Abstract
Although they are subjected to one of the most stressful physical environments on earth, wave-swept rocky shores support a highly diverse community of plants and animals. The surprising presence of such diversity amidst severe environmental adversity provides a unique opportunity for exploration of the role of extreme water flows in community ecology and natural selection. Methods are described by which the maximal water velocity and acceleration can be predicted for a site on the shore, and from these values maximal hydrodynamic forces are calculated. These forces can limit the range and foraging activity of some species, and can determine the rate of disturbance in others, but in general, wave-swept organisms have surprisingly high factors of safety. This apparent over-design can help to explain the diversity of forms present on wave-swept shores, and provides examples of how mechanics can limit the ability of natural selection to guide specialization. Although flow itself may commonly be prohibited from selecting for optima in morphology, it nonetheless continues to play a potentially important role in evolution by providing a mechanism for breaking or dislodging individuals that have been selected by other means.
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References
Abelson A. and Denny M. (1997). Settlement of marine organisms in flow. Ann. Rev. Ecol. Syst. 28: 317–339
Alexander R.McN. (1981). Factors of safety in the structure of animals. Sci. Progr. Oxf. 67: 109–130
Batchelor G.K. (1967). An Introduction to Fluid Dynamics. Cambridge University Press, Cambridge, UK
Bascomb W. (1980). Waves and Beaches. Ancor Press/Doubleday, Garden City, NY
Battjes J.A. 1970. Long-term wave height distribution at seven stations around the British Isles. National Institute of Oceanography (UK) Internal Report Number A.44.
Bell E.C. (1993). Mar. Biol. 117: 337–346
Bell E.C. and Denny M.W. (1994). Quantifying ‘wave exposure’: a simple device for recording maximum velocity and results of its use at several field sites. J. Exp. Mar. Biol. Ecol. 181: 9–29
Blanchette C.A. (1996). J. Exp. Mar. Biol. Ecol. 197: 1–14
Blanchette C.A. (1997). Ecology 78: 1563–1578
Carrington E. (1990). Drag and dislodgment of an intertidal macroalga: consequences of morphological variation in Mastocarpus papillatus Kützing. J. Exp. Mar. Biol. Ecol. 139: 185–200
Carrington E. (2002). Seasonal variation in the attachment strength of blue mussels: causes and consequences. Limnol. Oceanogr. 47: 1723–1733
Connell J.H. (1961). Ecology 42: 710–723
Connell J.H. (1972). Community interactions on marine rocky intertidal shores. Ann. Rev. Ecol. Syst. 3: 169–192
Connell J.H. (1978). Diversity in tropical rainforests and coral reefs. Science 199: 1302–1310
Dayton P.K. (1971). Competition, disturbanceand community organization: the provision and subsequent utilization of space in a rocky intertidal community. Ecol. Monogr. 45: 137–159
Dayton P.K. (1985). Ecology of kelp communities. Ann. Rev. Ecol. Syst. 16: 215–246
Dayton P.K., Currie V., Gerrodette T., Keller B. and Rosenthal R. (1984). Patch dynamics and stability of some southern California kelp communities. Ecol. Monogr. 54: 253–289
Dayton P.K., Tegner M.J., Parnell P.E. and Edwards P.B. (1992). Temporal and spatial patterns of disturbance and recovery in a kelp forest community. Ecol. Monogr. 62: 421–445
Denny M.W. (1988). Biology and the Mechanics of the Wave-Swept Environment. Princeton University Press, Princeton, NJ
Denny M. (1989). A limpet shell shape that reduces drag: laboratory demonstration of a hydrodynamic mechanism and an exploration of its effectiveness in nature. Can. J. Zool. 67: 2098–2106
Denny M.W. (1995). Predicting physical disturbance: mechanistic approaches to the study of survivorship on wave-swept shores. Ecol. Monogr. 65: 371–418
Denny M.W. (2000). Are there mechanical limits to size in wave-swept organisms?. J. Exp. Biol. 202: 3463–3467
Denny M.W., Daniel T.L. and Koehl M.A.R. (1985). Mechanical limits to size in wave-swept organisms. Ecol. Monogr. 51: 69–102
Denny M.W., Brown V., Carrington E., Kraemer G. and Miller A. (1989). Fracture mechanics and the survival of wave-swept macroalgae. J. Exp. Mar. Biol. Ecol. 127: 211–228
Denny M. and Gaylord B. (1996). Why the urchin lost its spines: hydrodynamic forces and survivorship in three echinoids. J. Exp. Biol. 199: 717–729
Denny M.W., Gaylord B.P. and Cowen E.A. (1997). J. Exp. Biol. 200: 3165–3183
Denny M., Gaylord B., Helmuth B. and Daniel T. (1998). The menace of momentum: dynamic forces on flexible organisms. Limnol. Oceanogr. 43: 955–968
Denny M.W. and Gaines S. (1999). Chance in Biology. Princeton University Press.
