David Bushek
Director & Professor
(856) 391-7653
Mailing Address:
  Haskin Shellfish Research Laboratory
  6959 Miller Avenue
  Port Norris, NJ  08349 USA


Ph.D. 1994 Rutgers, The State University of New Jersey
M.S. 1987 University of Houston
B.S. 1984 The Ohio State University

Research Interests

The predominant thrust of my research concerns host-parasite interactions in bivalve molluscs. A second focus aims to enhance our understanding of the ecological impact of bivalves, particularly the eastern oyster, at the population, community and ecosystem levels.

Current Research Projects

• Development of an Interactive Molluscan Shellfish Health Database for the East Coast for shellfish resource and aquaculture management. (NOAA SK, USDA APHIS VS, NJSG)

• Development of a Hatchery Certification Program to enhance biosecurity of interstate commerce of shellfish seed. (NJSG, USDA APHIS VS)

• DELSI = Delaware Estuary Living Shoreline Initiative (PDE, TNC, USFWS)

• Project PORTS (Promoting Oyster Restoration Through Schools).

• Delaware Bay New Jersey Oyster Stock Assessment and Long-term Monitoring of Disease and Mortality.

Selected Publications

Behringer, D. C., C. L. Wood, M. Krkosek, & D. Bushek. (2020). Disease in fisheries and aquaculture. In D. C. Behringer, B. R. Silliman, & K. D. Lafferty (Eds.), Marine Disease Ecology (pp. 183-212). New York, NY: Oxford University Press. 
Retrieved from

Yeh, H., S. A. Skubel, H. Patel, D. C. Shi, D. Bushek, & M. L. Chikindas. (2020). From farm to fingers: An exploration of probiotics for oysters, from production to human consumption. (2020). Probiotics and Antimicrobial Proteins Online First. DOI: 10.1007/s12602-019-09629-3 -or- Full article

Morris, R. L., D. M. Bilkovic, M. K. Boswell, D. Bushek, J. Cebrian, J. Goff, K. M. Kibler, M. K. La Peyre, G. McClenachan, J. Moody, P. Sacks, J. P. Shinn, E. L. Sparks, N. A. Temple, L. J. Walters, B. M. Webb, & S. E. Swearer. (2019). The application of oyster reefs in shoreline protection: Are we over‐engineering for an ecosystem engineer? Journal of Applied Ecology, 56 (7), 1703-1711. DOI: 10.1111/1365-2664.13390

Ben-Horin, T., C. A. Burge, D. Bushek, M. L. Groner, D. A. Proestou, L. I. Huey, G. Bidegain, & R. B. Carnegie. (2018). Intensive oyster aquaculture can reduce disease impacts on sympatric wild oysters. Aquaculture Environment Interactions, 10, 557–567. DOI: 10.3354/aei00290

Morson, J. M., D. M. Munroe, K. Ashton-Alcox, E. N. Powell, D. Bushek, & J. Gius. (2018). Density-dependent capture efficiency of a survey dredge and its influence on the stock assessment of eastern oysters (Crassostrea virginica) in Delaware Bay. Fisheries Research, 205, 115-121. DOI: 10.1016/j.fishres.2018.04.012

Paterno, J., L. Calvo, R. Jordan, & D. Bushek. (2017). Activity: One Fish, Two Fish-Assessing the Habitat Value of Restored Oyster Reefs: Using Scientific Research in the Classroom. Current: The Journal of Marine Education, 31 (1), 2-10.

Munroe, D. M., D. Bushek, P. Woodruff, & L. Calvo. (2017). Intertidal rack-and-bag oyster farms have limited interaction with horseshoe crab activity in New Jersey, USA. Aquaculture Environment Interactions, 9, 205-211. DOI: 10.3354/aei00227

Munroe. D. M., S. Borsetti, K. Ashton-Alcox, & D. Bushek. (2017). Early post-settlement growth in wild eastern oyster (Crassostrea virginica Gemlin 1791) populations. Estuaries and Coasts, 40 (3), 880-888. DOI: 10.1007/s12237-016-0185-y

Bidegain, G., E. N. Powell, J. M. Klinck, E. E. Hofmann, T. Ben-Horin, D. Bushek, S. E. Ford, D. M. Munroe, & X. Guo. (2017). Modeling the transmission of Perkinsus marinus in the eastern oyster Crassostrea virginica. Fisheries Research, 186 (1), 82–93. DOI: 10.1016/j.fishres.2016.08.006

