Marine Biology

, Volume 159, Issue 7, pp 1391–1402 | Cite as

Kelp as a trophic resource for marine suspension feeders: a review of isotope-based evidence

  • Robert J. Miller
  • Henry M. Page
Review, Concept, and Synthesis


Kelp forests are enormously productive, and they and adjacent habitats support large populations of suspension feeders. What do these suspension feeders eat? Intuitively, we might expect that kelp primary production is a key form of trophic support for these animals. Indeed, a large and growing number of studies using carbon stable isotope data, typically collected over short time periods, have asserted that detritus from kelps is an important, and in some cases the main, food source for coastal benthic suspension feeders. This view has been incorporated into several textbooks and review papers covering kelp forest ecosystems, and loss of trophic support for benthic suspension feeders is now often invoked as an ecosystem consequence of top-down or other impacts on kelp forests. More direct evidence, however, suggests that these animals mainly eat phytoplankton and, in some cases, bacteria or zooplankton. Because isotope values of pure coastal phytoplankton, uncontaminated with detritus, are difficult to obtain, present studies have largely relied on single measurements from offshore environments or from the literature, which typically reflects offshore values. We review the evidence showing that phytoplankton isotope values can, and are expected to, vary widely in coastal waters and that inshore phytoplankton may often be enriched in 13C compared to offshore phytoplankton. This unaccounted-for variation may have systematically biased the results of such trophic studies toward finding large contributions of kelp detritus to suspension-feeder diets. We review some key stable isotope studies and put forth evidence for alternative explanations of the isotope patterns presented. Finally, we make recommendations for future isotope studies and describe several approaches for progress in this area. New techniques, particularly flow cytometry and compound-specific stable isotope analysis, provide ways to shed light on this interesting and important ecological issue.


Phytoplankton Detritus Dissolve Inorganic Carbon Particulate Organic Matter Suspension Feeder 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



This manuscript benefitted from discussions and comments by M. Brzezinski and two anonymous reviewers. This work was supported by the U. S. National Science Foundation’s Long Term Ecological Research Program under Division of Ocean Sciences grant numbers 9982105 and 0620276 and by NSF Bio-Ocean award 0962306 to HM Page.


  1. Allan E, Ambrose S, Richoux N, Froneman P (2010) Determining spatial changes in the diet of nearshore suspension-feeders along the South African coastline: stable isotope and fatty acid signatures. Estuar Coast Shelf Sci 87:463–471Google Scholar
  2. Attwood C, Lucas M, Probyn T, McQuaid C, Fielding P (1991) Production and standing stocks of the kelp Macrocystis laevis Hay at the Prince Edward Islands, sub-Antarctic. Polar Biol 11:129–133Google Scholar
  3. Barnes C, Jennings S, Polunin NC, Lancaster JE (2008) The importance of quantifying inherent variability when interpreting stable isotope field data. Oecologia 155:227–235Google Scholar
  4. Beninger P, Decottignies P (2005) What makes diatoms attractive for suspensivores? The organic casing and associated organic molecules of Coscinodiscus perforatus are quality cues for the bivalve Pecten maximus. J Plankton Res 27:11–17Google Scholar
