Mark Denny Publications

Journal Articles (chronological)

Miller, L.P., B.J. Allen, F.A. King, D.R. Chilin, V.M. Reynoso and M.W. Denny (2015) Warm microhabitats drive both increased respiration and growth rates of intertidal consumers. Marine Ecology Progress Series 522: 127-143 doi:

Staaf, D.J., W.F. Gilly and M.W. Denny (2014). Aperture effects in squid jet propulsion. Journal of Experimental Biology 217(9): 1588-1600 doi:

Denny, M. (2014). Buzz Holling and the Functional Response. Bulletin of the Ecological Society of America 95(3): 200-203 doi:

Cole, A. and M. Denny (2014). United we fail: group versus individual strength in the California sea mussel, Mytilus californianus. The Biological Bulletin 227(1): 61-67

Denny, M. (2014). Buzz Holling and the Functional Response. Bulletin of the Ecological Society of America 95(3): 200-203 doi:

Staaf, D.J., W.F. Gilly and M.W. Denny (2014). Aperture effects in squid jet propulsion. Journal of Experimental Biology 217(9): 1588-1600 doi:

Denny, M., K. Mach, S. Tepler and P. Martone (2013). Indefatigable: an erect coralline alga is highly resistant to fatigue. Journal of Experimental Biology 216(20): 3772-3780 doi:

Rosman, J.H., M.W. Denny, R.B. Zeller, S.G. Monismith and J.R. Koseff (2013). Interaction of waves and currents with kelp forests (Macrocystis pyrifera): Insights from a dynamically scaled laboratory model. Limnology and Oceanography 58(3): 790-802

Denny, M. and L. Benedetti-Cecchi (2012). Scaling up in ecology: mechanistic approaches. Annual Review of Ecology, Evolution, and Systematics 43(1): 1-22 doi:

Booth, J.A.T., E.E. McPhee-Shaw, P. Chua, E. Kingsley, M. Denny, R. Phillips, S.J. Bograd, L.D. Zeidberg and W.F. Gilly(2012). Natural intrusions of hypoxic, low pH water into nearshore marine environments on the California coast. Continental Shelf Research 45(0): 108-115 doi:

Denny, M. (2012). Biophysics, bioenergetics and mechanistic approaches to ecology. The Journal of Experimental Biology 215(6): 871 doi:

Moláček, J., M. Denny and J.W.M. Bush (2012). The fine art of surfacing: Its efficacy in broadcast spawning. Journal of Theoretical Biology 294(0): 40-47 doi:

Denny, M.W. and W.W. Dowd (2012). Biophysics, environmental stochasticity, and the evolution of thermal safety margins in intertidal limpets. The Journal of Experimental Biology 215(6): 934-947. doi:

Tepler, S., K. Mach and M. Denny (2011). Preference versus performance: body temperature of the intertidal snail Chlorostoma funebralis. The Biological Bulletin 220(2): 107-117

Miller, L.P. and M.W. Denny (2011). Importance of behavior and morphological traits for controlling body temperature in littorinid snails. Biological Bulletin 220(3): 209-223.

Denny, M.W., W.W. Dowd, L. Bilir and K.J. Mach (2011). Spreading the risk: Small-scale body temperature variation among intertidal organisms and its implications for species persistence. Journal of Experimental Marine Biology and Ecology 400(1-2): 175-190. doi:

Mach, K.J., S.K. Tepler, A.V. Staaf, J.C. Bohnhoff and M.W. Denny (2011). Failure by fatigue in the field: a model of fatigue breakage for the macroalga Mazzaella, with validation. The Journal of Experimental Biology 214(9): 1571-1585. doi:

Denny, M.W. and B. Gaylord (2010). Marine Ecomechanics. The Annual Review of Marine Science 2: 89-114. doi:

Helmuth, B., B.R. Broitman, L. Yamane, S.E. Gilman, K. Mach, K.A.S. Mislan and M.W. Denny (2010). Organismal climatology: analyzing environmental variability at scales relevant to physiological stress. Journal of Experimental Biology 213: 995-1003. doi:

