica Marina 25:265-267. Anderson, E. K., and W. J. North. 1966. In situ studies of spore production and dispersal in the giant kelp, Macro- cystis. Proceedings of the Fifth International Seaweed Sym- posium 5:73-86. Anderson, E. K., and W. J. North. 1969. Light requirements of juvenile and microscopic stages of giant kelp, Macro- cystis. Proceedings of the Sixth International Seaweed Sym- posium 6:3-15. Ayling, A. M. 1981. The role of biological disturbance in temperate subtidal encrusting communities. Ecology 62: 830-847. Bernstein, B. B., S. C. Schroeter, and K. H. Mann. 1983. Sea urchin (Strongylocentrotus droebachiensk) aggregating behaviour investigated by a subtidal multifactorial exper- iment. Canadian Journal of Fisheries and Aquatic Sciences 40:1975-1986. Bernstein, B. B., B. E. Williams, and K. H. Mann. 198 1. The role of behavioural responses to predators in modifying urchins' (Strongylocentrotus droebachiensis) destructive grazing and seasonal foraging patterns. Marine Biology 63: 39-49. Bird, C. J., and C. R. Johnson. 1984. Seirospora seirosperma (Harvey) Dixon (Rhodophyta, Ceramiaceae)-a first record for Canada. Proceedings of the Nova Scotian Institute of Science 34: 173-1 75. Black, R. 1974. Some biological interactions affecting in- tertidal populations of the kelp Egregia laevigata. Marine Biology 28: 189-198. Breen, P. A,, T. A. Carson, J. B. Foster, and E. A. Stuart. 1982. Changes in subtidal community structure associated with British Columbia sea otter transplants. Marine Ecol- ogy Progress Series 7: 13-20. Breen, P. A,, and K. H. Mann. 1976. Destructive grazing of kelp by sea urchins in eastern Canada. Journal of the Fisheries Research Board of Canada 33: 1278-1283. Chapman, A. R. 0 . 1974. The genetic basis of morpholog- ical differentiation in some Laminaria populations. Marine Biology 24:85-91. 198 1. Stability of sea urchin dominated barren grounds following destructive grazing of kelp in St. Mar- garet's Bay, eastern Canada. Marine Biology 62:307-3 1 l . . 1984. Reproduction, recruitment and mortality in two species ofLaminaria in southwest Nova Scotia. Journal of Experimental Marine Biology and Ecology 78:99-109. . 1986. Age versus stage: an analysis of age- and size- specific mortality and reproduction in a population of Lam- inarla longicruris Pyl. Journal of Experimental Marine Bi- ology and Ecology 97: 1 13-122. Chapman, A. R. O., and J. S. Craigie. 1977. Seasonal growth in Laminaria longicruris: relations with dissolved inorganic nutrients and internal reserves of nitrogen. Marine Biology 40:197-205. Connell, J. H., and R. 0 . Slayter. 1977. Mechanisms of succession in natural communities and their role in com- munity stability and organization. American Naturalist 111: 11 19-1 144. Connell, J. H., and W. P. Sousa. 1983. On the evidence needed to judge ecological stability or persistence. Ameri- can Naturalist 121 :789-824. Cousens, R., and M. J. Hutchings. 1983. The relationship between density and mean frond weight in monospecific seaweed stands. Nature 301:240-24 1. Cowen, R. K., C. R. Agegian, and M. S. Foster. 1982. The maintenance of community structure in a central California giant kelp forest. Journal of Experimental Marine Biology and Ecology 64: 189-20 1. Dayton, P. K. 1971. Competition, disturbance, and com- munity organization: the provision and subsequent utili- zation of space in a rocky intertidal community. Ecological Monographs 41:35 1-389. . 1973. Dispersion, dispersal, and persistence of the annual intertidal alga, Postelsia palmaeformis Ruprecht. Ecology 54:433-438. . 1975a. Experimental evaluation of ecological dom- inance in a rocky intertidal algal community. Ecological Monographs 45: 137-1 59. . 19753. Experimental studies of algal canopy inter- actions in a sea otter-dominated kelp community at Am- chitka Island, Alaska. United States National Marine Fish- eries Service Fishery Bulletin 73:230-237. . 1985. The structure and regulation of some South American kelp communities. Ecological Monographs 55: 447-468. Dayton, P.K., Curric, T., Gerrodette, B.D., Keller, R. Rosenthal, and D. Ven Tresca. 1984. Patch dynamics and stability of some California kelp communities. Ecological Monographs 54:253-289. Dayton, P. K., and M. J. Tegner. 1984. Catastrophic storms, El Nifio, and patch stability in a southern California kelp community. Science 224:283-285. Dean, T. A., and F. R. Jacobsen. 1984. Growth of juvenile Macrocystis pyrifera (Laminariales) in relation to environ- mental factors. Marine Biology 83:301-3 1 1. Dean, T. A,, S. C. Schroeter, and J. D. Dixon. 