  5. Bidigare R, Kennicutt M, Keeneykennicutt W, Macko S (1991) Isolation and purification of chlorophyll-a and chlorophyll-b for the determination of stable carbon and nitrogen isotope compositions. Anal Chem 63:130–133Google Scholar
  6. Bode A, Alvarez-Ossorio MT, Varela M (2006) Phytoplankton and macrophyte contributions to littoral food webs in the Galician upwelling estimated from stable isotopes. Mar Ecol Prog Ser 318:89–102Google Scholar
  7. Brett M, Kainz M, Taipale S, Seshan H (2009) Phytoplankton, not allochthonous carbon, sustains herbivorous zooplankton production. Proc Natl Acad Sci 106:21197–21201Google Scholar
  8. Burton R, Snodgrass J, Gifford-Gonzalez D, Guilderson T, Brown T, Koch P (2001) Holocene changes in the ecology of northern fur seals: insights from stable isotopes and archaeofauna. Oecologia 128:107–115Google Scholar
  9. Bustamante RH, Branch GM (1996) The dependence of intertidal consumers on kelp-derived organic matter on the west coast of South Africa. J Exp Mar Biol Ecol 196:1–28Google Scholar
  10. Bustamante RH, Branch GM, Eekhout S (1995) Maintenance of an exceptional intertidal grazer biomass in South Africa—subsidy by subtidal kelps. Ecology 76:2314–2329Google Scholar
  11. Cifuentes L, Sharp J, Fogel M (1988) Stable carbon and nitrogen isotope biogeochemistry in the Delaware Estuary. Limnol Oceanogr 33:1102–1115Google Scholar
  12. Coma R, Ribes M, Gili J, Zabala M (2000) Seasonality in coastal benthic ecosystems. Trends Ecol Evol 15:448–453Google Scholar
  13. Corbisier T, Petti M, Skowronski R, Brito T (2004) Trophic relationships in the nearshore zone of Martel Inlet (King George Island, Antarctica): δ13C stable isotope analysis. Polar Biol 27:75–82Google Scholar
  14. Cranford P, Grant J (1990) Particle clearance and absorption of phytoplankton and detritus by the sea scallop Placopecten magellanicus Gmelin. J Exp Mar Biol Ecol 137:105–122Google Scholar
  15. Dayton P (1985) Ecology of kelp communities. Ann Rev Ecol Syst 16:215–245Google Scholar
  16. Duarte C, Cebrian J (1996) The fate of marine autotrophic production. Limnol Oceanogr 41:1758–1766Google Scholar
  17. Dugan J, Hubbard D, McCrary M, Pierson M (2003) The response of macrofauna communities and shorebirds to macrophyte wrack subsidies on exposed sandy beaches of southern California. Estuar Coast Shelf Sci 58:25–40Google Scholar
  18. Duggins D, Eckman J (1994) The role of kelp detritus in the growth of benthic suspension feeders in an understory kelp forest. J Exp Mar Biol Ecol 176:53–68Google Scholar
  19. Duggins D, Eckman J (1997) Is kelp detritus a good food for suspension feeders? effects of kelp species, age and secondary metabolites. Mar Biol 128:489–495Google Scholar
  20. Duggins D, Simenstad C, Estes J (1989) Magnification of secondary production by kelp detritus in coastal marine ecosystems. Science 245:170–173Google Scholar
  21. Dunton KH (1985) Trophic dynamics in marine nearshore systems of the Alaskan high Arctic, PhD thesis, University of Alaska Fairbanks, pp 247Google Scholar
  22. Dunton K (2001) δ15N and δ13C measurements of Antarctic peninsula fauna: Trophic relationships and assimilation of benthic seaweeds. Am Zool 41:99–112Google Scholar
  23. Dunton K, Schell D (1987) Dependence of consumers on macroalgal (Laminaria solidungula) carbon in an arctic kelp community: δ13C evidence. Mar Biol 93:615–625Google Scholar
  24. Dunton K, Weingartner T, Carmack E (2006) The nearshore western Beaufort Sea ecosystem: circulation and importance of terrestrial carbon in arctic coastal food webs. Prog Oceanogr 71:362–378Google Scholar