Luckenbach, M. W., J. N. Kraeuter, & D. Bushek. (2016). Assessing the effects of clam aquaculture on nektonic and benthic assemblages in two shallow-water estuaries. Journal of Shellfish Research, 35 (4), 757-775. DOI: 10.2983/035.035.0405

Bushek, D., & S. E. Ford. (2016). Anthropogenic impacts on an oyster metapopulation: Pathogen introduction, climate change and responses to natural selection. Elementa Science of the Anthropocene, 4: 000119. DOI: 10.12952/journal.elementa.000119

Powell, E. N., R. Mann, K. Ashton-Alcox, Y. Kim, & D. Bushek. (2016). The allometry of oysters: spatial and temporal variation in the length–biomass relationships for Crassostrea virginica. Journal of the Marine Biological Association of the United Kingdom, 96 (5), 1127-1144. DOI: 10.1017/S0025315415000703

McGurk, E. S., S. E. Ford, & D. Bushek. (2016). Unusually abundant and large ciliate xenomas in oysters, Crassostrea virginica, from Great Bay, New Hampshire, USA. Journal of Invertabrate Pathology, 137, 23-32. DOI: 10.1016/j.jip.2016.04.001

Carnegie, R. B., I. Arzul, & D. Bushek. (2016). Managing marine mollusc diseases in the context of regional and international commerce: Policy issues and emerging concerns. Philosophical Transactions of the Royal Society B, 371 (1689), 11pp. DOI: 10.1098/rstb.2015.0215

Powell, E. N., Y. Kim, & D. Bushek. (2015). Temporal structure and trends of parasites and pathologies in U.S. oysters and mussels: 16 years of mussel watch. Journal of Shellfish Research, 34 (3), 967-993. DOI: 10.2983/035.034.0325

Dungan, C. F., & D. Bushek. (2015). Development and applications of Ray's fluid thioglycollate media for detection and manipulation of Perkinsus spp. pathogens of marine molluscs. Journal of Invertebrate Pathology, 131, 68–82. DOI: 10.1016/j.jip.2015.05.004

Ben-Horin, T., G. Bidegain, L. Huey, D. A. Narváez, & D. Bushek. (2015). Parasite transmission through suspension feeding. Journal of Invertebrate Pathology, 131, 155–176. DOI: 10.1016/j.jip.2015.07.006

Baggett, L. P., S. P. Powers, R. D. Brumbaugh, L. D. Coen, B. M. DeAngelis, J. K. Greene, B. T. Hancock, S. M. Morlock, B. L. Allen, D. L. Breitburg, D. Bushek, J. H. Grabowski, R. E. Grizzle, E. D. Grosholz, M. K. La Peyre, M. W. Luckenbach, K. A. McGraw, M. F. Piehler, S. R. Westby, & P. S. E. zu Ermgassen. (2015). Guidelines for evaluating performance of oyster habitat restoration. Restoration Ecology, 23 (6), 737–745. DOI: 10.1111/rec.12262

Munroe, D. M., A. Tabatabai, I. Burt, D. Bushek, E. N. Powell & J. Wilkin. (2013). Oyster mortality in Delaware Bay: Impacts and recovery from Hurricane Irene and Tropical Storm Lee. Estuarine, Coastal and Shelf Science, 135, 209-219. DOI: 10.1016/j.ecss.2013.10.011

Ford, S. E., & D. Bushek. (2012). Development of resistance to an introduced marine pathogen by a native host. Journal of Marine Research 70 (2-3), 205-223. DOI: 10.1357/002224012802851922

Bushek, D., S. E. Ford, & I. Burt. (2012). Long-term patterns of an estuarine pathogen along a salinity gradient. Journal of Marine Research, 70 (2-3), 225-251. DOI: 10.1357/002224012802851968

Ford, S. E., E. Scarpa, & D. Bushek. (2012). Spatial and temporal variability of disease refuges in an estuary: Implications for the development of resistance. Journal of Marine Research 70 (2-3), 253-277. DOI: 10.1357/002224012802851850

Wang, Z., D. Haidvogel, D. Bushek, S. E. Ford, E. E. Hofmann, E. N. Powell, & J. Wilkins. (2012). Circulation and water properties and their relationship to the oyster disease MSX in Delaware Bay. Journal of Marine Research, 70 (2-3), 279-308. DOI: 10.1357/002224012802851931

He, Y., S. E. Ford, D. Bushek, E. N. Powell, Z. Bao, & X. Guo. (2012). Effective population sizes of eastern oyster Crassostrea virginica (Gmelin) populations in Delaware Bay, USA. Journal of Marine Research 70 (2-3), 357-379. DOI: 10.1357/002224012802851977

