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Senior Scientist
Executive Vice President and Director of Research
Woods Hole Oceanographic Institution
Woods Hole, Massachusetts 02543
July 28, 2008
A.B., University of California, Berkeley, 1969
Ph.D., University of California, Davis, 1974
University of California, Davis
Teaching Assistant, 1970-1971, 1972-1974
Research Assistant, 1972
Oregon State University
Visiting Associate Professor, 9/87 – 12/87
Woods Hole Oceanographic Institution:
Postdoctoral Scholar, 1974 -1975
Assistant Scientist, 1975 -1979
Associate Scientist, 1979 -1996
J. Seward Johnson Chair in Education, 1994 -1996
Senior Scientist, 1996 Chair, Biology Department, 1996 - 2000
Director, Ocean Life Institute, 2000 – 2006
Acting Director of Research, 2006 – 2008
Executive Vice President and Director of Research 2008 – present
Monterey Bay Aquarium Research Institute
Adjunct Scientist, 2000 SOCIETY MEMBERSHIPS:
American Geophysical Union
American Society of Limnology and Oceanography
Sigma Xi
Distribution, behavior, functional morphology, energetics, systematics and ecology of
gelatinous zooplankton, especially tunicates, ctenophores and medusae. Symbiotic
relationships among zooplankton. Vertical distribution and diel migration. Biology of
mesopelagic and benthopelagic zooplankton. Trophic structure of pelagic ecosystems.
Sensory biology and behavior. Flux of organic matter in the oceans. Instruments for
pelagic survey.
Graduate Students Advised:
David Caron, 1979-1984
Carol Diebel, 1983-1988
Rebecca Rapoza, 1994 - (LOA)
Annette Govindarajan, 1998 – 2004
Kelly Rakow, 2004 Postdoctoral Scholars/Investigators Sponsored:
Jennifer Purcell, 1981
Charles Greene, 1985
Anne Sell, 1998-2001
Sonke Johnsen, 1999-2000
Introductory Biological Oceanography, 1976, 1980
Lectures on Invertebrate Zoology and Zooplankton, Biological Oceanography Course,
Instructor for "Topics in Zooplankton Ecology” 1994, 1996, 2001, 2002
Biology Department Education Coordinator, 1982-85, 1995-96
Biology Curriculum Committee, 1995-2001
Biology Exam Committee, 2004, Chair, 2005
Associate Editor, Limnology and Oceanography Methods
Member, Marine Advisory Panel, Conservation International
Member, Steering Committee, Mount Hope Bay Natural Laboratory Project
Member, Zoning Working Group, Stellwagen Bank National Marine Sanctuary
Member, Steering Committee, Census of Marine Zooplankton
Scientific Advisor, New England Aquarium Jellies exhibit
Visiting Scholar, Auckland Museum, Auckland, New Zealand 2005
Adamík, P., S.M. Gallager, E. Horgan, L.P. Madin, A. Govindarajan and P. Alatalo.
2006. Effects of turbulence on the feeding rate of a pelagic predator: the
planktonic hydroid Clytia gracilis. J. Exp. Mar. Biol. Ecol. 333:159-165.
Madin, L.P., P. Kremer, P.H. Wiebe, J. E. Purcell, E. H. Horgan and D. A. Nemazie.
2006. Periodic swarms of the salp Salpa aspera in the Slope Water off the NE
United States: Biovolume, vertical migration, grazing, and vertical flux. Deep-sea
Res. I. 53:804-819.
Brown, H., S.M. Bollens, L.P. Madin and E.F. Horgan. 2005. Effects of warm water
intrusions on populations of macrozooplankton on Georges Bank, Northwest
Atlantic". Cont. Shelf Res. 25:143-156.
Stone, G.S., L.P. Madin, K. Stocks, G. Hovermale, P. Hoagland, M. Schumacher, P.
Etnoyer, C. Sotka, and H. Tausig. 2004. Chapter 2: Seamount Biodiversity,
Exploitation and Conservation. In: Glover, L.K. and S.A. Earle. (eds). Defying
Ocean’s End: An Agenda for Action. Island Press, Washington, D.C. pp. 43-70.
Madin, L.P., F. Grassle, F. Azam, D. Obura, M. Reaka-Kudla, M. Sibuet, G.S. Stone, K.
Stocks, A. Walls and G. R. Allen. 2004. Chapter 12: The Unknown Ocean. In:
Glover, L.K. and S.A. Earle. (eds). Defying Ocean’s End: An Agenda for Action.