1984. Effects of grazing by two species of sea urchins (Strongylocentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp (Macrocystis pyrifera and Pterygophora calfornica). Marine Biology 78:30 1-3 13. Dethier, M. N. 1984. Disturbance and recovery in intertidal pools: maintenance of mosaic patterns. Ecological Mono- graphs 54:99-118. De Wreede, R. E. 1984. Growth and age class distribution of Pterygophora calfornica (Phaeophyta). Marine Ecology Progress Series 19:93-100. Dring, M. J. 1982. The biology of marine plants. Edward Arnold, London, England. Druehl, L. D. 1978. The distribution of Macrocystis integ- rlfolia in British Columbia as related to environmental pa- rameters. Canadian Journal of Botany 56:69-79. Druehl, L. D., and P. A. Breen. 1986. Some ecological effects of harvesting Macrocystis integrifolia. Botanica Marina 29: 97-103. Druehl, L. D., and W. N. Wheeler. 1986. Population biology of Macrocystis integrifolia from British Columbia, Canada. Marine Biology 90: 173-1 79. Duggins, D. 0 . 1980. Kelp beds and sea otters: an experi- mental approach. Ecology 61:447-453. 198 1. Sea urchins and kelp: the effects of short term changes in urchin diet. Limnology and Oceanography 26: 391-394. Edelstein, T., J. S. Craigie, and J. McLachlan. 1969. Prelim- inary survey of the sublittoral flora of Halifax county. Jour- nal of the Fisheries Research Board of Canada 26:2703- 27 13. Espinoza, J., and A. R. 0 . Chapman. 1983. Ecotypic dif- ferentiation of Laminaria longicruris in relation to seawater nitrate concentration. Marine Biology 74:2 13-2 18. Estes, J. A., and J. F. Palmisano. 1974. Sea otters: their role in structuring nearshore communities. Science 185:1058- 1060. Estes, J. A, N. S. Smith, and J. F. Palmisano. 1978. Sea otter predation and community organization in the western Aleutian Islands, Alaska. Ecology 59:822-833. Foreman, R. E. 1977. Benthic community modification and recovery following intensive grazing by Strongylocentrotus droebachiensis. Helgolander wissenschaftliche Meeresun- tersuchungen 30:468-484. Foster, M. S. 1975a. Algal succession in a Macrocystis py- rifera Forest. Marine Biology 32:3 13-329. -. 19756. Regulation of algal community development in a Macrocystis pyrifera forest. Marine Biology 32:331- 342. Gagnk, J. A,, K. H. Mann, and A. R. 0 . Chapman. 1982. Seasonal patterns of growth and storage in Laminaria lon- gicruris in relation to differing patterns of availability of nitrogen in the water. Marine Biology 69:91-101. Gerard, V. A. 1982. Growth and utilization of internal ni- trogen reserves by the giant kelp Macrocystis pyrlfera in a low-nitrogen environment. Marine Biology 66:27-35. -. 1984a. Physiological effects of El Nifio on giant kelp in southern California. Marine Biology Letters 5:3 17-322. -. 1984b. The light environment in a giant kelp forest: influence of Macrocystis pyrifera on spatial and temporal variability. Marine Biology 84: 189-1 95. Gerard, V. A,, and K. H. Mann. 1979. Growth and repro- duction of Laminaria longicruris (Phaeophyta) populations exposed to different intensities of water movement. Journal of Phycology 15:33-41. Harrold, C., and D. C. Reed. 1985. Food availability, sea urchin grazing, and kelp forest community structure. Ecol- ogy 66:1160-1169. Hay, C. H., and G. R. South. 1979. Experimental ecology with particular reference to proposed commercial harvest- ing of Durv~llaea (Phaeophyta, Durvilleales) in New Zea- land. Botanica Marina 22:431-436. Hoffmann, A. J., and R. Ugarte. 1985. The anival of prop- agules of marine macroalgae in the intertidal zone. Journal of Experimental Marine Biology and Ecology 92:83-95. Holling, C. S. 1973. Resilience and stability of ecological systems. Annual Review of Ecology and Systematics 4:l- 23. Horn, H. S. 198 1. Succession. Pages 253-27 1 in R. M. May, editor. Theoretical ecology. Second edition. Sinauer As- sociates, Sunderland, Massachusetts, USA. JernakofF, P. 1985. Temporal and small-scale spatial vari- ability of algal abundance on an intertidal rocky shore. Botanica Marina 28:145-154. Johnson, C. R. 1984. Ecology of the kelp Laminaria lon- gicruris and its principal grazers in the rocky subtidal of Nova Scotia. Dissertation. Dalhousie University, Halifax, Nova Scotia, Canada. Johnson, C. R., and K. H. Mann. 1982. Adaptations of Strongylocentrotus droebachiensis for survival on barren grounds. Pages 277-283 in J. M. Lawrence, editor. Echi- noderms: Proceeedings of the International Conference, Tampa Bay. A. A. Balkema, Rotterdam, The Netherlands. Johnson, C. R., and K. H. Mann. 1986a. The crustose