Rosman, J.H., S.G. Monismith, M.W. Denny and J.R. Koseff (2010). Currents and turbulence within a kelp forest (Macrocystis pyrifera): insights from a dynamically scaled laboratory model. Limnology and Oceanography 55(3): 1145-1158. doi:

Miklasz, K.A. and M.W. Denny (2010). Diatom sinking speeds: Improved predictions and insight from a modified Stokes’ law. Limnology and Oceanography 55(6): 2513-2525. doi:

Denny, M.W. and B. Helmuth (2009). Confronting the physiological bottleneck: a challenge from ecomechanics. Integrative and Comparative Biology 49(3): 197-201. doi:

Denny, M.W., L.J.H. Hunt, L.P. Miller and C.D.G. Harley (2009). On the prediction of extreme ecological events. Ecological Monographs 79(3): 397-421. doi:

Denny, M.W., B.L. Lockwood and G.N. Somero (2009). Can the giant snake predict palaeoclimate? Nature 460(7255): E3-E4. doi:

Harley, C.D.G., M.W. Denny, K.J. Mach and L.P. Miller (2009). Thermal stress and morphological adaptations in limpets. Functional Ecology 23: 292-301. doi:

Martone, P.T., J.M. Estevez, F. Lu, K. Ruel, M.W. Denny, C. Somerville and J. Ralph (2009). Discovery of lignin in seaweed reveals convergent evolution of cell-wall structure. Current Biology 19: 169-175. doi:

Miller, L.P., C.D.G. Harley and M.W. Denny (2009). The role of temperature and desiccation stress in limiting the local-scale distribution of the owl limpet, Lottia giganteaFunctional Ecology 23(4): 756-767. doi:

Denny, M.W. (2008). Limits to running speed in dogs, horses and humans. Journal of Experimental Biology 211: 3836-3849. doi:

Hunt, L.J.H. and M.W. Denny (2008). Desiccation protection and disruption: a trade-off for an intertidal marine alga. Journal of Phycology 44: 1164-1170. doi:

Martone, P.T. and M.W. Denny (2008). To bend a coralline: effect of joint morphology on flexibility and stress amplification in an articulated calcified seaweed. Journal of Experimental Biology 211: 3421-3432. doi:

Martone, P.T. and M.W. Denny (2008). To break a coralline: mechanical constraints on the size and survival of a wave-swept seaweed. Journal of Experimental Biology 211: 3433-3441. doi:

O’Donnell, M.J. and M.W. Denny (2008). Hydrodynamic forces and surface topography: Centimeter-scale spatial variation in wave forces. Limnology and Oceanography 53(2): 579-588. doi:

Mach, K.J., D.V. Nelson and M.W. Denny (2007). Techniques for predicting the lifetimes of wave-swept macroalgae: a primer on fracture mechanics and crack growth. Journal of Experimental Biology 210: 2213-2230. doi:

Mach, K.J., B.B. Hale, M.W. Denny and D.V. Nelson (2007). Death by small forces: a fracture and fatigue analysis of wave-swept macroalgae. Journal of Experimental Biology 210: 2231-2243. doi:  

Denny, M.W. (2006). Ocean waves, nearshore ecology, and natural selection. Aquatic Ecology 40: 439-461. doi:  

Denny, M.W. and C.D.G. Harley (2006). Hot limpets: predicting body temperature in a conductance-mediated thermal system. Journal of Experimental Biology 209: 2409-2419. doi:

Denny, M.W., L.P. Miller and C.D.G. Harley (2006). Thermal stress on intertidal limpets: long-term hindcasts and lethal limits. Journal of Experimental Biology 209: 2420-2431. doi:

Denny, M. and L. Miller (2006). Jet propulsion in the cold: mechanics of swimming in the Antarctic scallop Adamussium colbeckiJournal of Experimental Biology 209: 4503-4514. doi:

Kitzes, J.A. and M.W. Denny (2005). Red algae respond to waves: Morphological and mechanical variation in Mastocarpus papillatus along a gradient of force. Biological Bulletin 208(2): 114-119.

Denny, M.W., B. Helmuth, G. Leonard, C.D.G. Harley, L.J.H. Hunt and E.K. Nelson (2004). Quantifying scale in ecology: lessons from a wave-swept shore. Ecological Monographs 74(3): 513-532. doi:

Li, N.K. and M.W. Denny (2004). Limits to phenotypic plasticity: flow effects on barnacle feeding appendages. Biological Bulletin206: 121-124.

