Distinguished Professor Emerita

Research Interests

Phytoplankton physiology and ecology; marine nitrogen cycle; nutrient use by phytoplankton and bacteria; macrophyte physiology and ecology; nitrogen metabolism in macrophytes.

Current Research

Nitrogen uptake and recycling in the Arctic Ocean; use of inorganic and organic nitrogen by marine bacteria; relationship between autotrophic and heterotrophic activity in the microbial food web; determination of bacterial biomass and activity in the euphotic zone.


BA, University of California, Irvine, 1971
MS, University of California, Irvine, 1974
PhD, University of California, Irvine, 1976


* = refereed journals

* Wheeler, P.A., B.B. North, and G.C. Stephens. 1974. Amino acid uptake by marine phytoplankters. Limnol. Oceanogr. 19: 249-259.

* Wheeler, P., B. North, M. Littler, and G. Stephens. 1977. Uptake of glycine by natural phytoplankton communities. Limnol. Oceanogr. 22: 900-910.

* Wheeler, P.A. and G.C. Stephens. 1977. Metabolic segregation of intracellular free amino acids in Platymonas (Chlorophyta) .J.Phycol. 13: 193-197.

* Wheeler, P.A. 1977. Effect of nitrogen source on Platymonas (Chlorophyta) cell composition and amino acid uptake rates. J. Phycol. 13: 304-306.

* Haines, K.C. and P.A. Wheeler. 1978. Ammonium and nitrate uptake by the marine macrophytes Hypnea musciformis (Rhodophyta) and Macrocystis pyrifera (Phaeophyta). J. Phycol. 14: 319-324.

North, W.J. and P.A. Wheeler. 1979. Nutritional requirements of the giant kelp Macrocystis (Phaeophyceae). Proc. Intl. Seaweed Symp. 9: 67-78.

* Wheeler, P.A. 1979. Uptake of methylamine (an ammonium analogue) by Macrocystis pyrifera (Phaeophyta). J. Phycol.. 15: 12-17.

Wheeler, P.A. 1980. Uptake of methylamine by the marine diatom Cyclotella cryptica. In: R.M. Spanswick, W.J. Lucas, and J. Dainty [Eds], Plant Membrane Transport: Current Conceptual Issues, Elsevier/North Holland Biomedical Press, pp. 633-634.

* Wheeler, P.A. 1980. Use of methylamine as an ammonium analogue in transport and assimilation studies with Cyclotella cryptica (Bacillariophyceae). J. Phycol. 16: 328-334.

* Wheeler, P.A. and W.J. North. 1980. Effect of nitrogen supply on nitrogen content and growth rates of juvenile Macrocystis pyrifera (Phaeophyceae) sporophytes. J. Phycol. 16: 577-582.

* Wheeler, P.A. and J.A. Hellebust. 1981. Uptake and concentration of alkylamines by a marine diatom: effects of H+ and K+ and implications for the transport and accumulation of weak bases. Plant Physiol. 67: 367-372.

* Wheeler, P.A. and W.J. North. 1981. Nitrogen supply, tissue composition, and frond growth rates for Macrocystis pyrifera off the coast of southern California. Mar. Biol. 64: 577-582.

* Wheeler, P.A., P.M. Glibert, and J.J. McCarthy. 1982. Ammonium uptake and incorporation by Chesapeake Bay phytoplankton: short-term uptake kinetics. Limnol. Oceanogr. 27: 1113-1128.

* Wheeler, P.A. and J.J. McCarthy. 1982. Methylammonium uptake by Chesapeake Bay phytoplankton: evaluation of the use of the ammonium analogue for field uptake measurements. Limnol. Oceanogr. 27: 1129-1140.

Wheeler, P.A. 1983. Phytoplankton nitrogen metabolism. In: E.J. Carpenter and D.C. Capone [Eds], Nitrogen in the Marine Environment, Academic Press, pp. 309-346.

* Wheeler, P.A., R.J. Olson, and S.W. Chisholm. 1983. The effects of photocycles and periodic ammonium supply on three marine phytoplankton species. II. Ammonium uptake and assimilation. J. Phycol. 19: 528-533.

Wheeler, P.A. 1985. Nutrients. In: M.M. Littler and D.S. Littler [Eds], Ecological Field Methods: Macroalgae, Handbook of Phycological Methods, Cambridge University Press, pp. 53-64.