cor- alline alga, Phymatolithon Foslie, inhibits the overgrowth of seaweeds without relying on herbivores. Journal of Ex- perimental Marine Biology and Ecology 96: 127-1 46. Johnson, C. R., and K. H. Mann. 1986b. The importance of plant defence abilities to the structure of subtidal seaweed communities: the kelp Laminaria longicruris de la Pylaie survives grazing by the snail Lacuna vincta (Montagu) at high population densities. Journal of Experimental Marine Biology and Ecology 97:231-267. Jones, G. M. 1985. Paramoeba invadens n. sp. (Amoebida, Paramoebidae), a pathogenic amoeba from the sea urchin, Strongylocentrotus droebachiensis, in eastern Canada. Jour- nal of Protozoology 32:564-569. Jones, G. M., A. J. Hebda, R. E. Scheibling, and R. J. Miller. 1985. Histopathology of the disease causing mass mortality of sea urchins (Strongylocentrotus droebachiensis) in Nova Scotia Journal of Invertebrate Pathology 45:260-27 1. Jones, G. M., and R. E. Scheibling. 1985. Paramoeba sp. (Amoebida, Paramoebidae) as the possible causative agent of sea urchin mass mortality in Nova Scotia. Journal of Parasitology 71 :559-565. Kain, J. M. 1979. A view of the genus Laminaria. Ocean- ography and Marine Biology Annual Reviews 17: 10 1-1 6 1. Keats, D. W. 1986. Comment on "Seaweeds, sea urchins, and lobsters: a reappraisal" by R. J. Miller. Canadian Jour- nal of Fisheries and Aquatic Sciences 43: 1675-1 676. Keats, D. W., G. R. South, and D. H. Steele. 1982. The occurrence of Agarum cribrosum (Mert.) Bory (Phaeophyta, Laminariales) in relation to some of its competitors and predators in Newfoundland. Phycologia 21: 189-1 9 1. Kimura, R. S., and M. S. Foster. 1984. The effects of har- vesting Macrocystis pyrifera on the algal assemblage in a giant kelp forest. Hydrobiologia 116/117:425-428. Kirkman, H. 198 1. The first year in the life history and the survival of the marine macrophyte, Ecklonia radiata (Turn.) J. Agardh. Journal of Experimental Marine Biology and Ecology 55:243-254. Leighton, D. L. 197 1. Grazing activities of benthic inver- tebrates in southern California kelp beds. Nova Hedwigia Beihefte, Supplement Number 32:42 1-453. Littler, M. M., D. S. Littler, and P. R. Taylor. 1983. Evolutionary strategies in a tropical barrier reef system: func- tional-form groups of marine macroalgae. Journal of Phy- cology 19:229-237. Lobban, C. S., P. J. Harrison, and M. J. Duncan. 1985. The physiological ecology of seaweeds. Cambridge University Press, Cambridge, England. Lubchenco, J. 1980. Algal zonation in the New England rocky intertidal community: an experimental analysis. Ecology 61:333-344. Lubchenco, J., and B. A. Menge. 1978. Community devel- opment and persistence in a low rocky intertidal zone. Eco- logical Monographs 48:67-94. MacFarlane, C. 1952. A survey of certain seaweeds of com- mercial importance in southwest Nova Scotia. Canadian Journal of Botany 30:78-97. MacFarlane, C., and H. P. Bell. 1933. Observations of the seasonal changes in the marine algae in the vicinity of Hal- ifax, with particular reference to winter conditions. Pro- ceedings of the Nova Scotian Institute of Science 18: 134- 176. Mann, K. H. 1972a. Ecological energetics of the seaweed zone in a marine bay on the Atlantic coast of Canada. I. Zonation and biomass of seaweeds. Marine Biology 12:l- 10. 19726. Ecological energetics of the seaweed zone in a marine bay on the Atlantic coast of Canada. 11. Produc- tivity of the seaweeds. Marine Biology 14: 199-209. 1977. Destruction of kelp-beds by sea-urchins: a cyclical phenomenon or irreversible degradation? Helgo- lfinder wissenschaftliche Meeresuntersuchungen 30:455467. 1985. Invertebrate behaviour and the structure of marine benthic communities. Pages 227-245 in R. M. Sibly and R. H. Smith, editors. Behavioural ecology: ecological consequences of adaptive behaviour. Blackwell Scientific, Oxford, England. Mathieson, A. C., and J. S. Prince. 1973. Ecology of Chon- drus crispus Stackhouse. Pages 53-79 in M. J. Harvey and J. McLachlan, editors. Chondrus crispus. Nova Scotian In- stitute of Science, Halifax, Nova Scotia, Canada. Mattison, J. E., J. D. Trent, A. L. Shanks, T. B. Akin, and J. S. Pearse. 1977. Movement and feeding activity of red sea urchins (Strongylocentrotusfranciscanus) adjacent to a kelp forest. Marine Biology 39:25-30. Menge, B. A,, and J. P. Sutherland. 1976. Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. American Naturalist 110:35 1- 369. Miller, R. J. 1 9 8 5a. Succession in sea urchin and seaweed abundance in Nova Scotia, Canada. Marine Biology 84: 275-286. -. 