Denny M.W. and Blanchette C.A. (2000). J. Exp. Biol. 203: 2623–2639
Denny M.W. and Wethey D. (2000). Physical processes that generate patterns in marine communities. In: Bertness, M.D., Gaines, S.D. and Hay, M.E. (eds) Marine Community Ecology, pp 3–38. Sinauer, New York
Denny M.W., Miller L.P., Stokes M.D., Hunt L.J.H. and Helmuth B.S.T. (2003). Topographical amplification of wave-induced flow in the surf zone of rocky shores. Limnol. Oceanogr. 48: 1–8
Dudgeon S.R. and Johnson A.S. (1992). Thick vs. thin: thallus morphology and tissue mechanics influence differential drag and dislodgment of two co-dominant seaweeds. J. Exp. Mar. Biol. Ecol. 165: 23–43
Ebling A.W., Laur D.R. and Rowley R.J. (1985). Severe storm disturbance and reversal of community structure in a southern California kelp forest. Mar. Biol. 84: 287–294
Eckart C. (1951). Surface waves on water of variable depth. Scripps Institute of Oceanography, La Jolla, CA
Etter R.J. (1988). Asymmetrical developmental plasticity in an intertidal snail. Evolution 42: 322–334
Friedland M. and Denny M.W. (1995). Surviving hydrodynamic forces in a wave-swept environment: consequences of morphology in the feather boa kelp, Egregia menziesii (Turner). J. Exp. Mar. Biol. Ecol. 190: 109–133
Galvin C.J. (1972). Wave breaking in shallow water. In: Meyers, R.E. (eds) Waves on Beaches and Resulting Sediment Transport, pp. Academic Press, NY
Gaylord B. (1999). Detailing agents of physical disturbance: wave-induced velocities and accelerations on a rocky shore. J. Exp. Mar. Biol. Ecol. 239: 85–124
Gaylord B. (2000). Biological implications of surf-zone flow complexity. Limnol. Oceanogr. 45: 174–188
Gaylord B., Blanchette C.A. and Denny M.W. (1994). Mechanical consequences of size in wave-swept algae. Ecol. Monogr. 64: 287–313
Gaylord B. and Denny M. (1997). J. Exp. Biol. 200: 3141–3164
George R., Flick R.E. and Guza R.T. (1994). Observation of turbulence in the surf zone. J. Geophys. Res. 99(C1): 801–810
Gerard V.A. (1987). Hydrodynamic streamlining of Laminaria saccharina Lamour. in response to mechanical stress. J. Exp. Mar. Biol. Ecol. 107: 237–244
Graham M. (1997). Factors determining the upper limit of the giant kelp, Macrocystis pyrifera Agardhalong the Monterey peninsulacentral California, USA. J. Exp. Mar. Biol. Ecol. 218: 127–149
Grant W.D. and Madsen O.S. (1986). The continental shelf bottom boundary layer. Ann. Rev. Fluid Mech. 18: 265–305
Hibberd S. and Peregrine D.H. (1979). Surf and run-up on a beach: a uniform bore. J. Fluid Mech. 95: 323–345
Hoffman G.E., Buckley B.A., Place S.P. and Zippay M.L. (2002). Molecular chaperones in ectothermic marine animals: biochemical function and gene expression. Int. Comp. Biol. 42: 808–814
Johnson A.S. and Koehl M.A.R. (1994). Maintenance of dynamic strain similarity and environmental stress factor in different flow habitats: thallus allometry and material properties of giant kelp. J. Exp. Biol. 195: 381–410
Johnson L.E. (1994). Limnol. Oceanogr. 39: 1893–1902
Judge M.L. (1988). The effects of increased drag on Lottia gigantea (Sowerby 1834) foraging behavior. Funct. Ecol. 2: 363–369
Kawamata S. (1998). Effect of wave-induced oscillatory flow on grazing by a subtidal sea urchin Strongylocentrotus nudus (A. Agassiz). J. Exp. Mar. Biol. Ecol. 224: 31–48
Kinsman B. (1965). Wind Waves. Prentice-Hall, Englewood Cliffs, NJ