Harlin, M.M. and P.A. Wheeler. 1985. Nutrient uptake. In: M.M. Littler and D.S. Littler [Eds], Ecological Field Methods: Macroalgae, Handbook of Phycological Methods, Cambridge University Press, pp. 493-508.

* Dudek, N., M.A. Brzezinski, and P.A. Wheeler. 1986. Measurement of 15N-ammonium isotope enrichment in seawater using a solvent extraction technique. Mar. Chem. 18: 59-69.

* Wheeler, P.A. and D.L. Kirchman. 1986. Utilization of inorganic and organic forms of nitrogen by bacteria in marine systems. Limnol. Oceanogr. 31: 998-1009.

* Smith, S.V., W.J. Wiebe, J.T. Hollibaugh, S.J. Dollar, S.W. Hager, B.E. Cole, G.W. Tribble, and P.A. Wheeler. 1987. Stoichiometry of C, N, P, and Si fluxes in a temperate-climate embayment. J. Mar. Res. 45: 427-460.

* Kokkinakis, S.A. and P.A. Wheeler. 1987. Nitrogen uptake and phytoplankton growth in temperate coastal upwelling regions. Limnol. Oceanogr. 32: 1112-1123.

* O'Brien, M.C. and P.A. Wheeler. 1987. Short term uptake of nutrients by Enteromorpha prolifera (Mull.) J. Ag. J. Phycol. 23: 547-556.

* Miller, C.B., K.L. Denman, A.E. Gargett, D.L. Mackas, P.A. Wheeler, B.C. Booth, B.W. Frost, M.R. Landry, J. Lewin, C.J. Lorenzen, M.J. Perry, M. Dagg, and N. Welschmeyer. 1988. Lower trophic level production dynamics in the oceanic subarctic Pacific Ocean. Bull. Ocean Res. Inst., Univ. Tokyo, No. 26: 1-26.

* Kokkinakis, S.A. and P.A. Wheeler. 1988. Uptake of ammonium and urea in the northeast Pacific: comparison between netplankton and nanoplankton. Mar. Ecol. Prog. Ser. 43: 113-124.

* Fujita, R.M., P.A. Wheeler, and R.L. Edwards. 1988. Metabolic regulation of ammonium uptake by Ulva rigida C. Ag. (Chlorophyta): A compartmental analysis of the rate-limiting step for uptake. J. Phycol. 24: 560-566.

* Fujita, R.M., P.A. Wheeler, and R.L. Edwards. 1989. Assessment of macroalgal nitrogen limitation in a seasonal upwelling region. Mar. Ecol. Prog. Ser. 53: 293-303.

* Wheeler, P.A., D.L. Kirchman, M.R. Landry, and S.A. Kokkinakis. 1989. Diel periodicity in ammonium uptake and regeneration in the oceanic subarctic Pacific: Implications for interactions in microbial foodwebs. Limnol. Oceanogr. 34: 1025-1033.

* Kirchman, D.L., R.G. Keil, and P.A. Wheeler. 1989. The effect of amino acids on ammonium utilization and regneration by heterotrophic bacteria in the subarctic Pacific. Deep-Sea Res. 36: 1763-1776.

* Kirchman, D.L., R.G. Keil, and P.A. Wheeler. 1990. Carbon limitation of ammonium uptake by heterotrophic bacteria in the subarctic Pacific. Limnol. Oceanogr. 35: 1258-1266.

* Wheeler, P.A. and S.A. Kokkinakis. 1990. Ammonium recycling limits nitrate use in the oceanic subarctic Pacific. Limnol. Oceanogr. 35: 1267-1278.

* Bjornsater, B.R. and P.A. Wheeler. 1990. Effect of nitrogen and phosphorus supply on growth and tissue composition of Ulva fenestrata and Enteromorpha intestinalis (Ulvales: Chlorophyta). J. Phycol. 26: 603-611.

* Miller, C.B., B.W. Frost, P.A. Wheeler, M.R. Landry, N. Welschmeyer, and T.M. Powell. 1991. Ecological dynamics in the subarctic Pacific, a possibly iron limited system. Limnol. Oceanogr. 36: 1600-1615.

* Miller, C.B., B.W. Frost, B. Booth, P.A. Wheeler, M.R. Landry, and N. Welschmeyer. 1991. Ecological processes in the subarctic Pacific: Iron Limitation cannot be the whole story. Oceanography 4: 71-78.