Denny, M. W. and B. Hale. (2003). Cyberkelp: an integrative approach to the modeling of flexible organisms. Phil. Trans. Roy. SocLondon B v358 no. 1437: 1535-1542 doi:

Denny, M.W., L.P. Miller, M.D. Stokes, L.J.H. Hunt and B.S.T. Helmuth (2003). Extreme water velocities: Topographical amplification of wave-induced flow in the surf zone of rocky shores. Limnology and Oceanography 48(1): 1-8.  

Gaylord, B., M.W. Denny and M.A.R. Koehl (2003). Modulation of wave forces on kelp canopies by alongshore currents. Limnology and Oceanography 48(2): 860-871.

Helmuth, B. and M.W. Denny (2003). Predicting wave exposure in the rocky intertidal zone: Do bigger waves always lead to larger forces? Limnology and Oceanography 48(3): 1338-1345. doi:  

Denny, M. W., E. K. Nelson, and K. S. Mead. (2002) Revised estimates of the effects of turbulence on fertilization in the purple sea urchin, Strongylocentrotus purpuratus. Biol. Bull. 203: 275-277

Denny, M. W. and B. Gaylord. (2002) The mechanics of wave-swept algae. J. Exp. Biol. 205: 1355-1362.

Denny, M. and L. Roberson. (2002) Blade motion and the nutrient flux to the kelp Eisenia arborea. Biol. Bull. 203: 1-13

Denny, M. W. (2001). The rewards of chance. Nat. History 110(4):72-77.

Denny, M.W. (2000) Limits to optimization: Fluid dynamics, adhesive strength, and the evolution of shape in limpet shells. J. Exp. Biol. 203: 2603-2622.

Denny, M.W. and C. A. Blanchette. (2000) Hydrodynamics, shell shape, behavior, and survivorship in the owl limpet, Lottia gigantea. J. Exp. Biol. 203: 2623-2639.

Gaylord, B., B. B. Hale, and M. W. Denny. (2001) Consequences of transient fluid forces for compliant benthic organisms. J. Exp. Biol. 204: 1347-1360.

Denny, M.W. (1999) Are there mechanical limits to size in wave-swept organisms? J. Exp. Biol. 202: 3463-3467.

Abelson, A. and M.W. Denny. (1998) The roles of hydrodynamic forces in the settlement of marine propagules. Ann Rev. Syst. Ecology 28: 317-339.

Denny, M.W. and R.T. Paine. (1998) Celestial mechanics, sea-level changes, and intertidal ecology. Biol. Bull. 194: 108-115.

Denny, M.W., B. P. Gaylord, B. Helmuth, and T.L. Daniel. (1998) The menace of momentum: Dynamic forces on flexible organisms. Limnol. Oceanogr. 43: 955-968

Cooper, S. and M.W. Denny. (1997) A conjecture on the relationship of bacterial shape to motility in rod-shaped bacteria. FEMS Microbiology Letters 148(2): 227-231

Denny, M.W., B.P. Gaylord, and E. Cowen. (1997) Flow and flexibility II: The roles of size and shape in determining wave forces on the bull kelp, Nereocystis luetkeana. J. Exp. Biol. 200: 3165-3183.

Gaylord, B. P. and M.W. Denny (1997) Flow and Flexibility I: Effects of size, shape, and stiffness in determining wave forces on stipitate kelps, Eisenia arborea and Pterygophora californica. J. Exp. Biol. 200: 3141-3164

Huryn, A.D. and M.W. Denny (1997). A biomechanical hypothesis explaining upstream movements by the freshwater snail Elimia. Functional Ecology 11(4): 472-483 doi:

Denny, M.W. and B. Gaylord. (1996) Why the urchin lost its spines: hydrodynamics and survivorship in three echinoid species. J. Exp. Biol. 199: 717-729

Leichter, J.J., S.R. Wing, S.L. Miller and M.W. Denny (1996). Pulsed delivery of subthermocline water to Conch Reef (Florida Keys) by internal tidal bores. Limnology and Oceanography 41(7): 1490-150