1985b. Seaweeds, sea urchins and lobsters: a reap- praisal. Canadian Journal of Fisheries and Aquatic Sciences 421206 1-2072. Miller, R. J., and A. G. Colodey. 1983. Widespread mass mortalities of the green sea urchin in Nova Scotia, Canada. Marine Biology 73:263-276. Moore, D. S., and R. J. Miller. 1983. Recovery of macro- algae following widespread sea urchin mortality with a de- scription of the nearshore hard-bottom habitat on the At- lantic coast of Nova Scotia. Canadian Technical Report of Fisheries and Aquatic Sciences 1230. Moreno, C. A,, and J. P. Sutherland. 1982. Physical and biological processes in a Macrocystis pyrifera community near Valdivia, Chile. Oecologia (Berlin) 55: 1-6. Neushul, M. 1967. Studies of subtidal marine vegetation in western Washington. Ecology 4833-94. North, W. J., and J. S. Pearse. 1969. Sea urchin population explosion in southern California coastal waters. Science 167:209. North, W. J., and R. C. Zimmerman. 1984. Influences of macronutrients and water temperatures on summertime survival of Macrocystis canopies. Hydrobiologia 116/117: 4 19-424. Novaczek, I., and J. McLachlan. 1986. Recolonization by algae of the sublittoral habitat of Halifax County, Nova Scotia, following the demise of sea urchins. Botanica Ma- rina 29:69-73. Pace, D. 1981. Kelp community development in Barkley Sound, British Columbia following sea urchin removal. Proceedings of the Eighth International Seaweed Sympo- sium 8:457-463. Padilla, D. K. 1984. The importance of form: differences in competitive ability, resistance to consumers and environ- mental stress in an assemblage of coralline algae. Journal of Experimental Marine Biology and Ecology 79: 105-127. Paine, R. T. 1979. Disaster, catastrophe, and local persis- tence of the sea palm Postelsia palmaeformis. Science 205: 685-687. -. 1980. Food webs: linkage, interaction strength and community infrastructure. Journal of Animal Ecology 49: 667-685. Paine, R. T., J. C. Castillo, and J. Cancino. 1985. Perturbation and recovery patterns of starfish-dominated inter- tidal assemblages in Chile, New Zealand, and Washington State. American Naturalist 125:679-69 1. Paine, R. T., and S. A. Levin. 1981. Intertidal landscapes: disturbance and the dynamics of pattern. Ecological Mono- graphs 51: 145-178. Paine, R. T., and R. L. Vadas. 1969. The effects of grazing by sea urchins, Strongylocentrotus spp., on benthic algal populations. Limnology and Oceanography 14:7 10-7 19. Pearse, J. S., and A. H. Hines. 1979. Expansion of a central California kelp forest following the mass mortality of sea urchins. Marine Biology 51:83-91. Peckol, P., and R. B. Searles. 1983. Effects of seasonality and disturbance on population development in a Carolina continental shelf community. Bulletin of Marine Science 33:67-86. Pimm, S. L. 1982. Food webs. Chapman and Hall, London, England. -. 1984. The complexity and stability of ecosystems. Nature 307:32 1-326. Press, S. J. 1972. Applied multivariate analysis. Holt, Rine- hart and Winston, New York, New York, USA. Pringle, J. D. 1986. A review of urchin-macroalgal associ- ations with a new synthesis for nearshore Canadian waters. Pages 191-218 in B. Santelices, editor. Usos y funciones ecologicas de las algas marinas bentonicas. Monografias Biologicas 4. Reed, D. C., and M. S. Foster. 1984. The effects of canopy shading on algal recruitment and growth in a giant kelp forest. Ecology 65:937-948. Rosenthal, R. J., W. D. Clarke, and P. K. Dayton. 1974. Ecology and natural history of a stand of giant kelp, Macro- cystis pyrifera, off Del Mar, California. United States Na- tional Marine Fisheries Service Fishery Bulletin 72:670- 684. Santelices, B., J. C. Castilla, J. Cancino, and P. Schmiede. 1980. Comparative ecology of Lessonia nigrescens and Durvillaea antarctica (Phaeophyta) in central Chile. Marine Biology 59: 1 19-1 32. Santelices, B., and F. P. Ojeda. 1984a. Effects of canopy removal on the understory algal community structure of coastal forests of Macrocystispyrifera from southern South America. Marine Ecology Progress Series 14: 165-1 73. Santelices, B., and F. P. Ojeda. 1984b. Population dynamics of coastal forests of Macrocystis pyrifera in Puerto Toro, Isla Navarino, Southern Chile. Marine Ecology Progress Series 14: 175-183. Scheibling, R. E. 1986. Increased macroalgal abundance following mass mortalities of sea urchins (Strongylocentro- tus droebachiensis) along the Atlantic coast of Nova Scotia. Oecologia (Berlin) 68: 186-1 98. Scheibling, R. E., and R. L. Stephenson. 