* Wheeler, P.A. and B.R. Bjornsater. 1992. Seasonal flucutations in tissue nitrogen, phosphorus, and N:P for five macroalgal species common to the Pacific Northwest coast. J. Phycol. 28: 1-6.

* Emerson, S., P. Quay, and P.A. Wheeler. 1993. Biological productivity determined from oxygen mass balance and incubation experiments. Deep-Sea Res. 40: 2351-2358.

* Dickson, M.-L. and P.A. Wheeler. 1993. Chlorophyll a concentrations in the North Pacific: Does a latitudinal gradient exist? Limnol. Oceanogr. 38: 1813-1818.

* Wheeler, P.A. 1993. New production in the oceanic subarctic Pacific: net changes in NO3- concentrations, rates of NO3- assimilation, and accumulation of particulate nitrogen. Progress in Oceanography 32: 137-161.

* Landry, M.R., D.J. Gifford, D.L. Kirchman, P.A. Wheeler, and B.C. Monger. 1993. Direct and indirect effects of grazing by Neocalanus plumchrus on plankton community dynamics in the subarctic Pacific. Progress in Oceanography 32: 239-258.

* Kristiansen, S., T. Farbrot, and P.A. Wheeler. 1994. Nitrogen cycling in the Barents Sea - Seasonal dynamics of new and regenerated production in the marginal ice zone. Limnol. Oceanogr. 39: 1630-1642.

* Libby, P.S. and P.A. Wheeler. 1994. A wet-oxidation method for determination of particulate organic nitrogen on glass fiber and 0.2 µm membrane filters. Mar. Chem. 48: 31-41.

* Dickson, M.-L. and P. A. Wheeler. 1995. Comment on the chlorophyll a retention properties of glass fiber GF/F filters. Limnol. Oceanogr. 40: 434-436.

* Dickson, M.-L. and P.A. Wheeler. 1995. Nitrogen uptake rates in a coastal upwelling regime: A comparison of PN-specific, absolute and Chl a specific rates. Limnol. Oceanogr. 40: 533-543.

* Dickson, M.-L. and P.A. Wheeler. 1995. Ammonium uptake and regeneration rates in a coastal upwelling regime. Mar. Ecol. Prog. Ser. 121:239-248.

* Menge, B.A., B. Daley, and P.A. Wheeler. 1995. Control of interaction strength in marine benthic communities.pp. 258-274 In: Food Webs: Integration of Pattern and Dynamics, G.A. Polis and R. Winemiller, [Eds.], Chapman and Hall, New York.

* Wheeler, P.A., M. Gosselin, E. Sherr, D. Thibault, D.L. Kirchman, R. Benner, and T.E. Whitledge. 1996. Active cycling of organic carbon in the central Arctic Ocean. Nature 380:697-699.

* Aagaard, K., L.A. Barrie, E.C. Carmack, C. Garrity, E.P. Jones, D. Lubin, R.W. MacDonald, J.H. Swift, W.B. Tucker, P.A. Wheeler, and R.H. Whritner. 1996. U.S., Canadian researchers explore Arctic Ocean. EOS 77: 209-213.

* Libby, P.S. and P.A. Wheeler.1997. Particulate and dissolved organic nitrogen in the central equatorial Pacific. Deep-Sea Res. 44: 345-361.

* Menge, B.A., B. Daley, P.A. Wheeler and P.T. Strub. 1997. Rocky intertidal oceanography: an association between community structure and nearshore phytoplankton concentration. Limnol. Oceanogr. 42: 57-66.

* Menge, B.A., B.A. Daley, P. A. Wheeler, E. Dahlhoff, E. Sanford, and P.T. Strub. 1997. Benthic- pelagic links and rocky intertidal communities: Bottom-up effects on top-down control. Proc. Natl. Acad. Sci. USA 94: 14530-14535.

* Wheeler, P.A. 1997. Preface: The 1994 Arctic Ocean Section. Deep-Sea Res. 44: 1483-1486.

* Wheeler, P.A., J.M. Watkins, and R.L. Hansing.1997. Nutrients, organic carbon and organic nitrogen in the surface water and halocline of the Arctic Ocean: Implications for the sources of dissolved organic carbon. Deep-Sea Res. 44: 1571-1592.

