The benthos: the ocean’s last boundary?

Josep-Maria  Gili; Begoña  Vendrell-Simón; Wolf  Arntz; Francesc  Sabater; Joandomènec  Ros

doi:10.3989/scimar.05091.24A

Authors

Josep-Maria Gili Institut de Ciències del Mar-CSIC https://orcid.org/0000-0001-9542-2385
Begoña Vendrell-Simón Institut de Ciències del Mar-CSIC https://orcid.org/0000-0002-7906-268X
Wolf Arntz Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research https://orcid.org/0000-0002-3315-1070
Francesc Sabater Department of Evolutionary Biology, Ecology and Environmental Sciences, University of Barcelona https://orcid.org/0000-0001-6767-231X
Joandomènec Ros Department of Evolutionary Biology, Ecology and Environmental Sciences, University of Barcelona https://orcid.org/0000-0002-5153-5238

DOI:

https://doi.org/10.3989/scimar.05091.24A

Keywords:

ocean boundaries, benthos, benthic-pelagic and pelagic-benthic coupling, Ramon Margalef

Abstract

Benthic communities depend on receiving much of their food from the water column. While sinking, particles are transformed in a discontinuous process and are temporally retained in transitional physical structures, which act as boundaries and contribute to their further transformation. Motile organisms are well-acquainted with boundaries. The number, width and placement of boundaries are related to the degree of particle degradation or transformation. Progressively deepening within each boundary, particles are degraded according to their residence time in the discontinuity and the activity of the organisms temporarily inhabiting that boundary. Finally, particles reach the seafloor and represent the main food source for benthic organisms; the quality and quantity of this food have a strong impact on the development of benthic communities. However, benthic communities not only play the role of a sink of matter: they act as an active boundary comparable to other oceanic boundaries, in accordance with the boundary concept proposed by the ecologist Ramon Margalef.

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References

Abell J., Emerson S., Renaud P. 2000. Distributions of TOP, TON and TOC in the North Pacific subtropical gyre: Implications for nutrient supply in the surface ocean and remineralization in the upper thermocline. J. Mar. Res. 58: 203-222. https://doi.org/10.1357/002224000321511142

Alldredge A.L. 1979. The chemical composition of macroscopic aggregates in two neritic seas. Limnol. Oceanogr. 24: 855-866. https://doi.org/10.4319/lo.1979.24.5.0866

Amouroux J.M., Amouroux J., Bastide J., et al. 1990. Interrelations in a microcosm with a suspension-feeder and a deposit-feeder. I. Experimental study. Oceanol. Acta 13: 61-68.

Arntz W.E., Gili J.M. 2001. A case for tolerance in marine ecology: let us not put out the baby with the bathwater. Sci. Mar. 65 (Suppl. 2): 283-299. https://doi.org/10.3989/scimar.2001.65s2283

Arntz W.E., Brey T., Gallardo V.A. 1994. Antarctic zoobenthos. Oceanogr. Mar. Biol. Ann. Rev. 32: 241-304.

Arntz W.E., Gili J.M., Reise K. 1999. Unjustifiably ignored: reflections on the role of benthos in marine ecosystems. In: Gray J.S., Ambrose W. Jr., et al. (eds) Biogeochemical Cycling and Sediment Ecology, Kluwer Academic Publishers, Dordrecht, pp. 105-124. https://doi.org/10.1007/978-94-011-4649-4_5

Arntz W.E., Gallardo V.A., Gutiérrez D., et al. 2006. El Niño and similar perturbation effects on the benthos of the Humboldt, California, and Benguela Current upwelling ecosystems. Adv. Geosci. 6: 243-265. https://doi.org/10.5194/adgeo-6-243-2006

Barry J.P. 1988. Hydrographic patterns in McMurdo Sound, Antarctica and their relationship to local benthic communities. Polar Biol. 8: 377-391. https://doi.org/10.1007/BF00442029

Berasategui A.D., Menu Marque S., Gómez-Erache M., et al. 2006. Copepod assemblages in a highly complex hydrographic region. Est. Coast. Shelf Sci. 66: 483-492. https://doi.org/10.1016/j.ecss.2005.09.014

Bhaskar P.V., Bhosle N.B. 2005. Microbial extracellular polymeric substances in marine biogeochemical processes. Curr. Sci. 88: 45-53.