1984. Mass mor- tality of Strongylocentrotus droebachiensis (Echinodermata: Echinoidea) off Nova Scotia, Canada. Marine Biology 78: 153-164. Schiel, D. R. 1985. Growth, survival and reproduction of two species of marine algae at different densities in natural stands. Journal of Ecology 73: 199-2 17. Simenstad, C. A,, J. A. Estes and K. W. Kenyon. 1978. Aleuts, sea otters, and alternate stable-state communities. Science 200:403-4 1 1. Smith, B. D. 1985. Recovery following experimental har- vesting of Laminaria longzcruris and L. digitata in south- western Nova Scotia. Helgolander wissenschaftliche Meeresuntersuchungen 39:83-10 1. Sousa, W. P. 1979a. Disturbance in marine intertidal boul- der fields: the nonequilibrium maintenance of species di- versity. Ecology 60: 1225-1239. -. 19796. Experimental investigations of disturbance and ecological succession in a rocky intertidal algal com- munity. Ecological Monographs 49:227-254. -. 1984. Intertidal mosaics: patch size, propagule availability, and spatially variable patterns of succession. Ecology 65:1918-1935. Sousa, W. P., S. C. Schroeter, and S. D. Gaines. 1981. Lat- itudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation. Oeco- logia (Berlin) 48:297-307. South, G. R., and R. G. Hooper. 1980. A catalogue and atlas ofthe benthic marine algae of the island of Newfound- land. Memorial University of Newfoundland Occasional Papers in Biology Number 13. Steneck, R. S. 1983. Escalating herbivory and resulting adaptive trends in calcareous algal crusts. Paleobiology 9: 44-6 1. -. 1986. The ecology of coralline algal crusts: conver- gent patterns and adaptive strategies. Annual Review of Ecology and Systematics 17:273-303. Sutherland, J. P. 198 1. The fouling community at Beaufort, North Carolina: a study in stability. American Naturalist 1181499-5 19. Sutherland, J. P., and R. H. Karlson. 1977. Development and stability of the fouling community at Beaufort, North Carolina. Ecological Monographs 47:425-446. Tremblay, C., and A. R. 0 . Chapman. 1980. The local occurrence of Agarum cribrosum in relation to the presence or absence of its competitors and predators. Proceedings of the Nova Scotian Institute of Science 30: 165-170. Vadas, R. L. 1968. The ecology of Agarum and the kelp bed community. Dissertation. University of Washington, Seattle, Washington, U.S.A. Vadas, R. L., R. W. Elner, P. E. Ganvood, and I. G. Babb. 1986. Experimental evaluation of aggregation behaviour in the sea urchin Strongylocentrotus droebachiensis. Marine Biology 90:433-448. Velimirov, B., and C.L. Griffiths. 1979. Wave-induced kelp movement and its importance for community structure. Botanica Marina 22: 169-1 72. Wharton, W. G., and K. H. Mann. 198 1. Relationship be- tween destructive grazing by the sea urchin, Strongylocentrotus droebachiensis, and the abundance of American lob- ster, Homarus americanus, on the Atlantic coast of Nova Scotia. Canadian Journal of Fisheries and Aquatic Sciences 38: 1339-1349. Yodzis, P. 198 1. The stability of real ecosystems. Nature 2891674-676." name="eprints.referencetext" /> <meta content="Johnson, Craig R. and Mann, Kenneth H. (1988) Diversity, patterns of adaptation, and stability of Nova Scotian kelp beds. Ecological Monographs, 58 (2). pp. 129-154." name="eprints.citation" /> <meta content="http://eprints.utas.edu.au/1237/1/Johnson_and_Mann_1988.pdf" name="eprints.document_url" /> <link rel="schema.DC" href="http://purl.org/DC/elements/1.0/" /> <meta content="Diversity, patterns of adaptation, and stability of Nova Scotian kelp beds" name="DC.title" /> <meta content="Johnson, Craig R." name="DC.creator" /> <meta content="Mann, Kenneth H." name="DC.creator" /> <meta content="270702 Marine and Estuarine Ecology (incl. Marine Ichthyology)" name="DC.subject" /> <meta content="There are two alternate community states in the rocky subtidal of the Atlantic coast of Nova Scotia, an unproductive sea urchin/coralline alga community, and highly productive kelp beds dominated by Laminaria longicruris. Disease-induced mortality of the sea urchins (Strongylocentrotus droebachiensis) triggered a switch from the first state to the second and provided a unique opportunity to study (1) the ability of L. longicruris to recover its former dominant status, and (2) its stability when competing with other seaweeds and when perturbed by storms and grazers other than urchins. Rates of recolo- nization of L. longicruris depended on the proximity of a refugial source of spores. When reproductive plants were nearby, a closed canopy developed within 18 mo of urchin mortality. When a reproductive population was several kilometres away, there was