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Professor;
Ph.D., University of Delaware, 1989

Research and Professional Interests:

I am interested in the ecology and evolution of marine invertebrates. My work examines the interactions between ecological processes, natural and sexual selection, and molecular evolution. For example, how does population density influence selection on gamete recognition proteins with cascading effects on reproductive isolation and speciation (Levitan and Ferrell 2006)? I enjoy integrating field and laboratory studies into a theoretical framework. I use a diverse array of tools to answer questions including field experiments on gamete fertilization and reproductive isolation, molecular studies of paternity, hybridization and protein evolution, phylogenetic analysis of trait evolution and theoretical explorations of sexual selection and gamete evolution. Click on the pictures below to see my ongoing projects.

Selected Publications:

2008 Levitan, D.R. Gamete traits influence the variance in reproductive success, the intensity of sexual selection, and the outcome of sexual conflict among congeneric sea urchins. Evolution 62:1305-1316 [PDF]

2007 Tomaiuolo, M., T.F. Hansen, D. R. Levitan. A theoretical investigation of sympatric evolution of temporal reproductive isolation as illustrated by marine broadcast spawners. Evolution 61:2584-2595 [PDF]

2007 Munguia, P., C. Mackie and D.R. Levitan. The influence of stage-dependent dispersal on the population dynamics of three amphipod species. Oecologia 153:533-541 [PDF]

2007 Levitan, D.R., C.P. terHorst, N.D. Fogarty. The risk of polyspermy in three congeneric sea urchins and its implications for gametic incompatibility and reproductive isolation. Evolution 61:2007-2014 [PDF]

2006 Levitan, D.R. and D.L. Ferrell. Selection on gamete recognition proteins depends on sex, density and genotype frequency. Science 312:267-269 [PDF] [On-line Supplement PDF]

2006 Levitan, D.R. Relationship between egg size and fertilization success in broadcast spawning marine invertebrates. Integrative and Comparative Biology 46:298-311 [PDF]

2005 Zigler, K.S., M.A. McCartney, D.R. Levitan, H.A. Lessios. Sea urchin bindin divergence predicts gamete compatibility. Evolution 59:2399-2404 [PDF]

2005 Levitan, D.R. The distribution of male and female reproductive success in a broadcast spawning marine invertebrate. Integrative and Comparative Biology 45:848-855 [PDF]

2005 Levitan, D.R. Sex specific spawning behavior and its consequences in an external fertilizer. American Naturalist 165:682-694 [PDF]

2005 Levitan, D.R. and T.M. McGovern. The Allee effect in the Sea. Pages 47-57 in (E.A. Norse and L.B. Crowder eds) Marine Conservation Biology: The Science of Maintaining the Sea’s Biodiversity. Island Press

2004 McCartney, M.A., K. Brayer and D.R. Levitan. Polymorphic microsatellite lo ci from the red urchin, Strongylocentrotus franciscanus, with comments on heterozyogote deficit. Molecular Ecology Notes 4: 226-228 [PDF]

2004 Levitan, D.R. Density-dependent Sexual Selection in External Fertilizers: Variances in Male and Female Reproductive Success Along the Continuum from Sperm Limitation to Sexual Conflict in the Sea Urchin Strongylocentrotus franciscanus.American Naturalist 164:298-309 [PDF]

2004 Fukami, H., A.F. Budd, D.R. Levitan, J. Jara, R. Kersanach and N. Knowlton. Geographical differences in species boundaries among members of the Montastraea annularis complex based onmolecular and morphological markers. Evolution 58:324-337.[PDF]

2004 Levitan, D.R., H. Fukami, J. Jara, D. Kline, T.A. McGovern, K.M. McGhee, C.A. Swanson and N. Knowlton. Mechanisms of reproductive isolation among sympatric broadcast-spawning corals of the Montastraea annularis complex. Evolution 58: 308-323.[PDF]

2002 Levitan, D.R. The relationship between conspecific fertilization success and reproductive isolation among three congeneric sea urchins. Evolution 56:1599-1609. [PDF]

