Department of Biological Science

Biological Science Faculty Member

Personal Home Page

Professor
Ph.D., University of Delaware, 1989
Graduate Faculty Status

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.

Selected Publications:

2019 Levitan, D.R., R. Buchwalter and Y. Hao.  The Evolution of gametic compatibility and compatibility groups in the sea urchin Mesocentrotus franciscanus: an avenue for speciation in the sea.  Evolution 73:1428-1442 [PDF].

2019 Olsen, K., J. Moscoso and D.R. Levitan.  Somatic mutation is a function of clone size and depth in Orbicella reef-building corals. Biological Bulletin 236:1-12 [PDF].

2018 Betters, M. and D.R. Levitan. The role of compatibility in determining the costs and benefits of polyandry in the broadcast spawning sea urchin Lytechinus variegatus. Biological Bulletin 235:63-70 [PDF].

2018 Levitan, D.R. Do sperm really compete and do eggs ever have a choice? Adult distribution and gamete mixing influence sexual selection, sexual conflict and the evolution of gamete recognition proteins in the sea.  American Naturalist 191: 88-105 [PDF].

2017 Kosman, E.T., B. Hipp and D.R. Levitan.  Chemoattractant-mediated preference of non-self eggs in Ciona robustas sperm. Biological Bulletin 233:183-189 [PDF].

2017 Burgess, S.C., W H. Ryan, N.W. Blackstone, P.J Edmunds, M. O. Hoogenboom, D.R. Levitan, J.L Wulff.  Metabolic scaling in modular animals.  Invertebrate Biology 136:456-472, DOI: 10.1111/ivb.12199 [PDF].

2016  Prada, C., B. Hanna, A.F. Budd, C.M. Woodley, J. Schmutz, J. Grimwood, R. Iglesia-Prieto, J.M. Pandolfi, D. R. Levitan, K.G. Johnson, N. Knowlton, H. Kitano, M. DeGregorio and M. Medina. Empty niches after extinctions increase population size of modern corals.  Current Biology 26:1-5 [PDF].

2016 Campbell, A.L., D.R. Levitan, D.J. Hosken and C. Lewis.  Ocean acidification changes the male fitness landscape. Scientific Reports 6:31250 [PDF].

2015 Stapper, A.P., P. Beerli and D.R. Levitan.  Assortative mating drives linkage-disequilibrium between sperm and egg recognition proteins in the sea urchin Strongylocentrotus purpuratus. Molecular Biology and Evolution.  Advanced access msv010v2
[PDF].

2014 Levitan, D.R., W. Boudreau, J. Jara and N. Knowlton.  Long-term reproductive consequences of bleaching stress on the Caribbean corals of the Orbicella (formerly Montastraea) annularis species complex. Marine Ecology Progress Series 515:1-10 [PDF].

2014 Kosman, E.T. and D.R. Levitan.  Sperm competition and the evolution of gametic compatibility in externally fertilizing taxa.  Molecular Human Reproduction doi:1093/molehr/gau069 [PDF].

2014 Edmunds, P.J., X. Ponchon, D.R. Levitan, D.M. Yost, M. Belcaid, H. M. Putnam, R.D. Gates.  Long-term changes in Symbiodinium communities in Orbicella annularis in St. John, U.S. Virgin Islands.  Marine Ecology Progress Series 506:129-144 [PDF].

2014 Levitan, D.R., P.J. Edmunds, K.E. Levitan.  What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–84 mass mortality.   Oecologia 175:117-128 [PDF].

2013 Lessios, H.A., H.A. Lasker and D.R. Levitan.  The use of SCUBA to study the early life history of marine invertebrates. In Research and Discoveries: The Revolution of Science through Scuba, Smithsonian Contributions to Marine Sciences 39: 87-97 [PDF].