sparse recolonization for 3 yr, then a massive recruitment occurred with closure of the canopy in the 4th yr. Laminaria is clearly the competitive dominant in the seaweed community. Manipulative experiments showed that the kelp limits the abundance of several understory species, but there was no evidence that the abundant annual seaweeds limited kelp recruitment. When sea urchins were rare, the density and growth rates of Laminaria were influenced mostly by intraspecific competition. When the canopy of adult plants was removed there was a dramatic increase in kelp recruitment, but the recruits that grew in dense patches in the clearings were significantly smaller than those of a similar age that grew more sparsely beneath the canopy. Once the kelp recovered from destructive grazing and formed a mature forest, it was able to maintain its dominance, even in habitats subject to severe nutrient stress for 8 mo of the year. For most ofthe year mortality and erosion of laminae outweighed the effects of recruitment and growth, and the canopy declined, especially during winter when storms were frequent. Erosion was exacerbated by grazing of the gastropod Lacuna vincta. However, in late winter and early spring, recruitment and rapid growth restored the canopy. When severe storm damage was simulated by completely removing Laminaria in patches, the kelp rapidly recolonized and soon outgrew other seaweeds. Unlike the competitive dominants in kelp bed systems in the northeast Pacific, L . longicruris in Nova Scotia manifests multiple patterns of adaptation that enable it to dominate early and late stages of succession in a range of habitats of different levels of nutrient stress and of disturbance from storms and grazers. The principal threat to the stability of the kelp beds is destructive grazing by sea urchins. We suggest that the consid- erable differences between the dynamios of kelp beds in Nova Scotia and those of the northeast Pacific, and the high degree of stability of L . longicruris stands in Nova Scotia, is attributable to the low diversity of kelps and therefore low levels of competition in Nova Scotia, and to the multiple adaptations of L . longicruris that enable it to tolerate several stresses and disturbances. We argue that the dynamics of community organization, and therefore the stability properties of this system are determined primarily by biological interactions and not by physical variables. This differs from the kelp communities in the northeast Pacific, in which both biological and physical factors influence dynamics significantly at a primary level. We offer a qualitative model of the dynamics of community structure in Nova Scotia that may be viewed as a set of deterministic "subroutines," in which each subroutine describes the outcome of a particular biological interaction. The subroutine(s) that predominate at one point in time and space are probably determined mostly by physical hydrographic variables that have a large stochastic component. " name="DC.description" /> <meta content="1988-06" name="DC.date" /> <meta content="Article" name="DC.type" /> <meta content="PeerReviewed" name="DC.type" /> <meta content="application/pdf" name="DC.format" /> <meta content="http://eprints.utas.edu.au/1237/1/Johnson_and_Mann_1988.pdf" name="DC.identifier" /> <meta content="http://dx.doi.org/10.2307/1942464" name="DC.relation" /> <meta content="Johnson, Craig R. and Mann, Kenneth H. (1988) Diversity, patterns of adaptation, and stability of Nova Scotian kelp beds. 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border: solid 1px #ccc; padding: 3px"><tr> <td align="left"><a href="http://eprints.utas.edu.au/cgi/users/home">Login</a> | <a href="http://eprints.utas.edu.au/cgi/register">Create Account</a></td> <td align="right" style="white-space: nowrap"> <form method="get" accept-charset="utf-8" action="http://eprints.utas.edu.au/cgi/search" style="display:inline"> <input class="ep_tm_searchbarbox" size="20" type="text" name="q" /> <input class="ep_tm_searchbarbutton" value="Search" type="submit" name="_action_search" /> <input type="hidden" name="_order" value="bytitle" /> <input type="hidden" name="basic_srchtype" value="ALL" /> <input type="hidden" name="_satisfyall" value="ALL" /> </form> </td> </tr></table></td></tr> <tr> <td class="toplinks"><!-- InstanceBeginEditable name="content" --> <div align="center"> <table width="720" class="ep_tm_main"><tr><td align="left"> <h1 class="ep_tm_pagetitle">Diversity, patterns of adaptation, and stability of Nova Scotian kelp beds</h1> <p style="margin-bottom: 1em" class="not_ep_block"><span class="person_name">Johnson, Craig R.