Koehl M.A.R. (1977). Effects of sea anemones on the flow forces they encounter. J. Exp. Biol. 69: 87–105
Koehl M.A.R. (1984). How do benthic organisms withstand moving water. Am. Zool. 24: 57–70
Koehl M.A.R. (1986). Seaweeds in moving water: forma and mechanical function. In: Givnish, T.J. (eds) On the Economy of Plant Form and Function, pp 603–634. Cambridge University Press, Cambridge
Koehl M.A.R. (1999). Ecological biomechanics of benthic organisms: life history, mechanical design and temporal patterns of mechanical stress. J. Exp. Biol. 202: 3469–3476
Koehl M.A.R. and Wainwright S.A. (1977). mechanical design of a giant kelp. Limnol. Oceanogr. 22: 1067–1071
Koehl M.A.R. and Alberte R.S. (1988). Flow, flapping and photosynthesis of Nereocystis luetkeana: a functional comparison of undulate and flat blade morphologies. Mar. Biol. 99: 435–444
Leigh E.G., Paine R.T., Quinn J.F. and Suchanek T.H. (1987). Wave energy and intertidal productivity. Proc. Natl. Acad. Sci. USA 84: 1314–1318
Lewis J.R. (1964). The Ecology of Rocky Shores. English Universities Press, London
Levitan D.R. (1995). The ecology of fertilization in free-spawning invertebrates. In: McEdwards, L.M. (eds) Ecology of Marine Invertebrate Larvae, pp. CRC Press, Boca Raton, FL
Longuet-Higgins M.S. (1952). On the statistical distribution of the heights of sea waves. J. Mar. Res. 11: 245–266
Longuet-Higgins M.S. (1980). On the distribution of heights of sea waves: some effects of nonlinearity and finite bandwidth. J. Geophys. Res. 85(C3): 1519–1523
Lowell R.B. (1985). Selection for increased safety factors of biological structures as environmental unpredictability increases. Science 228: 1009–1011
Lowell R.B. (1987). Safety factors of tropical versus temperate limpet shells: multiple selection pressures on a single structure. Evolution 41: 638–650
Lubchenco J. (1978). Plant diversity in a marine intertidal community: importance of herbivore food preference and algal competitive abilities. Am. Nat. 112: 23–39
Massel S.R. (1996). Ocean Surface Waves: Their Physics and Prediction. World Scientific, London
Massel S.R. and Done T.J. (1993). Effects of cyclone waves on massive coral assemblages on the Great Barrier Reef: meteorology, hydrodynamics and demography. Coral Reefs 12: 153–166
Menge B.A. (1976). Organization of the New England rocky intertidal community: role of predation, competition and environmental heterogeneity. Ecol. Monogr. 46: 355–393
Menge B.A. (1978). Predation intensity in a rocky intertidal community: relation between predator foraging activity and environmental harshness. Oecologia 34: 1–16
Munk W.H. (1949). The solitary wave theory and its application to surf problems. Ann. New York Acad. Sci. 51: 376–424
Newell R.C. (1979). Biology of Intertidal Organisms. Marine Ecological Surveys, Faversham, UK
Paine R.T. (1966). Food web complexity and species diversity. Am. Nat. 100: 65–75
Paine R.T. (1974). Intertidal community structure: experimental studies on the relationship between a dominant competitor and its principal predator. Oecologia 15: 93–120
Paine R.T. (1979). Disastercatastropheand local persistence of the sea palmPostelsia palmaeformis. Science 205: 685–687
Paine R.T. and Levin S.A. (1981). Intertidal landscapes: disturbance and the dynamics of pattern. Ecol. Monogr. 51: 145–178