2002 Levitan, D.R. Density-dependent selection on gamete traits: Are traits associated with fertilization related to sperm availability in three congeneric sea urchins? Ecology 83:464-479. [ PDF ]

2001 Levitan, D.R. and S.D. Irvine. Fertilization selection on egg and jelly-coat size in the sand dollar Dendraster excentricus. Evolution 55: 2479-2483. [ PDF ]

2001 Petersen, C. and D.R. Levitan. The Allee effect: a barrier to repopulation of exploited species. Pages 281-300, in (J.D. Reynolds, G.M. Mace, K.H. Redford and J.G. Robinson eds) Conservation of Exploited Species. Cambridge University Press.

2001 Farley, G.S. and D.R. Levitan. The role of jelly coats in sperm-egg encounters, fertilization success, and selection on egg size in echinoids. American Naturalist 157:626-236. [ PDF ]

2000 Levitan, D.R. Optimal egg size in marine invertebrates: theory and phylogenetic analysis of the critical relationship between egg size and development time in echinoids. American Naturalist156:175-192. [ PDF ]

2000 Levitan D.R. Sperm velocity and endurance trade-off and influence fertilization in the sea urchin Lytechinus variegatus. Proceedings of the Royal Society of London: Biological Sciences267:531-534. [ PDF ]

1998 Levitan, D.R. Does Bateman's Principle apply to broadcast-spawning organisms? Egg traits influence in situ fertilization rates among congeneric sea urchins. Evolution 52:1043-1056. [ PDF ]

1998 Levitan, D.R. Sperm limitation, sperm competition and sexual selection in external fertilizers. Pages 173-215, in (T. Birkhead and A. Moller eds) Sperm Competition and Sexual Selection. AcademicPress. [ PDF ]

1998 Levitan, D.R. and M.A. Sewell. Male abundance and fertilization success in free-spawning marine invertebrates: Review of the evidence and fisheries implications. In Proceedings of the North PacificSymposium on Invertebrate Stock Assessment and Management. Edited by G.S. Jamieson and A. Campbell. Canadian Special Publication in Fisheries and Aquatic Science 125:159-164.

1998 Edmunds, P.J. R.B. Aronson, D.W. Swanson, D.R. Levitan, and W.F. Precht. Photographic versus visual census techniques for the quantification of juvenile corals. Bulletin of Marine Science62:937-946.

1997 Grosberg, R.K., M.W. Hart and D.R. Levitan. Is allorecognition specificity in Hydractinia symbiologicarpus controlled by a single gene? Genetics 145:857-860. [PDF]

1996 Grosberg, R.K., Levitan, D.R. and B.B. Cameron. Evolutionary Genetics of allorecognition in the colonial hydroid Hyractinia symbiologicarpus. Evolution 50:2221-2240. [PDF]

1996 Levitan, D.R. Effects of gamete traits on fertilization in the sea and the evolution of sexual dimorphism. Nature 382:153-155. [PDF]

1996 Levitan, D.R. Predicting optimal and unique egg sizes in free-spawning marine invertebrates. American Naturalist 148:174-188. [PDF]

1996 Grosberg, R.K., Levitan, D.R. and B.B. Cameron. Characterization of genetic structure and genealogies using RAPD-PCR markers: a random primer for the novice and nervous. Pages 67-100 in(J. Ferraris and S. Palumbi eds.) Molecular Zoology: Advances, Strategies & Protocols. WileyPress.

1995 Levitan, D.R. The ecology of fertilization in free-spawning invertebrates. Pages 123-156 in (L. McEdward ed.) Ecology of Marine Invertebrate Larvae. CRC Press.

1995 Levitan, D.R. and C.M. Young. Reproductive success in large populations: empirical measures and theoretical predictions of fertilization in the sea biscuit Clypeaster rosaceus. Journal ofExperimental Marine Biology and Ecology 190:221-241. [PDF]

1995 Levitan, D.R. and C. Petersen. Sperm limitation in the sea. Trends in Ecology and Evolution. 10:228-231. [PDF]

1994 Aronson, R.B., P.J Edmunds, W.F. Precht, D.W. Swanson, and D.R. Levitan. Large-scale, long-termmonitoring of Caribbean coral reefs: simple, quick inexpensive techniques. Atoll Research Bulletin 42:11-19.