* Gosselin, M., M. Levasseur, P.A. Wheeler, R. Horner, and B.C. Booth. 1997. New measurements of phytoplankton and ice algal production in the Arctic Ocean. Deep-Sea Res. 44: 1623-1644.

* Kirchman, D.L. and P.A. Wheeler. 1998. Uptake of inorganic nitrogen by heterotrophic bacteria and phytoplankton in the subarctic Pacific. Deep-Sea Res. 45: 347-365.

Grebmeier, J.M., T.E. Whitledge, L.A. Codispoti, K.H. Dunton, J.J. Walsh, and P.A. Wheeler. [Eds] 1998. Arctic System Science Ocean-Atmosphere-Ice Interactions Western Arctic Shelf-Basin Interaction Science Plan. ARCSS/OAII Report Number 7, Old Dominion University, Norfolk, VA, 65 pp.

* Thibault, D., E.J.H. Head, and P.A. Wheeler. 1999. Mesozooplankton in the Arctic Ocean in summer. Deep-Sea Res. 46: 1931-1415.

* Naldi, M. and P.A. Wheeler. 1999. Changes in nitrogen pools in Ulva fenestrata (Chlorophyta) and Gracilaria pacifica (Rhodophyta) under nitrate and ammonium enrichment. J. Phycol. 35: 70-77.

* Wheeler, P.A. 1999. Cell geometry revisited: Realistic shapes and accurate determination of cell volume and surface area from microscopic measurements. J. Phycol. 35: 209-210.

Fleischbein, J., J. Hill, A. Huyer, R.L. Smith, and P.A. Wheeler. Hydrographic data from GLOBEC Long-term Observation Program off Oregon, 1997 and 1998. 1999. College of Oceanic and Atmospheric Sciences, Oregon State University, Data Report 172, Reference 99-1.

* Ortiz, J.D., A.C. Mix, P.A. Wheeler, and R.M. Key. 2000. An estimate of the anthropogenic offset of oceanic 13C DIC based on the ventiliation of the California Current at 42o N. Global Biogeochem. Cycles 14: 917-929.

* Durazo, R., T.R. Baumgartner, S.J. Bograd, C.A. Collins, S. de la Campa, J. Garcia, G. Gaxiola- Castro, A. Huyer, K.D. Hydrenbach, D. Loya, R.J. Lynn, F.B. Schwing, R.L. Smith, W.J. Sydeman, and P. A. Wheeler. 2001. The state of the California Current, 2000-2001: A third straight La Niña Year. CalCOFI Rep. Vol. 42: 1-32.

* Uttal, T., J.A. Curry, M.G. McPhee, D.K. Perovich, R.E. Moritz, J.A. Maslanik, P.S. Guest, H.L. Stern, J.A. Moore, R. Turenne, A. Heiberg, M.C. Serreze, D.P. Wylie, O.G. Persson, C.A. Paulson, C. Halle, J.H. Morison, P.A. Wheeler, A. Makshtas, H. Welch, M.D. Shupe, J.M. Intieri, K. Stamnes, R.W. Lindsey, R. Pinkel, W.S. Pegau, T.P. Stanton, and T.C. Grenfeld. 2002. Surface heat budget of the Arctic Ocean. Bull. Amer. Metero. Soc. 83: 255-275.

* Hill, J.K. and P.A. Wheeler. 2002. Organic carbon and nitrogen in the northern California Current system during July 1997: comparison of offshore, river plume, and coastally upwelled water. Progress in Oceanography, 53:369-387.

* Naldi, M. and P.A. Wheeler. 2002. 15N measurements of ammonium and nitrate uptake by Ulva fenestrata (Chlorophyta) and Gracilaria pacifica (Rhodophyta): Comparison of net nutrient disappearance, release of ammonium, and 15N acculmulation in algal tissue. J. Phycol. 38:135-144.

* Chase, Z., A. van Geen, P.M. Kosro, J. Marra, and P.A. Wheeler. 2002. Iron, nutrient, and phytoplankton distributions in Oregon coastal water. J. Geophys. Res. 10.1029/2000JC000444, 26 October 2002.

* Corwith, H.L. and P.A. Wheeler. 2002. El Niño related variations in nutrient and chlorophyll distibutions off Oregon. Progress in Oceanography, 54: 361-380.