Powell, E. N., J. M. Klinck, X. Guo, E. E. Hofmann, S. E. Ford, & D. Bushek. (2012). Can oysters Crassostrea virginica develop resistance to dermo disease in the field: The impediment posed by climate cycles. Journal of Marine Research, 70 (2-3), 309-355. DOI: 10.1357/002224012802851896

Ford, S. E., J. Paterno, E. Scarpa, N. A. Stokes, Y. Kim, E. N. Powell, & D. Bushek. (2011). Widespread survey finds no evidence of Haplosporidium nelsoni (MSX) in Gulf of Mexico oysters. Diseases of Aquatic Organisms, 93 (3), 251-256. DOI: 10.3354/dao02306

Kraeuter, J. N., S. E. Ford, D. Bushek, W. C. Walton, D. C. Murphy, G. Flimlin, & G. Mathis. (2011). Evaluation of three northern quahog (=hard clam) Mercenaria mercenaria (Linnaeus) strains grown in Massachusetts and New Jersey for QPX-resistance. Journal of Shellfish Research, 30 (3), 805-812. DOI: 10.2983/035.030.0321

Powell, E. N., J. M. Klink, X. Guo, E. E. Hofmann, S. E. Ford, & D. Bushek. (2011). The potential for oysters, Crassostrea virginica, to develop resistance to dermo disease in the field: Evaluation using a gene-based population dynamics model. Journal of Shellfish Research, 30 (3), 685-712. DOI: 10.2983/035.030.0310

Ford, S. E., J. Paterno, E. Scarpa, Y. Kim, E. N. Powell, & D. Bushek. (2011). Widespread survey finds no evidence of Haplosporidium nelsoni (MSX) in Gulf of Mexico oysters. Diseases of Aquatic Organisms, 93 (3), 251-256. DOI: 10.3354/dao02306

Wang S., E. Peatman, H. Liu, D. Bushek, S. E. Ford, H. Kucuktas, J. Quilang, P. Li, R. Wallace, Y. Wang, X. Guo, & Z. Liu. (2010). Microarray analysis of gene expression in eastern oyster (Crassostrea virginica) reveals a novel combination of antimicrobial and oxidative stress host responses after dermo (Perkinsus marinus) challenge. Fish & Shellfish Immunology, 29 (6), 921-929. DOI: 10.1016/j.fsi.2010.07.035

Hofmann, E. E., D. Bushek, S. E. Ford, X. Guo, D. Haidvogel, D. Hedgecock, J. M. Klinck, C. Milbury, D. A. Narváez, E. N. Powell, Y. Wang, Z. Wang, J. Wilkin, L. Zhang. (2009). Understanding how disease and environment combine to structure resistance in estuarine bivalve populations. Oceanography, 22 (4), 212-231. DOI: 10.5670/oceanog.2009.110

Ford, S. E., N. Stokes, E. Burreson, E. Scarpa, R. Carnegie, J. N. Kraeuter, & D. Bushek. (2009). Minchinia mercenariae n. sp. (Haplosporidia) in the hard clam Mercenaria mercenaria: Implications of a rare parasite in a commercially important host. Journal of Eukaryotic Microbiology, 56 (6), 542-551. DOI: 10.1111/j.1550-7408.2009.00432.x

Gray, B. R., D. Bushek, J. W. Drane, & D. Porter. (2009). Associations between land use and Perkinsus marinus infection of eastern oysters in a high salinity, partially urbanized estuary. Ecotoxicology, 18 (2), 259-269. DOI: 10.1007/s10646-008-0279-9

Powell, E. N., J. N. Kraeuter, K. Ashton-Alcox, S. E. Ford, & D. Bushek. (2008). Long-term trends in oyster population dynamics in Delaware Bay: Regime shifts and response to disease. Journal of Shellfish Research, 27 (4), 729-755. DOI: 10.2983/0730-8000(2008)27[729:LTIOPD]2.0.CO;2

Taylor, J., & D. Bushek. (2008). Intertidal oyster reefs can persist and function in a temperate North American Atlantic estuary. Marine Ecology Progress Series, 361, 301-306. DOI: 10.3354/meps07429

Richards, G. P., M. A. Watson, E. J. Crane, I. Burt, & D. Bushek. (2008). Shewanella and Photobacterium spp. in oysters and seawater from the Delaware Bay. Applied and Environmental Microbiology, 74 (11), 3323-3327. DOI: 10.1128/AEM.00060-08