1993 Levitan, D.R. and R.K. Grosberg. The analysis of paternity and maternity in the marine hydrozoan Hydractinia symbiolongicarpus using randomly amplified polymorphic (RAPD) markers. Molecular Ecology 2:315-326.

1993 Levitan, D.R. The importance of sperm limitation to the evolution of egg size in marine invertebrates. American Naturalist 141:517-536. [PDF]

1992 Sewell, M.A. and D.R. Levitan. Fertilization success in a natural spawning of the dendrochirote sea cucumber Cucumaria miniata. Bulletin of Marine Science 51:161-166.

1992 Levitan, D.R. Community structure in times past: influence of human fishing pressure on algal- urchin interactions. Ecology 73:1597-1605. [ PDF ]

1992 Grosberg, R.K. and D.R. Levitan. For adults only? Supply side ecology and the history of larval ecology. Trends in Ecology and Evolution 7:130-133.

1992 Grosberg, R.K. and D.R. Levitan. Reply to Dayton. Trends in Ecology and Evolution 7:392- 393.1992 Levitan, D.R., M.A. Sewell, and F.-S. Chia. How distribution and abundance influencefertilization success in the sea urchin Strongylocentrotus franciscanus. Ecology 73:248-254. [PDF]

1991 Levitan, D.R. Skeletal changes in the test and jaws of the sea urchin Diadema antillarum when food-limited. Marine Biology 106:431-435. [PDF]

1991 Levitan, D.R. Influence of body size and population density on fertilization success and reproductive output in a free-spawning invertebrate. Biological Bulletin 181261-268. [PDF]

1991 Levitan, D.R., M.A. Sewell, and F.-S. Chia. Kinetics of fertilization in the sea urchin Strongylocentrotus franciscanus: interaction of gamete dilution, age, and contact time. BiologicalBulletin 181:371-378. [PDF]

1990 Karlson, R.H. and D.R. Levitan. Recruitment-limitation in open populations of Diadema antillarum: an evaluation. Oecologia 82:40-44. [PDF]

1990 Levitan, D.R. Population consequences of indeterminate growth in the tropical sea urchin Diadema antillarum: A sea urchin's response to mass-mortalities. Park Science 10:8.

1989 Levitan, D.R. and S.J. Genovese. Substratum-dependent predator-prey dynamics: Patch reefs as refuges from gastropod predation. Journal of Experimental Marine Biology and Ecology 130:111-118. [PDF]

1989 Levitan, D.R. Density-dependent size regulation in Diadema antillarum: effects on fecundity and survivorship. Ecology 70:1414-1424. [PDF]

1988 Ketcham, R., D. Levitan, A. Shenk, and R. Eisenberg. Do interactions of cellular slime molds species regulate their densities in soil? Ecology 69:193-199. [PDF]

1988 Levitan, D.R. Density-dependent size regulation and negative growth in the sea urchin Diadema antillarum Philippi. Oecologia 76:627-629.[PDF]

1988 Levitan, D.R. Algal-urchin biomass responses following the mass mortality of the sea urchin Diadema antillarum Philippi at SaintJohn, U.S. Virgin Islands. Journal of Experimental MarineBiology and Ecology 119:167-178.[PDF]

1988 Levitan, D.R. Asynchronous spawning and aggregative behavior in the sea urchin Diadema antillarum Philippi. In (R.D. Burke et al. eds.) Echinoderm Biology, pp 181-186, A.A. BalkemaPress, Rotterdam. [PDF]

Graduate Students:

Adreani, Maria (Mia)
Fogarty, Nicole (Nikki) D
Hollensead, Lisa
Lotterhos, Kathleen
Plata Stapper, Andres
terHorst, Casey

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