Island Press, Washington, D.C. pp. 213-236.
Vargas, C.A. and L.P. Madin. 2004. Zooplankton feeding ecology: Clearance and
ingestion rates of the salps Thalia democratica, Cyclosalpa affinis and Salpa
cylindrica on naturally occurring particles in the Mid-Atlantic Bight. J. Plankton
Res. 26:827-833.
Roman, M.R., H.A. Adolf, M.R. Landry, L.P. Madin, D.K. Steinberg and X. Zhang. 2002.
Estimates of oceanic mesozooplankton production: A comparison using the
Bermuda and Hawaii time-series data. Deep-Sea Res. II 49:175-192.
Bathmann, U., Bundy, M.H., Clarke, M.E., Cowles, T.J., Daly, K., Dam, H.G.,
Dekshenieks, M.M., Donaghay, P.L., Gibson, D.M., Gifford, D.J., Hansen, B.W.,
Hartline, D.K., Head, E.J.H., Hofmann, E.E., Hopcroft, R.R., Jahnke, R.A.,
Jonasdottir, S.H., Kiorboe, T., Kleppel, G.S., Klinck, J.M., Kremer, P.M., Landry,
M.R., Lee, R.F., Lenz, P.H., Madin, L.P., Manahan, D.T., Mazzocchi, M.G.,
McGillicuddy, D.J., Miller, C.B., Nelson, J.R., Osborn, T.R., Paffenhofer, G.A.,
Pieper, R.E., Prusova, I., Roman, M.R., Schiel, S., Seim, H.E., Smith, S.L.,
Torres, J.J., Verity, P.G., Wakeham, S.G., Wishner, K.F. 2001. Future marine
zooplankton research -- a perspective. Mar. Ecol. Progr. Ser. 222:297-308.
Madin, L.P. and G.R. Harbison. 2001. Gelatinous Zooplankton. In. J. Steele, S. Thorpe, K.
Turekian (eds.), Encyclopedia of Ocean Sciences. Academic Press. London.
Madin, L.P. 2001. Taxonomic Outline of Marine Organisms. In. J. Steele, S. Thorpe, K.
Turekian (eds.), Encyclopedia of Ocean Sciences. Academic Press. London.
Butler, M., S.M. Bollens, B. Burkhalter, L. P. Madin, and E. Horgan. 2001.
Mesopelagic fishes of the Arabian Sea: Distribution, Abundance and Diet of
Chauliodus pammelas, C. sloani, Stomias affinis, and S. nebulosus. Deep-Sea
Res. II. 48:1369-1383.
Madin, L.P., E.F. Horgan and D.K. Steinberg. 2001. Zooplankton at the Bermuda
Atlantic Time-series Study (BATS) station: Diel, Seasonal and Interannual
Variation in Biomass, 1994-1998. Deep-Sea Res. II 48:2063-2082
Sell, A.F., D. van Keuren and L.P. Madin. 2001. Predation by omnivorous copepods on
early developmental stages of Calanus finmarchicus and Pseudocalanus spp.
Limnol. Oceanogr. 46:953-959.
Bollens, S.M., E. Horgan, S. Concelman, L.P. Madin and M. Butler. 2001. Planktonic
Hydroids on Georges Bank: Effects of mixing and food supply on feeding and
growth. Deep-Sea Res. II 48:659-672.
Concelman, S. S.M. Bollens, B. Sullivan, L.P. Madin, E. Horgan, M. Butler and D.
vanKeuren. 2001. Distribution, Abundance and Benthic-Pelagic Coupling of
Suspended Hydroids on Georges Bank. Deep-Sea Res. II 48:645-658.
Mincks, S.L., S. M. Bollens, L. P. Madin, E. Horgan, M. Butler, P. Kremer, and J. E.
Craddock. 2000. Distributions, Abundance, and Feeding Ecology of Decapods in
the Arabian Sea, With Implications for Vertical Flux. Deep-Sea Res. 47:14751516.
Steinberg, D.K., C.A. Carlson, N.R. Bates, S.A. Goldthwait, L.P. Madin, and A.F.
Michaels. 2000. Zooplankton vertical migration and the active transport of
dissolved organic and inorganic carbon in the Sargasso Sea. Deep-Sea Res.
Steinberg, D.K., Madin, L.P. and F.T. Thingstad. 2000. Influence of zooplankton
community structure on export processes at the Bermuda Atlantic Time-series
Study (BATS) Site. In: Ocean Biogeochemistry: A New Paradigm. pp. 89-90.