  25. Evans S, Anderson W, Jochem F (2006) Spatial variability in Florida Bay particulate organic matter composition: combining flow cytometry with stable isotope analyses. Hydrobiol 569:151–165Google Scholar
  26. France R, Cattaneo A (1998) Delta C-13 variability of benthic algae: effects of water colour via modulation by stream current. Freshw Biol 39:617–622Google Scholar
  27. Fredriksen S (2003) Food web studies in a Norwegian kelp forest based on stable isotope (delta C-13 and delta N-15) analysis. Mar Ecol Prog Ser 260:71–81Google Scholar
  28. Fry B, Sherr E (1984) δ13C measurements as indicators of carbon flow in marine and fresh-water ecosystems. Contrib Mar Sci 27:13–47Google Scholar
  29. Fry B, Wainright S (1991) Diatom sources of 13C-rich carbon in marine food webs. Mar Ecol Prog Ser 76:149–157Google Scholar
  30. Gaye-Siessegger J, Focken U, Muetzel S, Abel H, Becker K (2004) Feeding level and individual metabolic rate affect δ13C and δ15N values in carp: implications for food web studies. Oecologia 138:175–183Google Scholar
  31. Gili J-M, Coma R (1998) Benthic suspension feeders: their paramount role in littoral marine food webs. Trends Ecol Evol 13:316–324Google Scholar
  32. Goering J, Alexander V, Haubenstock N (1990) Seasonal variability of stable carbon and nitrogen isotope ratios of organisms in a North Pacific bay. Estuar Coast Shelf Sci 30:239–260Google Scholar
  33. Gollety C, Riera P, Davoult D (2010) Complexity of the food web structure of the Ascophyllum nodosum zone evidenced by a δ13C and δ15N study. J Sea Res 64:304–312Google Scholar
  34. Graham MH (2004) Effects of local deforestation on the diversity and structure of Southern California giant kelp forest food webs. Ecosystems 7:341–357Google Scholar
  35. Graham BS, Koch PL, Newsome SD, McMahon KW, Aurioles D (2010) Using isoscapes to trace the movements and foraging behavior of top predators in oceanic ecosystems. In: West JB, Bowen GJ, Dawson TE (eds) Isoscapes: understanding movement, pattern, and process on earth through isotope mapping. Springer, Dordrecht, pp 299–318Google Scholar
  36. Hamilton S, Sippel S, Bunn S (2005) Separation of algae from detritus for stable isotope or ecological stoichiometry studies using density fractionation in colloidal silica. Limnol Oceanogr Methods 3:149–157Google Scholar
  37. Hill JM, McQuaid CD, Kaehler S (2006) Biogeographic and nearshore-offshore trends in isotope ratios of intertidal mussels and their food sources around the coast of southern Africa. Mar Ecol Prog Ser 318:63–73Google Scholar
  38. Hill J, McQuaid C, Kaehler S (2008) Temporal and spatial variability in stable isotope ratios of SPM link to local hydrography and longer term SPM averages suggest heavy dependence of mussels on nearshore production. Mar Biol 154:899–909Google Scholar
  39. Hunt G, Stabeno P (2005) Oceanography and ecology of the Aleutian Archipelago: spatial and temporal variation. Fish Oceanogr 14:292–306Google Scholar
  40. Jacob U, Brey T, Fetzer I, Kaehler S, Mintenbeck K, Dunton K, Beyer K, Struck U, Pakhomov EA, Arntz WE (2006) Towards the trophic structure of the Bouvet Island marine ecosystem. Polar Biol 29:106–113Google Scholar
  41. Jaschinski S, Brepohl D, Sommer U (2008) Carbon sources and trophic structure in an eelgrass Zostera marina bed, based on stable isotope and fatty acid analyses. Mar Ecol Prog Ser 358:103–114Google Scholar
  42. Jørgensen CB (1990) Bivalve filter feeding : hydrodynamics, bioenergetics, physiology and ecology. Olsen & Olsen, FredensborgGoogle Scholar