2012 Fogarty, N.D., M. Lowenberg, M.N. Ojima, N. Knowlton and D.R. Levitan. Asymmetric conspecific sperm precedence in relation to spawning times in the Montastraea annularis species complex (Cnidaria: Scleractinia).  Journal of Evolutionary Biology 25:2481-2488 [PDF].

2012  Levitan, D. R. Contemporary evolution of sea urchin gamete-recognition proteins: experimental evidence of density-dependent gamete performance predicts shifts in allele frequencies over time. Evolution 66:1722-1736 [PDF].

2012  Fogarty, N. D.; S. V. Vollmer, and D. R. Levitan. Weak prezygotic isolating mechanisms in threatened caribbean Acropora corals. PLoS One 7:e30486 [PDF].

2011  Levitan, D. R., N. D. Fogarty, J. Jara, K. E. Lotterhos, and N. Knowlton. Genetic, spatial, and temporal components of precise spawning synchrony in reef building corals of the Montastraea annularis species complex. Evolution 65:1254-1270 [PDF].

2011  Simon, T. N., and D. R. Levitan. Measuring fertilization success of broadcast-spawning marine invertebrates within seagrass meadows. Biological Bulletin 220:32-38 [PDF].

2010  Reuter, K. E., and D. R. Levitan. Influence of sperm and phytoplankton on spawning in the echinoid Lytechinus variegatus. Biological Bulletin 219:198-206 [PDF].

2010  terHorst, C. P., T. E. Miller, and D. R. Levitan. Evolution of prey in ecological time reduces the effect size of predators in experimental microcosms. Ecology 91:629-636 [PDF].

2010  Tomaiuolo, M., and D. R. Levitan. Modeling how reproductive ecology can drive protein diversification and result in linkage-disequilibrium between sperm and egg proteins. American Naturalist 176:12-25 [PDF].

2010  Levitan, D. R., and A. P. Stapper. Simultaneous positive and negative frequency dependent selection on sperm bindin, a gamete recognition protein in the sea urchin Strongylocentrotus purpuratus. Evolution 64:785-797 [PDF].

2010  Lotterhos, K., and D. R. Levitan. Gamete release and spawning behavior in broadcast spawning marine invertebrates. Pages 99-120 in J. Leonard, ed., The Evolution of Primary Sexual Characters. Oxford University Press, Oxford, New York.

2010  Levitan, D. R. Sexual selection in external fertilizers. Pages 365-378 in D. F. Westneat and C. W. Fox, eds., Evolutionary Behavioral Ecology. Oxford University Press, New York.

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, and D. R. Levitan. A theoretical investigation of sympatric evolution of temporal reproductive isolation as illustrated by marine broadcast spawners. Evolution 612584-2595 [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, and 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, and 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 Seas Biodiversity. Island Press, Washington, DC.

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 on molecular 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, Cambridge, UK.

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 157:626-636 [ 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 Naturalist 156:175-192 [ PDF ].

2000  Levitan D. R. Sperm velocity and endurance trade off each other and influence fertilization in the sea urchin Lytechinus variegatus. Proceedings of the Royal Society of London: Biological Sciences 267:531-534 [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, San Diego [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. Pages 159-164 in G. S. Jamieson and A. Campbell, eds., Proceedings of the North PacificSymposium on Invertebrate Stock Assessment and Management. Canadian Special Publication in Fisheries and Aquatic Science 125.

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 Science 62:937-946.

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., D. R. Levitan, 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., D. R. Levitan, 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 and Protocols. Wiley-Liss, New York

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, Boca Raton.

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 of Experimental 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-term monitoring 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 181:261-268 [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 Saint John, U.S. Virgin Islands. Journal of Experimental Marine Biology and Ecology119:167-178 [PDF].

1988  Levitan, D. R. Asynchronous spawning and aggregative behavior in the sea urchin Diadema antillarum Philippi. Pages 181-186 in R. D. Burke, ed., Echinoderm Biology. A.A. Balkema Press, Rotterdam [PDF].

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