Bidle K., Azam F. 1999. Accelerated dissolution of diatom silica by marine bacterial assemblages. Nature 397: 508-512. https://doi.org/10.1038/17351

Billet D.S.M., Lampitt R.S., Rice A.L., et al. 1983. Seasonal sedimentation of phytoplankton to the deep-sea benthos. Nature 302: 520-522. https://doi.org/10.1038/302520a0

Boero F., Belmonte G., Fanelli G., et al. 1996. The continuity of living matter and the discontinuities of its constituents: do plankton and benthos really exist? Trends Ecol. Evol. 11: 177-180. https://doi.org/10.1016/0169-5347(96)20007-2

Bonsdorff E., Blomqvist E.M. 1993. Biotic coupling on shallow water soft bottoms - Examples from the Northern Baltic Sea. Oceanogr. Mar. Biol. Ann. Rev. 31: 153-176.

Brey T., Gerdes D. 1998. High Antarctic macrobenthic community production. J. Exp. Mar. Biol. Ecol. 231: 191-200. https://doi.org/10.1016/S0022-0981(98)00060-4

Buesseler K.O., Andrews J.E., Pike S.M., et al. 2005. Particle export during the Southern Ocean Iron Experiment (SOFeX). Limnol. Oceanogr. 50: 311-327. https://doi.org/10.4319/lo.2005.50.1.0311

Caron G., Michel C., Gosselin M. 2004. Seasonal contributions of phytoplankton and faecal pellets to the organic carbon sinking flux in the North Water (northern Baffin Bay). Mar. Ecol. Prog. Ser. 283: 1-13. https://doi.org/10.3354/meps283001

Charpy-Roubaud C., Sournia A. 1990. The comparative estimation of phytoplanktonic, microphytobenthic and macrophytobenthic primary production in the oceans. Mar. Micro. Food Webs 4: 31-57.

Clarke A., Tyler P.A. 2008. Adult Antarctic krill feeding at abyssal depths. Curr. Biol. 18: 282-285. https://doi.org/10.1016/j.cub.2008.01.059 PMid:18302926

Daly M., Rack F., Zook R. 2013. Edwardsiella andrillae, a new species of sea anemone from Antarctic Ice. PloS ONE 8: e83476. https://doi.org/10.1371/journal.pone.0083476 PMid:24349517 PMCid:PMC3859642

Davoult D., Gounin F., Richard A. 1990. Dynamique et reproduction de la population d'Ophiothrix fragilis (Abildgaard) du détroit du Pas-de Calais (Manche orientale). J. Exp. Mar. Biol. Ecol. 138: 201-216. https://doi.org/10.1016/0022-0981(90)90167-B

Del Giorgio P.A., Duarte C.M. 2002. Respiration in the open ocean. Nature 420: 379-384. https://doi.org/10.1038/nature01165 PMid:12459775

Eckman J.E., Duggins D.O. 1993. Effects of flow speed on growth of benthic suspension feeders. Biol. Bull. 185: 28-41. https://doi.org/10.2307/1542128 PMid:29300603

Fagan W., Fortin M-J., Soykan C. 2003. Integrating edge detection and dynamic modeling in quantitative analyses of ecological boundaries. BioScience 53: 730-738. https://doi.org/10.1641/0006-3568(2003)053[0730:IEDADM]2.0.CO;2

Fanning K.A., Carter K.L., Betzer P.R. 1982. Sediment resuspension by coastal waters: a potential mechanism for nutrient re-cycling on the ocean's margins. Deep-Sea Res. Part A 29: 953-965. https://doi.org/10.1016/0198-0149(82)90020-6