  43. Kaehler S, Pakhomov EA, McQuaid CD (2000) Trophic structure of the marine food web at the Prince Edward Islands (Southern Ocean) determined by δ13C and δ15N analysis. Mar Ecol Prog Ser 208:13–20Google Scholar
  44. Kaehler S, Pakhomov EA, Kalin RM, Davis S (2006) Trophic importance of kelp-derived suspended particulate matter in a through-flow sub-Antarctic system. Mar Ecol Prog Ser 316:17–22Google Scholar
  45. Kang C, Sauriau P, Richard P, Blanchard G (1999) Food sources of the infaunal suspension-feeding bivalve Cerastoderma edule in a muddy sandflat of Marennes Oleron Bay, as determined by analyses of carbon and nitrogen stable isotopes. Mar Ecol Prog Ser 187:147–158Google Scholar
  46. Kang C, Choy E, Son Y, Lee J, Kim J, Kim Y, Lee K (2008) Food web structure of a restored macroalgal bed in the eastern Korean peninsula determined by C and N stable isotope analyses. Mar Biol 153:1181–1198Google Scholar
  47. Kokkinakis SA, Wheeler PA (1987) Nitrogen uptake and phytoplankton growth in coastal upwelling regions. Limnol Oceanogr 32:1112–1123Google Scholar
  48. Kostadinov T, Siegel D, Maritorena S, Guillocheau N (2007) Ocean color observations and modeling for an optically complex site: Santa Barbara Channel, California, USA. J Geophys Res, Oceans 112: C07011. doi: 10.1029/2006JC003526
  49. Lastra M, Page H, Dugan J, Hubbard D, Rodil I (2008) Processing of allochthonous macrophyte subsidies by sandy beach consumers: estimates of feeding rates and impacts on food resources. Mar Biol 154:163–174Google Scholar
  50. Laws E, Popp B, Cassar N, Tanimoto J (2002) 13C discrimination patterns in oceanic phytoplankton: likely influence of CO2 concentrating mechanisms, and implications for palaeoreconstructions. Funct Plant Biol 29:323–333Google Scholar
  51. Levinton JS, Ward JE, Shumway SE (2002) Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus. Mar Biol 141:367–376Google Scholar
  52. Linley E, Newell R, Bosma S (1981) Heterotrophic utilization of mucilage released during fragmentation of kelp (Ecklonia maxima and Laminaria pallida): 1. development of microbial communities associated with the degradation of kelp mucilage. Mar Ecol Prog Ser 4:31–41Google Scholar
  53. Lubetkin S, Simenstad C (2004) Multi-source mixing models to quantify food web sources and pathways. J Appl Ecol 41:996–1008Google Scholar
  54. Lucas AJ, Dupont CL, Tai V, Largier JL, Palenik B, Franks PJS (2011) The green ribbon: multiscale physical control of phytoplankton productivity and community structure over a narrow continental shelf. Limnol Oceanogr 56:611–626Google Scholar
  55. McLeod R, Wing S (2007) Hagfish in the New Zealand fjords are supported by chemoautotrophy of forest carbon. Ecology 88:809–816Google Scholar
  56. Michener RH, Kaufman L (2007) Stable isotope ratios as tracers. In: Michener RH, Lajtha K (eds) Stable isotopes in ecology and environmental science. Blackwell Pub, Malden, pp 238–282Google Scholar
  57. Miller T, Brodeur R, Rau G (2008) Carbon stable isotopes reveal relative contribution of shelf-slope production to the northern California Current pelagic community. Limnol Oceanogr 53:1493–1503Google Scholar
  58. Monteiro P, James A, Sholtodouglas A, Field J (1991) The δ13C trophic position isotope spectrum as a tool to define and quantify carbon pathways in marine food webs. Mar Ecol Prog Ser 78:33–40Google Scholar
  59. Moore J, Semmens B (2008) Incorporating uncertainty and prior information into stable isotope mixing models. Ecol Lett 11:470–480Google Scholar