</span> and <span class="person_name">Mann, Kenneth H.</span> (1988) <xhtml:em>Diversity, patterns of adaptation, and stability of Nova Scotian kelp beds.</xhtml:em> Ecological Monographs, 58 (2). pp. 129-154.</p><p style="margin-bottom: 1em" class="not_ep_block"></p><table style="margin-bottom: 1em" class="not_ep_block"><tr><td valign="top" style="text-align:center"><a href="http://eprints.utas.edu.au/1237/1/Johnson_and_Mann_1988.pdf"><img alt="[img]" src="http://eprints.utas.edu.au/style/images/fileicons/application_pdf.png" border="0" class="ep_doc_icon" /></a></td><td valign="top"><a href="http://eprints.utas.edu.au/1237/1/Johnson_and_Mann_1988.pdf"><span class="ep_document_citation">PDF</span></a> - Full text restricted - Requires a PDF viewer<br />5Mb</td><td><form method="get" accept-charset="utf-8" action="http://eprints.utas.edu.au/cgi/request_doc"><input value="1605" name="docid" accept-charset="utf-8" type="hidden" /><div class=""><input value="Request a copy" name="_action_null" class="ep_form_action_button" onclick="return EPJS_button_pushed( '_action_null' )" type="submit" /> </div></form></td></tr></table><p style="margin-bottom: 1em" class="not_ep_block">Official URL: <a href="http://dx.doi.org/10.2307/1942464">http://dx.doi.org/10.2307/1942464</a></p><div class="not_ep_block"><h2>Abstract</h2><p style="padding-bottom: 16px; text-align: left; margin: 1em auto 0em auto">There are two alternate community states in the rocky subtidal of the Atlantic coast of Nova Scotia, an unproductive sea urchin/coralline alga community, and highly productive kelp beds dominated by Laminaria longicruris. Disease-induced mortality of the sea urchins (Strongylocentrotus droebachiensis) triggered a switch from the first state to the second and provided a unique opportunity to study (1) the ability of L. longicruris to recover its former dominant status, and (2) its stability when competing with other seaweeds and when perturbed by storms and grazers other than urchins. Rates of recolo- nization of L. longicruris depended on the proximity of a refugial source of spores. When reproductive plants were nearby, a closed canopy developed within 18 mo of urchin mortality. When a reproductive population was several kilometres away, there was sparse recolonization for 3 yr, then a massive recruitment occurred with closure of the canopy in the 4th yr. Laminaria is clearly the competitive dominant in the seaweed community. Manipulative experiments showed that the kelp limits the abundance of several understory species, but there was no evidence that the abundant annual seaweeds limited kelp recruitment. When sea urchins were rare, the density and growth rates of Laminaria were influenced mostly by intraspecific competition. When the canopy of adult plants was removed there was a dramatic increase in kelp recruitment, but the recruits that grew in dense patches in the clearings were significantly smaller than those of a similar age that grew more sparsely beneath the canopy. Once the kelp recovered from destructive grazing and formed a mature forest, it was able to maintain its dominance, even in habitats subject to severe nutrient stress for 8 mo of the year. For most ofthe year mortality and erosion of laminae outweighed the effects of recruitment and growth, and the canopy declined, especially during winter when storms were frequent. Erosion was exacerbated by grazing of the gastropod Lacuna vincta. However, in late winter and early spring, recruitment and rapid growth restored the canopy. When severe storm damage was simulated by completely removing Laminaria in patches, the kelp rapidly recolonized and soon outgrew other seaweeds. Unlike the competitive dominants in kelp bed systems in the northeast Pacific, L . longicruris in Nova Scotia manifests multiple patterns of adaptation that enable it to dominate early and late stages of succession in a range of habitats of different levels of nutrient stress and of disturbance from storms and grazers. The principal threat to the stability of the kelp beds is destructive grazing by sea urchins. We suggest that the consid- erable differences between the dynamios of kelp beds in Nova Scotia and those of the northeast Pacific, and the high degree of stability of L . longicruris stands in Nova Scotia, is attributable to the low diversity of kelps and therefore low levels of competition in Nova Scotia, and to the multiple adaptations of L . longicruris that enable it to tolerate several stresses and disturbances. We argue