Utter, B.D. and M.W. Denny. (1996) Wave-induced forces on the giant kelp, Macrocystis pyrifera: Field test of a computational model. J. Exp. Biol. 199: 2645-2654

Denny, M.W. (1995). Predicting physical disturbance: mechanistic approaches to the study of survivorship on wave swept shores. Ecological Monographs 65(4): 371-418 doi:

Denny, M.W. (1995) Size and survival in the surf zone. American Scientist 83: 166-173

Friedland, M.T. and M.W. Denny (1995). Surviving hydrodynamic forces in a wave-swept environment: Consequences of morphology in the feather boa kelp, Egregia menziesii (Turner). Journal of Experimental Marine Biology and Ecology 190(1): 109-133 doi:

Mead, K. and M.W. Denny. (1995) Effects of turbulence on fertilization in the purple sea urchin, Strongylocentrotus purpuratus. Biol. Bull. 188: 46-56

Bell, E.C. and M.W. Denny (1994). Quantifying “wave exposure”: a simple device for recording maximum velocity and results of its use at several field sites. Journal of Experimental Marine Biology and Ecology 181(1): 9-29 doi:  

Denny, M.W. (1994) Extreme drag forces and the survival of wind- and water-swept organisms. J. Exp. Biol. 194: 97-115

Gaylord, B, C. Blanchette and M.W. Denny. (1994) Mechanical consequences of size in wave-swept algae. Ecological Monographs64: 287-313

Gosline, J.M., C.C. Pollack, P.A. Guerette, A. Cheng, M.E. DeMont and M.W. Denny. (1994) Elastomeric network models for the frame and viscid silks from the orb web of the spider Araneus diadematus. Amer. Chem Soc. Symp. Series 544: 328-341

Denny, M.W. (1993) Disturbance, natural selection, and the prediction of maximal wave-induced forces. Fluid Dynamics in Biology. Contemporary Mathematics 141: 65-90

Gaines, S.D. and M.W. Denny (1993) The largest, smallest, highest, lowest, longest and shortest: extremes in ecology. Ecology 74: 1677-1692

Wing, S.R., J.J. Leichter and M.W. Denny. (1993) A dynamic model for wave-induced light fluctuations in a kelp forest. Limnol. Oceanogr. 38: 396-407

Denny, M.W., J. Dairiki and S. Destefano. (1992) Biological consequences of topography on wave- swept rocky shores: I. enhancement of external fertilization. Biol. Bull. 183: 220-232

Denny, M.W. (1991) Biology, natural selection and the prediction of maximal wave-induced forces. S. Afr. J. Mar. Sci. 10: 353-363

Holbrook, M.N., M.W. Denny and M.A.R. Koehl (1991). Intertidal “trees”: consequences of aggregation on the mechanical and photosynthetic properties of sea-palms Postelsia palmaeformis Ruprecht. Journal of Experimental Marine Biology and Ecology146(1): 39-67 doi:

Denny, M.W. (1990) Terrestrial versus aquatic biology: the medium and its message. Amer. Zool. 30: 111-121

Denny, M.W. and S. Gaines. (1990) On the prediction of maximum intertidal wave forces. Limnol. Oceanogr. 35: 1-15

Denny, M.W. (1989) Invertebrate mucous secretions: Functional alternatives to vertebrate paradigms. Symp. Soc. Exp. Biol. 43: 337-366

Denny, M.W. (1989) A limpet shell 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., V. Brown, E. Carrington, G. Kraemer and A. Miller (1989). Fracture mechanics and the survival of wave-swept macroalgae. Journal of Experimental Marine Biology and Ecology 127(3): 211-228 doi:

Denny, M.W. and J.A.C. Humphrey. (1989) Fluid mechanical constraints on biotic functions. Mechanical Engineering 111:49-53.

Denny, M.W. and M.F. Shibata. (1989) Consequences of surf-zone turbulence for settlement and external fertilization. Amer. Nat.134: 859-889.

Hahn, T. and M. Denny (1989). Tenacity-mediated selective predation by oystercatchers on intertidal limpets and its role in maintaining habitat partitioning by Collisella scabra and Lottia digitalis. Marine Ecology Progress Series 53(1): 1-10

Humphrey, J.A.C. and M.W. Denny. (1989) Ballooning spiders. Mechanical Engineering 111(11): 84-87.