* Ikeda, T. and P.A. Wheeler. 2002. Ocean Impacts from the bottom of the food web to the top: Biological Oceanography Committee Retrospective. PICES Scientific Report No.22, pp. 25-34.

*Schwing, F. B., S.J. Bograd, C.A. Collins, G. Gaxiola-Castro, J. Garcia, R. Goericke, J. Gomez- Valdez, A. Huyer, K.D. Hyrenbach, P.M. Kosro, B.E. Lavaniegos, R.J. Lynn, A.W. Mantyla, M.D. Ohman, W.T. Peterson, R.L. Smith, W.J. Sydeman, E. Venrick, and P.A. Wheeler. 2002. The state of the California Curent, 2001-2002: Will the CCS keep its cool or is El Niño looming? CalCOFI Rep. Vol. 43: 31-68.

* Sherr, E.B., B.F. Sherr, P.A. Wheeler, and K.Thompson. 2003. Temporal and spatial variation in stocks of autotrophic and heterotrophic microbes in the upper water column of the central Arctic Ocean. Deep-Sea Research I. 50: 557-571.

* Wetz, M.S. and P.A. Wheeler. 2003. Production and partitioning of organic matter during simulated phytoplankton blooms. Limnol. Oceanogr. 48: 1808-1817.

* Wheeler, P.A., A. Huyer, and J. Fleischbein. 2003. Cold halocline, increased nutrients and higher chlorophyll off Oregon in 2002. Geophys. Res. Lett. 30 (15), 8021, doi:10.109/2003 GL017395.

* McLaughlin, F., E.C. Carmack, R.W. Macdoanld, H. Melling, J.H. Swift, P.A. Wheeler, B.F. Sherr, and E.B. Sherr. 2003. The juxtaposition of Atlantic and Pacific-origin waters in the Canada Basin, 1997-1998: A basin in transition. Deep-Sea Research I. 51:107-128.

* Vendrick, E., S.J. Bograd, D. Checkley, R. Durazo, G. Gaxiola-Castro, J. Hunter, A. Huyer, K.D. Hyrenbach, B.E. Laveniegos, A. Mantyla, F.B. Schwing, R.L. Smith, W.J. Sydeman, and P.A. Wheeler. 2003. The state of the California Current, 2002-2003: Tropical and subarctic influences vie for dominance. CalCOFI Rep. Vol. 44: 28-60.

* Wetz, M.S. and P.A. Wheeler. 2004. Responses of bacteria and heterotrophic nanoflagellates to simulated phytoplankton blooms. Mar. Ecol. Prog. Ser. 272:49-57.

* Karp-Boss, L., P.A. Wheeler, B. Hales, and P. Covert. 2004. Distributions and variability of POM in a coastal upwelling system. J. Geophys. Res. 109: doi:10.1029/2003JC002184, 2004.

* Wetz, M.S., P.A. Wheeler, and R.M. Letelier. 2004. Light-induced growth of phytoplankton collected during the winter from the benthic boundary layer off Oregon. Mar. Ecol. Prog. Ser. 280: 95-104.

* Sherr, E.B., B.F. Sherr, and P.A. Wheeler. 2004. Distribution of coccoid cyanobacteria and small eukaryotic phytoplankton in the upwelling ecosystem off the Oregon coast during 2001 and 2002. Deep-Sea Research doi:10.1016/j.dsr2.2004.09.020

Moses, W.C. and P.A. Wheeler. 2004. Seasonal and across-shelf trends of the phytoplankton community of the Oregon coastal environment. College of Oceanic and Atmospheric Sciences, Oregon State University, Data Report 194, COAS Reference 2004-2.

* Goericke, R., E. Vendrick, E., A. Mantyla, D. Checkley, S.J. Bograd, F.B. Schwing, , A. Huyer, R.L. Smith, P.A. Wheeler, R. Hooff, W.T. Peterson, G. Gaxiola-Castro, J. Gomez-Valdes, B.E. Laveniegos, , K.D. Hyrenbach, and W.J. Sydeman. 2004. The state of the California Current, 2003-2004: A “rare” normal year. CalCOFI Rep. Vol. 45: 27-59.

Wetz, J.J., Corwith, H., and Wheeler, P.A. 2005. Nutrient and extracted chlorophyll data from the GLOBEC Long-term observation program 1997-2004. COAS, Oregon State University, Data Report 193, Reference 2004-1, 2004, Revised 2005.