Madin, L.P. and D. Deibel. 1998. Chapter 5: Feeding and energetics of Thaliaceans, In
Bone, Q. ed., The Biology of Pelagic Tunicates, Oxford Univ. Press. pp 43-64.
Couwelaar, M. van, M.V. Angel and L.P. Madin. 1997. The distribution and biology of the
swimming crab Charybdis smithii McLeay 1838 (Crustacea; Brachyura; Portunidae)
in the NW Indian Ocean. Deep-Sea Res. 44(6-7):1251-1280.
Madin, L.P., J.E. Purcell and C.B. Miller. 1997. Abundance and grazing effects of
Cyclosalpa bakeri (Tunicata, Thaliacea) in the subarctic Pacific. Mar. Ecol. Prog.
Ser. 157:175-183.
Madin, L.P., P. Kremer and S. Hacker. 1996. Distribution and vertical migrations of salps
(Tunicata, Thaliacea) near Bermuda. J. Plankton Res. 18:747-755.
Madin, L.P., S.M. Bollens, E. Horgan, M. Butler, J. Runge, B.K. Sullivan, G. KleinMacPhee, E. Durbin, A. Durbin, A. Bucklin, M.E. Clarke. 1996. Voracious
Planktonic Hydroids: Unexpected Predatory Impact on a coastal marine ecosystem.
Deep-Sea Res. 43:1823-1829.
Roman, M.R., D.A. Caron, P. Kremer, E.J. Lessard, L.P. Madin, T.C. Malone, J.M. Napp,
E.R. Peele and M.J. Youngbluth. 1995. Spatial and temporal changes in the
partitioning of organic carbon in the plankton community of the Sargasso Sea off
Bermuda. Deep-Sea Res. 42:973-992.
Caron, D.A., H.G. Dam, P. Kremer, E.J. Lessard, L.P. Madin, T.C. Malone, J.M. Napp,
E.R. Peele, M.R. Roman and M.J. Youngbluth. 1995. The contribution of
microorganisms to particulate carbon and nitrogen in surface waters of the Sargasso
Sea near Bermuda. Deep-Sea Res. 42:943-972.
Madin, L.P. and P. Kremer. 1995. Determination of the filter feeding rates of salps
(Tunicata, Thaliacea). ICES J. Mar. Sci. 52:583-595.
Madin, L.P. 1995. Sensory ecology of salps (Tunicata, Thaliacea): More questions than
answers. Mar. Freshw. Behav. Physiol. 26:175-195.
Tebeau, C.M. and L.P. Madin. 1994. Grazing rates for three life history stages of the
doliolid Dolioletta gegenbauri Uljanin. (Tunicata, Thaliacea). J. Plankton Res.
Hart, M.W., R.L. Miller and L.P. Madin. 1994. Form and feeding mechanism of a living
Planctosphaera pelagica (Phylum Hemichordata). Mar. Biol. 120:521-533.
Kremer, P. and L.P. Madin. 1992. Particle retention efficiency of salps. J. Plankton Res.
Madin, L.P. and J.E. Purcell. 1992. Feeding, metabolism and growth of Cyclosalpa bakeri
in the Subarctic Pacific. Limnol. Oceanogr. 37:1236-1251.
Larson, R.J., G.I. Matsumoto, L.P. Madin and L.M. Lewis. 1992. Deep-sea benthic and
benthopelagic medusae: recent observations using submersibles and a remotely
operated vehicle. Bull. Mar. Sci. 51:277-286.
Hacker, S. and L.P. Madin. 1991. Why habitat architecture and color are important to
shrimp living in pelagic Sargassum: The use of camouflage and plant-part mimicry.
Mar. Ecol. Progr. Ser. 70:143-155.
Purcell, J.E. and L. P. Madin. 1991. Diel vertical migration, feeding and spawning of the
salp Cyclosalpa bakeri in the Subarctic Pacific. Mar. Ecol. Prog. Ser. 73:211-217.
Atema, J., P. A. Moore, L.P. Madin and G.A. Gerhardt. 1991. Subnose-1: Electrochemical
tracking of odor plumes at 900 m beneath the ocean surface. Mar. Ecol. Prog. Ser.
Madin, L. P. 1990. Aspects of jet propulsion in salps. Can. J. Zool. 68: 765-777.
Madin, L.P. 1990. Being there: the role of in-situ science in oceanography. Marine
Technology Soc. J. 24: 19-21.