Pearson G.A. and Brawley S.H. (1998). Sensing hydrodynamic conditions via carbon acquisition: control of gamete release in fucoid seaweeds. Ecology 79: 1725–1739
Rayleigh J.W.S. (1880). On the resultant of a large number of vibrations of the same pitch and arbitrary phase. Phil. Mag. 10: 73–78
Ricketts E.F., Calvin J., Hedgpeth J.W. and Phillips D.W. (1985). Between Pacific Tides. Stanford University Press, Stanford, CA
Schlichting H. (1979). Boundary-Layer Theory. McGraw-Hill, NY
Serrão E.A., Pearson G.A., Kautsky L. and Brawley S.H. (1996). Successful external fertilization in turbulent environments. Proc. Natl. Acad. Sci. USA 93: 5286–5290
Seymour R.J., Tegner M.J., Dayton P.K. and Parnell P.E. (1989). Storm wave induced mortality of the giant kelp, Macrocystis pyriferain southern California. Est. Coast. Shelf Sci. 28: 277–292
Shanks A.L. and Wright W.G. (1986). Adding teeth to wave action: the destructive effects of wave-borne rocks on intertidal organisms. Oecologia 69: 420–428
Shaughnessy F.J., DeWreede R.E. and Bell E.C. (1996). Consequences of morphology and tissue strength to blade survivorship of two closely related Rhodophyta species. Mar. Ecol. Progr. Ser. 136: 257–266
Somero G. (2002). Thermal physiology and vertical zonation of intertidal animals: Optimalimits and costs of living. Int. Comp. Biol. 42: 779–780
Sousa W.P. (1979). Disturbance in marine intertidal boulder fields: the nonequilibrium maintenance of species diversity. Ecology 60: 1225–1239
Stillman J. 2002. Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus Petrolisthes. Int. Comp. Biol.: 790–796.
Svendsen I.A. (1987). Analysis of surf zone turbulence. J. Geophys. Res. 92: 5115–5124
Thornton E.B. and Guza R.T. (1983). Transformation of wave height distribution. J. Geophys. Res. 88(C10): 5925–5938
Tomanek L. (2002). The heat-shock response: its variation, regulation and ecological importance in intertidal gastropods (genus Tegula). Int. Comp. Biol. 42: 797–807
Tomanek L. and Helmuth B. (2002). Physiological ecology of inertial organisms: a synergy of concepts. Int. Comp. Biol. 42: 771–775
Tricker R.A.R. (1964). Bores, Breakers, Waves and Wakes. Mills and Boon, London
Trussell G.C. (1997). Phenotypic plasticity in the foot size of an intertidal snail. Ecology 78: 1033–1048
(1984). Shore Protection Manual. US Government Printing Office, Washington, DC
Utter B.D. and Denny M.W. (1996). Wave-induced forces on the giant kelp Macrocystis pyrifera (Agardh): field test of a computational model. J. Exp. Biol. 199: 2645–2654
Van Dorn W.G. 1976. Set-up and run-up in shoaling breakers. Proc. 15th Int. Coastal Eng. Conf., pp. 738–751.
Van Dorn W.G. (1978). Breaking invariants in shoaling waves. J. Geophys. Res. 83: 2981–2988
Vogel S. (1994). Life in Moving Fluids. Princeton University Press, Princeton, NJ
Wing S.R. and Patterson M.R. (1993). Effects of wave-induced lightflecks in the intertidal zone on photosynthesis in the macroalgae Postelsia palmaeformis and Hedophylum sessile (Phaeophyceae). Mar. Biol. 116: 519–525
Witman J.D. (1987). Subtidal coexistence: storms, grazing, mutualismand the zonation of kelps and mussels. Ecol. Monogr. 57: 167–187
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Denny, M.W. Ocean waves, nearshore ecology, and natural selection. Aquat Ecol 40, 439–461 (2006). https://doi.org/10.1007/s10452-004-5409-8
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DOI: https://doi.org/10.1007/s10452-004-5409-8