that the dynamics of community organization, and therefore the stability properties of this system are determined primarily by biological interactions and not by physical variables. This differs from the kelp communities in the northeast Pacific, in which both biological and physical factors influence dynamics significantly at a primary level. We offer a qualitative model of the dynamics of community structure in Nova Scotia that may be viewed as a set of deterministic "subroutines," in which each subroutine describes the outcome of a particular biological interaction. The subroutine(s) that predominate at one point in time and space are probably determined mostly by physical hydrographic variables that have a large stochastic component. </p></div><table style="margin-bottom: 1em" border="0" cellpadding="3" class="not_ep_block"><tr><th valign="top" class="ep_row">Item Type:</th><td valign="top" class="ep_row">Article</td></tr><tr><th valign="top" class="ep_row">Additional Information:</th><td valign="top" class="ep_row">Copyright by the Ecological Society of America.</td></tr><tr><th valign="top" class="ep_row">Keywords:</th><td valign="top" class="ep_row">community structure; competition; disturbance; diversity; kelp; Laminaria; multiple adaptations; Nova Scotia; sea urchins; stability; stress; succession. </td></tr><tr><th valign="top" class="ep_row">Subjects:</th><td valign="top" class="ep_row"><a href="http://eprints.utas.edu.au/view/subjects/270702.html">270000 Biological Sciences > 270700 Ecology and Evolution > 270702 Marine and Estuarine Ecology (incl. Marine Ichthyology)</a></td></tr><tr><th valign="top" class="ep_row">Collections:</th><td valign="top" class="ep_row">UNSPECIFIED</td></tr><tr><th valign="top" class="ep_row">ID Code:</th><td valign="top" class="ep_row">1237</td></tr><tr><th valign="top" class="ep_row">Deposited By:</th><td valign="top" class="ep_row"><span class="ep_name_citation"><span class="person_name">Professor Craig R. Johnson</span></span></td></tr><tr><th valign="top" class="ep_row">Deposited On:</th><td valign="top" class="ep_row">25 Jun 2007</td></tr><tr><th valign="top" class="ep_row">Last Modified:</th><td valign="top" class="ep_row">04 Feb 2008 17:28</td></tr><tr><th valign="top" class="ep_row">ePrint Statistics:</th><td valign="top" class="ep_row"><a target="ePrintStats" href="/es/index.php?action=show_detail_eprint;id=1237;">View statistics for this ePrint</a></td></tr></table><p align="right">Repository Staff Only: <a href="http://eprints.utas.edu.au/cgi/users/home?screen=EPrint::View&eprintid=1237">item control page</a></p> </td></tr></table> </div> <!-- InstanceEndEditable --></td> </tr> <tr> <td><!-- #BeginLibraryItem "/Library/footer_eprints.lbi" --> <table width="795" border="0" align="left" cellpadding="0" class="footer"> <tr valign="top"> <td colspan="2"><div align="center"><a href="http://www.utas.edu.au">UTAS home</a> | <a href="http://www.utas.edu.au/library/">Library home</a> | <a href="/">ePrints home</a> | <a href="/contact.html">contact</a> | <a href="/information.html">about</a> | <a href="/view/">browse</a> | <a href="/perl/search/simple">search</a> | <a href="/perl/register">register</a> | <a href="/perl/users/home">user area</a> | <a href="/help/">help</a></div><br /></td> </tr> <tr><td colspan="2"><p><img src="/images/eprints/footerline.gif" width="100%" height="4" /></p></td></tr> <tr valign="top"> <td width="68%" class="footer">Authorised by the University Librarian<br /> © University of Tasmania ABN 30 764 374 782<br /> <a href="http://www.utas.edu.au/cricos/">CRICOS Provider Code 00586B</a> | <a href="http://www.utas.edu.au/copyright/copyright_disclaimers.html">Copyright & Disclaimers</a> | <a href="http://www.utas.edu.au/accessibility/index.html">Accessibility</a> | <a href="http://eprints.utas.edu.au/feedback/">Site Feedback</a> </td> <td width="32%"><div align="right"> <p align="right" class="NoPrint"><a href="http://www.utas.edu.au/"><img src="http://www.utas.edu.au/shared/logos/unioftasstrip.gif" alt="University of Tasmania Home Page" width="260" height="16" border="0" align="right" /></a></p> <p align="right" class="NoPrint"><a href="http://www.utas.edu.au/"><br /> </a></p> </div></td> </tr> <tr valign="top"> <td><p> </p></td> <td><div align="right"><span class="NoPrint"><a href="http://www.eprints.org/software/"><img src="/images/eprintslogo.gif" alt="ePrints logo" width="77" height="29" border="0" align="bottom" /></a></span></div></td> </tr> </table> <!-- #EndLibraryItem --> <div align="center"></div></td> </tr> </table> </body> </html>