Denny, M.W. (1987). Lift as a mechanism of patch initiation in mussel beds. Journal of Experimental Marine Biology and Ecology113(3): 231-245 doi:

Denny, M.W. (1987). Life in the maelstrom: The biomechanics of wave-swept rocky shores. Trends in Ecology & Evolution 2(3): 61-66 doi:

Gosline, J.M., E.M. Demont and M.W. Denny (1986) The structure and properties of spider silk. Endeavor 10(1): 37-44. (translated and reprinted in Natuur en Techniek, Holland)

Denny, M.W. (1985) Wave forces on intertidal organisms: a case study. Limnol. Oceanogr. 30: 1171-1187.

Denny, M.W., T. Daniel and M.A.R. Koehl (1985) Mechanical limits to size in wave-swept organisms. Ecol. Monogr. 55: 69-102

Denny, M.W. (1984). Mechanical properties of pedal mucus and their consequences for gastropod structure and performance. American Zoologist 24: 23-36 doi:

Gosline, J.M., M.W. Denny and M.E. DeMont (1984). Spider silk as rubber. Nature 309(5968): 551-552 doi:

Denny, M.W. (1983) A simple device for measuring the maximum force imposed on intertidal organisms. Limnol. Oceanogr. 28(6):1269-1274.

Denny, M.W. (1982) Forces on intertidal organisms: design and application of a wave-force telemetry system. Limnol. Oceanogr. 27:178-183.

Denny, M.W. (1981) A quantitative model for the adhesive locomotion of the terrestrial slug, Ariolimax columbianus. J. Exp. Biol.91:195-217.

Denny, M.W. and J.M. Gosline (1980) The physical properties of the pedal mucus of the terrestrial slug, Ariolimax columbianus. J. Exp. Biol. 88:375-93.

Denny, M.W. (1980) The silks – their properties and functions. Symposia of the Soc. for Exp. Biol. 34:247-72.

Denny, M. (1980). Locomotion: the cost of gastropod crawling. Science 208(4449): 1288-1290 doi:

Denny, M. (1980). The role of gastropod pedal mucus in locomotion. Nature 285(5761): 160-161 doi:

Denny, M.W. (1976) The physical properties of spiders’ silk and their role in the design of orb-webs. Journal of Experimental Biology65:483- 506.

Book Chapters

Denny, M.W. and D.Wethey. (2001) Physical processes that generate patterns in marine communities. Chapter 1 in M. Bertness, M. Hay, and S. Gaines (eds.) Marine Community Ecology, Sinauer Press, N.Y.

Denny, M.W. (1994) Roles of hydrodynamics in the study of life on wave-swept shores. pp. 169-204 in P.C. Wainwright and S.M. Reilly, editors. Ecomorphology: Integrative Organismal Biology. University of Chicago Press.

Gosline, J.M., R.E. Shadwick, M.E. Demont, and M.W. Denny. (1988) Non-Gaussian elastic properties in biopolymer networks. in Biological and Synthetic Polymer Networks, (O. Kramer, ed.), Plenum Press. pp. 57-77.

Denny, M.W. (1985) Water motion, pp. 7-29 in Handbook of Phycological Methods Vol. IV Ecology, M. Littler (ed.) Cambridge Univ. Press

Denny, M.W. (1983) “Molecular Biomechanics of Molluscan Mucous Secretions” in Biology of Mollusca, Vol. l Metabolic Biochemistry and Molecular Biomechanics. K. Wilbur, K. Simkiss, and P. Hochachka (Ed.) Academic Press, N.Y. pp. 431- 465.

Book Reviews

Denny, M. (1983) Structural Biomaterials by J.F.V. Vincent, Quarterly Review of Biology 58(2):310- 311.

Denny, M. (1985) Animal Mechanics by R. McN. Alexander. American Scientist, September issue

Denny, M.W. (1993) Plant Biomechanics by K. Niklas, Quarterly Review of Biology 68(4): 597.

Denny, M.W. (1995) Life In Moving Fluids by S. Vogel, Bulletin of Mathematical Biology 57: 949-951