Bushek, D., B. Landau, & E. Scarpa. (2008). Perkinsus chesapeaki in stout razor clams Tagelus pleius from Delware Bay. Diseases of Aquatic Organisms, 78 (3), 243-247. DOI: 10.3354/dao01871

Bushek, D., A. Kornbluh, H. Wang, X. Guo, G. DeBrosse, & J. Quinlan. (2008). Fertilization interference between Crassostrea ariakensis and Crassostrea virginica: A gamete sink? Journal of Shellfish Research, 27 (3), 593-600. DOI: 10.2983/0730-8000(2008)27[593:FIBCAA]2.0.CO;2

Coen L. D., R. D. Brumbaugh, D. Bushek, R. Grizzle, M. W. Luckenbach, M. H. Posey, S. P. Powers, & G. S. Tolley. (2007). AS WE SEE IT: Ecosystem services related to oyster restoration. Marine Ecology Progress Series, 341, 303-307. DOI: 10.3354/meps341303

Bushek, D., M. Heidenreich, & D. Porter. (2007). The effects of several common anthropogenic contaminants on proliferation of the parasitic oyster pathogen Perkinsus marinus. Marine Environmental Research, 64 (4), 535-540. DOI: 10.1016/j.marenvres.2007.03.004

Allen, D. M., S. S. Haertel-Borer, B. J. Milan, D. Bushek, & R. D. Dame. (2007). Geomorphological determinants of nekton use of intertidal salt marsh creeks. Marine Ecology Progress Series, 329, 57-71. DOI: 10.3354/meps329057

Ellin, R., & D. Bushek. (2006). Adaptation of Ray's fluid thioglycollate medium assay to detect and quantify planktonic stages of Perkinsus spp. parasites. Journal of Shellfish Research, 25 (3), 1037-1042. DOI: 10.2983/0730-8000(2006)25[1037:AORFTM]2.0.CO;2

Bushek, D., & S. Boyd. (2006). Seasonal abundance and occurrence of the Asian isopod Synidotea laevidorsalis in Delaware Bay, USA. Biological Invasions, 8 (4), 697-702. DOI: 10.1007/s10530-005-2890-5

Toline, C. A., P. Kenny, & D. Bushek. (2005). Cement-coated stakes enhance recruitment of damaged oyster reefs (South Carolina). Ecological Restoration, 23 (4), 277-288. DOI: 10.3368/er.23.4.271

Bushek, D., & D. M. Allen. (2005). Motile suspension-feeders in estuarine and marine ecosystems. In R. F. Dame & S. Olenin (Eds.) The Comparative Role of Suspension Feeders in Ecosystems (pp. 53-71). NATO Science Series: IV - Earth and Environmental Sciences, Vol. 47. Dordrecht, The Netherlands: Springer. DOI: 10.1007/1-4020-3030-4_4

Bushek, D., D. Richardson, M. Y. Bobo, & L. D. Coen. (2004). Quarantine of oyster shell cultch reduces the abundance of Perkinsus marinus. Journal of Shellfish Research, 23 (2), 369-373.

Courses Taught




Oceanographic Methods & Data Analysis

Current Post-Doctoral Researchers

Recent Post-Doctoral Researchers

Tal Ben-Horin

Host pathogen interactions in marine and estuarine systems

Paola Lopez-Duarte

Can elemental signatures from larval shells reveal source populations of settling oyster larvae?

Current Students

Heidi Yeh Ph.D.

Oyster microbiome dynamics

Recent Students

Rachel Eggleston RIOS Intern

Long-term shifts in timing of oyster reproduction in Delaware Bay

Casey Jones B.S. (2018)

NSF RIOS Intern - sea level rise impacts on horseshoe crabs and red knots

Amanda Wenczel Ph.D. (2018)

Filter feeder impacts in estuarine ecosystems

Mike DiLorenzo B.S. (2017)

Summer Intern - shellfish hatchery phytoplankton cultivation

Bree Cerione B.S. (2016)

Summer Intern - Living shoreline nekton use and oyster monitoring

Jenny Paterno M.S. (2015)

Assessment of fish utilization on an oyster restoration area in the Delaware Bay

Emily Scarpa M.S. (2013)


Joshua Moody M.S. (2012)

The relationship of the ribbed mussel, Geukensia demissa, to shoreline erosion in Delaware Bay


President 2007-2009, National Shellfisheries Association

[Haskin Shellfish Research Laboratory]