  60. Mordy C, Stabeno P, Ladd C, Zeeman S, Wisegarver D, Salo S, Hunt G (2005) Nutrients and primary production along the eastern Aleutian Island Archipelago. Fish Oceanogr 14:55–76Google Scholar
  61. Nadon MO, Himmelman JH (2006) Stable isotopes in subtidal food webs: have enriched carbon ratios in benthic consumers been misinterpreted? Limnol Oceanogr 51:2828–2836Google Scholar
  62. Nakatsuka T, Handa N, Wada E, Wong C (1992) The dynamic changes of stable isotopic ratios of carbon and nitrogen in suspended and sedimented particulate organic matter during a phytoplankton bloom. J Mar Res 50:267–296Google Scholar
  63. Nelson J (1993) Rates and possible mechanism of light-dependent degradation of pigments in detritus derived from phytoplankton. J Mar Res 51:155–179Google Scholar
  64. Newell RIE (2004) Ecosystem influences of natural and cultivated populations of suspension-feeding bivalve molluscs: a review. J Shellfish Res 23:51–61Google Scholar
  65. Newell R, Field J, Griffiths C (1982) Energy balance and significance of microorganisms in a kelp bed community. Mar Ecol Prog Ser 8:103–113Google Scholar
  66. Newell RIE, Kemp WM, Hagy JD III, Cerco CF, Testa JM, Boynton WR (2007) Top-down control of phytoplankton by oysters in Chesapeake Bay, USA: comment on Pomeroy et al. (2006). Mar Ecol Prog Ser 341:293–298Google Scholar
  67. Norderhaug KM, Fredriksen S, Nygaard K (2003) Trophic importance of Laminaria hyperborea to kelp forest consumers and the importance of bacterial degradation to food quality. Mar Ecol Prog Ser 255:135–144Google Scholar
  68. Norkko A, Thrush SF, Cummings VJ, Gibbs MM, Andrew NL, Norkko J, Schwarz AM (2007) Trophic structure of coastal antarctic food webs associated with changes in sea ice and food supply. Ecology 88:2810–2820Google Scholar
  69. Officer C, Smayda T, Mann R (1982) Benthic filter feeding: a natural eutrophication control. Mar Ecol Prog Ser 9:203–210Google Scholar
  70. Overmyer J, MacNeil M, Fisk A (2008) Fractionation and metabolic turnover of carbon and nitrogen stable isotopes in black fly larvae. Rap Comm Mass Spec 22:694–700Google Scholar
  71. Page HM, Lastra M (2003) Diet of intertidal bivalves in the Ria de Arosa (NW Spain): evidence from stable C and N isotope analysis. Mar Biol 143:519–532Google Scholar
  72. Page HM, Reed DC, Brzezinski MA, Melack JM, Dugan JE (2008) Assessing the importance of land and marine sources of organic matter to kelp forest food webs. Mar Ecol Prog Ser 360:47–62Google Scholar
  73. Paine R (2002) Trophic control of production in a rocky intertidal community. Science 296:736–739Google Scholar
  74. Parnell A, Inger R, Bearhop S, Jackson A (2010) Source partitioning using stable isotopes: coping with too much variation. Plos One. doi: 10.1371/journal.pone.0009672 Google Scholar
  75. Pel R, Floris V, Gons H, Hoogveld H (2004) Linking flow cytometric cell sorting and compound-specific 13C analysis to determine population-specific isotopic signatures and growth rates in cyanobacteria-dominated lake plankton. J Phycol 37:857–866Google Scholar
  76. Perga M, Kainz M, Mazumder A (2008) Terrestrial carbon contribution to lake food webs: could the classical stable isotope approach be misleading? Can J Fish Aquat Sci 65:2719–2727Google Scholar
  77. Perissinotto R, Duncombe Rae C (1990) Occurrence of anticyclonic eddies on the Prince Edward Plateau (Southern Ocean): effects on phytoplankton biomass and production. Deep-Sea Res Part A 37:777–793Google Scholar