Caron, D. A., L. P. Madin and J. J. Cole. 1989. Salp fecal pellets: Microbial degradation
and the vertical flux of material in oceanic plankton communities. J. Mar. Res. 47:
Larson, R. J., L. P. Madin and G. R. Harbison. 1988. In situ observations of the rare
deepwater scyphomedusa Deepstaria, with description of a new species. J. Mar.
Biol. Ass. U.K. 68: 689-699.
Madin, L. P. 1988. Feeding behavior of tentaculate predators: In situ observations and a
conceptual model. Bull. Mar. Sci. 43: 413-429.
Madin, L. P. 1988. Beebe Project. Zooplankton studies in the 1987 field season. In:
Global Venting, Midwater and Benthic Ecological Processes, M. DeLuca and I.
Babb (eds.), National Undersea Research Program Research Report 88-4: 291-306.
Cetta, C. M., L. P. Madin and P. Kremer. 1986. Respiration and excretion by oceanic salps.
Mar. Biol. 91: 529-537.
Caron, D. A., P. G. Davis, L. P. Madin and J. McN. Sieburth. 1986. Enrichment of
microbial populations in macroaggregates (marine snow) from surface waters of the
North Atlantic. J. Mar. Res. 44: 543-565.
Madin, L. P. 1985. In situ photography of gelatinous zooplankton. In: D. George, G.
Lythgue and J. Georges (eds.), Underwater Photography and Television for
Scientists. Clarendon Press, Oxford. pp. 65-82.
Madin, L. P. and C. M. Cetta. 1984. The use of gut fluorescence to estimate grazing by
oceanic salps. J. Plank. Res. 6: 475-482.
Madin, L. P. and N. R. Swanberg. 1984. In situ methods for the study of planktonic
organisms. In: N. D. Flemming (ed.), Divers, Submersibles and Marine Science,
Memorial University of Newfoundland Occasional Papers in Biology No. 9: 4-16.
Harbison, G. R. and L. P. Madin. 1982. The Ctenophora. In: S. P. Parker (ed.), Synopsis
and Classification of Living Organisms. McGraw-Hill, New York.
Madin, L. P. 1982. The production, composition and sedimentation of salp fecal pellets in
oceanic waters. Mar. Biol. 67: 39-45.
Alldredge, A. L. and L. P. Madin. 1982. Pelagic tunicates: Unique herbivores in the
marine plankton. BioScience 32: 655-663.
Caron, D. A., P. G. Davis, L. P. Madin and J. McN. Sieburth. 1982. Enrichment of
heterotrophic bacteria and bacteriovorous protozoa in oceanic macroaggregates.
Science 218: 795-797.
Madin, L. P., C. M. Cetta and V. L. McAlister. 1981. Elemental and biochemical
composition of salps (Tunicata: Thaliacea). Mar. Biol. 63: 217-226.
Wiebe, P. H., L. P. Madin, L. R. Haury, G. R. Harbison and L. M. Philbin. 1979. Diel
vertical migration by Salpa aspera: Potential for large-scale particulate organic
matter transport to the deep sea. Mar. Biol. 53: 249-255.
Madin, L. P. and G. R. Harbison. 1978. Salps of the genus Pegea Savigny 1816. Bull.
Mar. Sci. 28: 335-344.
Harbison, G. R., L. P. Madin and N. R. Swanberg. 1978. On the natural history and
distribution of oceanic ctenophores. Deep-Sea Res. 25: 233-256.
Madin, L. P. and G. R. Harbison. 1978. Bathocyroe fosteri gen. et sp. nov., a mesopelagic
ctenophore observed and collected from a submersible. J. Mar. Biol. Assoc. U.K.
58: 559-564.
Madin, L. P. and G. R. Harbison. 1978. Thalassocalyce inconstans gen. n. sp. n. an
enigmatic ctenophore representing a new family and order. Bull. Mar. Sci. 28: 680687.
Madin, L. P. and G. R. Harbison. 1977. The associations of amphipods Hyperiidea with
gelatinous zooplankton - I. Association with Salpidae. Deep-Sea Res. 24: 449463.
Harbison, G. R., D. C. Biggs and L. P. Madin. 1977. The association of amphipods
Hyperiidea with gelatinous zooplankton - II. Associations with Cnidaria,
Ctenophora and Radiolaria. Deep-Sea Res. 24: 465-488.