Fleischbein, J., Huyer, A., Kosro, P.M., Smith, R.L., and Wheeler, P.A. 2005. Upper ocean water properties and currents along paired sections in the northern California Current Summer 1998-2003. COAS, Oregon State University, Data Report 201, Reference 2005-4.

* Huyer, A., J.H. Fleischbein, J. Keister, P.M. Kosro, N. Perlin, R.L. Smith, and P.A. Wheeler. 2005. Two coastal upwelling domains in the northern California Current system. J. Mar. Res. 63: 901-929.

Wetz, J.J. and Wheeler, P.A. 2005. Pump station data report for the May 2001, August 2001, and January 2003 COAST cruises: Nutrients, extracted chlorophyll, and dissolved and particulate organic carbon and nitrogen. COAS, Oregon State University, Data Report 202, Reference 2005-5.

* Barth, J.A. and P.A. Wheeler 2005. The coastal ocean advances in shelf transport (COAST) project Motivation and Overview. J. Geophys. Res. 110: C10S01, doi:10.1029/2005JC003124.

Fleischbein, J., Huyer, A., Kosro, P.M., Smith, R.L., and Wheeler, P.A. 2005. Upper ocean water properties and currents along paired sections in the northern California Current Spring 1998-2003. COAS, Oregon State University, Data Report 203, Reference 2005-6.

* Goericke, R., E. Vendrick, A. Mantyla, S.J. Bograd, F.B. Schwing, A. Huyer, R.L. Smith, P.A. Wheeler, R. Hoff, W.T. Peterson, N. Lo, G. Gaxiola-Castro, R. Durazo, K.D. Hyrebach, W.J. Sydeman. 2005. The state of the California Current, 2004-2005: Has a new regime begun? CalCOFI Rep. Vol. 46: 32-71.

* Hales, B.L. Karp-Boss, A. Perlin, and P.A. Wheeler. 2006. Oxygen production and carbon sequestration in an upwelling coastal margin. Global Biogeochemical Cycles 20: GB3001, doi:10.1029/2005GB002517, 2006.

* Wetz, M. S., B. Hales, Z. Chase, P.A. Wheeler, M.M. Whitney. 2006. Riverine input of macronutrients, iron, and organic matter to the coastal ocean off Oregon, USA, during the winter. Limnol. Oceanogr. 51: 2221-2231.

Fleischbein, J.H., A. Huyer, P.M. Kosro, R.L. Smith, J.J. Wetz, and P.A. Wheeler. 2006. Upper ocean properties and currents along paired sections in the Northern California Current, September 1999-2003. College of Oceanic and Atmospheric Sciences, Oregon State University, Data Report 204, Reference 2006-1.

Wetz, J.J., J. Arrington, and P.A. Wheeler. 2006. Particulate and dissolved organic carbon and nitrogen data from the GLOBEC Long-term Observation Program, 1997-2004. College of Oceanic and Atmospheric Sciences, Oregon State University, Data Report 204, Reference 2006-1.

*Wetz, M.S. and P.A. Wheeler. 2007. Release of dissolved organic matter by coastal diatoms. Limnol. Oceanogr. 52: 798-807.

* Huyer, A., P.A. Wheeler, P. T. Strub, R.L. Smith, R. Letelier, and P.M. Kosro. 2007. The Newport Line off Oregon – Studies in the North East Pacific. Progress in Oceanography 75: 126-160.

*Wetz, M.S., B. Hales, and P.A. Wheeler. 2008. Degradation of diatom-derived organic matter: Implications for carbon and nitrogen biogeochemistry in coastal environments. Estuarine, Coastal and Shelf Science. 77: 422-432.

*Christensen, J.P., K. Shimada, I. Semiletov, and P.A. Wheeler. 2008. Chlorophyll response to shelf-break upwelling and winds in the Chukchi Sea, Alaska, in Autumn. The Open Oceanography Journal 2: 34-53.

* Hauri, C., Gruber, N. Plattner, G-K. Alin, S., Feely, R.A., Hales, B. and Wheeler, P.A. 2009. Ocean acidification in the California Current System. Oceanography 22: 59-69.

Miller, C.B. and P.A. Wheeler. 2012. Biological Oceanography, Second Edition, Wiley-Blackwell, Hoboken, N.J., 464 pp.