  78. Perissinotto R, Duncombe Rae C, Boden B, Allanson B (1990) Vertical stability as a controlling factor of the marine phytoplankton production at the Prince-Edward archipelago (Southern Ocean). Mar Ecol Prog Ser 60:205–209Google Scholar
  79. Perry R, Thompson P, Mackas D, Harrison P, Yelland D (1999) Stable carbon isotopes as pelagic food web tracers in adjacent shelf and slope regions off British Columbia, Canada. Can J Fish Aquat Sci 56:2477–2486Google Scholar
  80. Phillips D, Gregg J (2003) Source partitioning using stable isotopes: coping with too many sources. Oecologia 136:261–269Google Scholar
  81. Post D (2002) Using stable isotopes to estimate trophic position: models, methods, and assumptions. Ecology 83:703–718Google Scholar
  82. Rau G, Riebesell U, Wolf-Gladrow D (1996) A model of photosynthetic 13C fractionation by marine phytoplankton based on diffusive molecular CO2 uptake. Mar Ecol Prog Ser 133:275–285Google Scholar
  83. Ribes M, Coma R, Gili J (1999) Heterogeneous feeding in benthic suspension feeders: the natural diet and grazing rate of the temperate gorgonian Paramuricea clavata (Cnidaria : Octocorallia) over a year cycle. Mar Ecol Prog Ser 183:125–137Google Scholar
  84. Ricciardi A, Bourget E (1999) Global patterns of macroinvertebrate biomass in marine intertidal communities. Mar Ecol Prog Ser 185:21–35Google Scholar
  85. Rooker J, Turner J, Holt S (2006) Trophic ecology of sargassum-associated fishes in the Gulf of Mexico determined from stable isotopes and fatty acids. Mar Ecol Prog Ser 313:249–259Google Scholar
  86. Sackmann B, Mack L, Logsdon M, Perry M (2004) Seasonal and inter-annual variability of SeaWiFS-derived chlorophyll a concentrations in waters off the Washington and Vancouver Island coasts, 1998–2002. Deep-Sea Res Part I 51:945–965Google Scholar
  87. Salomon A, Shears N, Langlois T, Babcock R (2008) Cascading effects of fishing can alter carbon flow through a temperate coastal ecosystem. Ecol Appl 18:1874–1887Google Scholar
  88. Salomon AK, Gaichas SK, Shears NT, Smith JE, Madin EMP, Gaines SD (2010) Key features and context-dependence of fishery-induced trophic cascades. Cons Biol 24:382–394Google Scholar
  89. Sargent J, Bell M, Henderson R, Tocher D (1990) Polyunsaturated fatty acids in marine and terrestrial food webs. In: Mellinger J (ed) Animal nutrition and transport processes 1: Nutrition in wild and domestic animals, pp 11–23Google Scholar
  90. Saupe S, Schell D, Griffiths W (1989) Carbon isotope ratio gradients in western Arctic zooplankton. Mar Biol 103:427–432Google Scholar
  91. Savoye N, Aminot A, Treguer P, Fontugne M, Naulet N, Kerouel R (2003) Dynamics of particulate organic matter δ15N and δ13C during spring phytoplankton blooms in a macrotidal ecosystem (Bay of Seine, France). Mar Ecol Prog Ser 255:27–41Google Scholar
  92. Schaal G, Riera P, Leroux C (2009) Trophic significance of the kelp Laminaria digitata (Lamour.) for the associated food web: a between-sites comparison. Estuar Coast Shelf Sci 85:565–572Google Scholar
  93. Schaal G, Riera P, Leroux C (2010) Trophic ecology in a Northern Brittany (Batz Island, France) kelp (Laminaria digitata) forest, as investigated through stable isotopes and chemical assays. J Sea Res 63:24–35Google Scholar
  94. Schell D, Barnett B, Vinette K (1998) Carbon and nitrogen isotope ratios in zooplankton of the Bering, Chukchi and Beaufort seas. Mar Ecol Prog Ser 162:11–23Google Scholar
  95. Schlacher T, Connolly R (2009) Land-ocean coupling of carbon and nitrogen fluxes on sandy beaches. Ecosystems 12:311–321Google Scholar