Carpenter, E. J., G. R. Harbison, L. P. Madin, N. R. Swanberg, D. C. Biggs, E. M. Hulburt,
V. L. McAlister and J. J. McCarthy. 1977. Rhizosolenia mats. Limnol. Oceanogr.
22: 739-741.
Harbison, G. R. and L. P. Madin. 1976. Description of the female Lycaea nasuta Claus
1879 with an illustrated key to the species of Lycaea (Hyperiidea, Lycaeidae). Bull.
Mar. Sci. 26: 165-171.
Hamner, W. M., L. Madin, A. Alldredge, R. Gilmer and P. Hamner. 1975. Underwater
observations of gelatinous zooplankton: sampling problems, feeding biology and
behavior. Limnol. Oceanogr. 20: 907-917.
Madin, L. P. 1974. Field observations of the feeding behavior of salps (Tunicata:
Thaliacea). Mar. Biol. 25: 143-147.
Madin, L.P. Pelagic Tunicates on the California Coast. In. J. Carlton, ed. The Light and
Smith Manual. Univ. Calif. Press.
Jiang, S., T. D. Dickey, D. K. Steinberg, L. P. Madin. 2007. Temporal Variability of
Zooplankton Biomass from ADCP Backscatter Time Series Data at the Bermuda
Testbed Mooring Site. Deep-Sea Res. I. In press.
Stoltz, G.S., S.M. Bollens, H. Brown, L.P. Madin and E.F. Horgan. The distribution and
population biology of the ctenophore Pleurobrachia pileus on Georges Bank,
Northwest Atlantic. Deep-Sea Res.
Madin, L.P., E. Horgan, S. Gallager, J. Eaton and A. Girard. 2006. LAPIS: A new imaging
tool for macrozooplankton. Proceedings, IEEE/MTS Oceans ’06 Meeting.
Madin, L.P. (ed.) 2002. Fisheries, Oceanography and Society. Marine Protected Areas:
Design and Implementation for Conservation and Fisheries Restoration. Proceedings
of Symposium of the Ocean Life Institute. 68 pp.
Madin, L.P. 2001. Initial Report and Proposed Science Plan for Bahia Honda. Ocean Life
Institute internal document, 16 pp.
Madin, L.P. 1997. Synopsis of Diversity of Invertebrate Micronekton (except
Cephalopods) in the World Ocean. White Paper for Non-Fish Nekton Workshop,
New England Aquarium, Boston.
Madin, L.P., E.F. Horgan, B.K. Sullivan, G. Klein-MacPhee, S.M. Bollens and M. Butler.
1997. Estimating the impact of invertebrate predation on target species of copepods
on Georges Bank. ICES 1997 Annual Science Conference, Baltimore, MD.
Madin, L.P. and M.R. Landry, eds. 1995. U.S. GLOBEC Workshop on Dynamics of Open
Ocean Populations. U.S. GLOBEC Report No. 14, 24 pp.
Madin, L.P. 1991. Distribution and taxonomy of zooplankton in the Alboran Sea and
adjacent western Mediterranean. A literature survey and field guide. Report for
Harbor Branch Oceanographic Institution and Naval Oceanographic and
Atmospheric Research Laboratory. W.H.O.I. Technical Report 91-26, 148 pp.
Madin, L.P. 1991. The use of submersibles for research on deepwater zooplankton.
Proceedings of the UJNR Diving Panel Session on Deep Ocean Science, Hakone,
Japan. p. 269-280.
Dacey, J.W.H., Craddock, J.E., Madin, L.P., Michaels, A.F. and J.R. Weinberg. 1989.
Biological impact of deep sea carbon dioxide disposal. p. 64-76. In: Golomb,D.,
Herzog, H., Tester, J., White, D. and S. Zemba. Feasibility, modeling and economics
of sequestering power plant CO2 emissions in the deep ocean. Energy Laboratory,
Massachusetts Institute of Technology, Cambridge, MA. p. 1-76
Madin, L. 2005. The Ocean Life Institute. Oceanus Magazine 43:4-5.
Madin, L. 2005. A muster of scientists for a dearth of whales. Oceanus Magazine 43:3435.
Madin, L.P. and K.A.C. Madin. 1995. Diversity in a vast and stable habitat. Oceanus
Madin, K.A.C. and L.P. Madin. 1991. Sex (and Asex) in the Jellies. Oceanus 34:27-35.
Harbison, G. R. and L. P. Madin. 1979. Diving - A new view of plankton biology.
Oceanus 22:18-27.
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