  96. Seiderer L, Newell R (1985) Relative significance of phytoplankton, bacteria and plant detritus as carbon and nitrogen resources for the kelp bed filter-feeder Choromytilus meridionalis. Mar Ecol Prog Ser 22:127–139Google Scholar
  97. Seiderer L, Newell R (1988) Exploitation of phytoplankton as a food resource by the kelp bed ascidian Pyura stolonifera. Mar Ecol Prog Ser 50:107–115Google Scholar
  98. Simenstad CA, Duggins DO, Quay PD (1993) High turnover of inorganic carbon in kelp habitats as a cause of δ13C variability in marine food webs. Mar Biol 116:147–160Google Scholar
  99. Smayda T (1997) Harmful algal blooms: their ecophysiology and general relevance to phytoplankton blooms in the sea. Limnol Oceanogr 42:1137–1153Google Scholar
  100. Soares AG, Schlacher TA, McLachlan A (1997) Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region. Mar Biol 127:657–664Google Scholar
  101. Steneck R, Graham M, Bourque B, Corbett D, Erlandson J, Estes J, Tegner M (2002) Kelp forest ecosystems: biodiversity, stability, resilience and future. Environ Conserv 29:436–459Google Scholar
  102. Stuart V, Field J, Newell R (1982) Evidence for absorption of kelp detritus by the ribbed mussel Aulacomya ater using a new Cr-51 labeled microsphere technique. Mar Ecol Prog Ser 9:263–271Google Scholar
  103. Takahashi K, Wada E, Sakamoto M (1992) Carbon isotope ratio and photosynthetic activity of phytoplankton in the eutrophic Mikawa Bay, Japan. Ecol Res 7:355–361Google Scholar
  104. Tallis H (2009) Kelp and rivers subsidize rocky intertidal communities in the Pacific Northwest (USA). Mar Ecol Prog Ser 389:85–96Google Scholar
  105. Thomas A, Strub P, Carr M, Weatherbee R (2004) Comparisons of chlorophyll variability between the four major global eastern boundary currents. Int J Remote Sens 25:1443–1447Google Scholar
  106. Thompson M, Schaffner L (2001) Population biology and secondary production of the suspension feeding polychaete Chaetopterus cf. variopedatus: Implications for benthic-pelagic coupling in lower Chesapeake Bay. Limnol Oceanogr 46:1899–1907Google Scholar
  107. Tortell PD, Rau GH, Morel FMM (2000) Inorganic carbon acquisition in coastal Pacific phytoplankton communities. Limnol Oceanogr 45:1485–1500Google Scholar
  108. Tremblay J, Michel C, Hobson K, Gosselin M, Price N (2006) Bloom dynamics in early opening waters of the Arctic Ocean. Limnol Oceanogr 51:900–912Google Scholar
  109. van Duyl F, Moodley L, Nieuwland G, van Ijzerloo L, van Soest R, Houtekamer M, Meesters E, Middelburg J (2011) Coral cavity sponges depend on reef-derived food resources: stable isotope and fatty acid constraints. Mar Biol 158:1–14Google Scholar
  110. Verity P, Beatty T, Williams S (1996) Visualization and quantification of plankton and detritus using digital confocal microscopy. Aquat Microb Ecol 10:55–67Google Scholar
  111. Vetter E, Dayton P (1999) Organic enrichment by macrophyte detritus, and abundance patterns of megafaunal populations in submarine canyons. Mar Ecol Prog Ser 186:137–148Google Scholar
  112. Wildish D, Kristmanson DD (1997) Benthic suspension feeders and flow. Cambridge University Press, CambridgeGoogle Scholar
  113. Williams S, Verity P, Beatty T (1995) A new staining technique for dual identification of plankton and detritus in seawater. J Plankton Res 17:2037–2047Google Scholar

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© Springer-Verlag 2012

Authors and Affiliations

  1. 1.Marine Science InstituteThe University of California Santa BarbaraSanta BarbaraUSA

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