Fukuchi M., Sasaki H., Hattori H., et al. 1993. Temporal variability of particulate flux in the northern Bering Sea. Cont. Shelf Res. 13: 693-704. https://doi.org/10.1016/0278-4343(93)90100-C

Gili J.M., Coma R. 1998. Benthic suspensión feeders: their paramount role in littoral marine food webs. Trends Ecol. Evol. 13: 316-321. https://doi.org/10.1016/S0169-5347(98)01365-2

Gili J.M., Coma R., Orejas C., et al. 2001. Are Antarctic suspension-feeding communities different from those elsewhere in the world? Polar Biol. 24: 473-485. https://doi.org/10.1007/s003000100257

Gili J.M., Rossi S., Pagès F., et al. 2006. A new trophic link between the pelagic and benthic systems on the Antarctic shelf. Mar. Ecol. Prog. Ser. 322: 43-49. https://doi.org/10.3354/meps322043

Gooday A.J. 2002. Biological responses to seasonally varying fluxes of organic matter to the ocean floor: A review. J. Oceanogr. 58: 305-332. https://doi.org/10.1023/A:1015865826379

Graf G. 1992. Benthic-pelagic coupling: a benthic view. Oceanogr. Mar. Biol. Annu. Rev. 30: 149-190.

Graf G., Rosenberg R. 1997. Bioresuspension and biodeposition: a review. J. Mar. Syst. 11: 269-278. https://doi.org/10.1016/S0924-7963(96)00126-1

Grebmeier J.M., Barry J.P. 1991. The influence of oceanographic processes on pelagic-benthic coupling in polar regions: a benthic perspective. J. Mar. Syst. 2: 495-518. https://doi.org/10.1016/0924-7963(91)90049-Z

Grebmeier J.M., McRoy C.P. 1989. Pelagic-benthic coupling on the shelf of the northern Bering and Chukchi Seas -III. Benthic food supply and carbon cycling. Mar. Ecol. Prog. Ser. 53: 253-268. https://doi.org/10.3354/meps051253

Gutt J., Siegel V. 1994. Benthopelagic aggregations of krill (Euphausia superba) on the deeper shelf of the Weddell Sea (Antarctic). Deep Sea Res. I 41: 169-178. https://doi.org/10.1016/0967-0637(94)90031-0

Harrold C., Light K., Lisin S. 1998. Organic enrichment of submarine-canyon and continental-shelf benthic communities by macroalgal drift imported from nearshore kelp forest. Limnol. Oceanogr. 43: 669-678. https://doi.org/10.4319/lo.1998.43.4.0669

Henson S.A., Sanders R., Holeton C., et al. 2006. Timing of nutrient depletion, diatom dominance and a lower-boundary estimate of export production for Irminger Basin, North Atlantic. Mar. Ecol. Prog. Ser. 313: 73-84. https://doi.org/10.3354/meps313073

Hodges B.A., Rudnick D.L. 2004. Simple models of steady deep maxima in chlorophyll and biomass. Deep-Sea Res. Part I 51: 999-1015. https://doi.org/10.1016/j.dsr.2004.02.009

Huthnance J.M. 1995. Circulation, exchange and water masses at the ocean margin: the role of physical processes at the shelf edge. Progr. Oceanogr. 35: 353-431. https://doi.org/10.1016/0079-6611(95)80003-C

Ichino M.C., Clark M.R., Drazen J.C., et al. 2015. The distribution of benthic biomass in hadal trenches: a modelling approach to investigate the effect of vertical and lateral organic matter transport to the seafloor. Deep-Sea Res. Part I 100: 21-33. https://doi.org/10.1016/j.dsr.2015.01.010

Isla E., Gerdes D., Palanques A., et al. 2006. Particle flux and tides near the continental ice edge on the eastern Weddell Sea shelf. Deep-Sea Res. Part II 53: 866-874. https://doi.org/10.1016/j.dsr2.2006.02.010

Johnson A.S. 1988. Hydrodynamic study of the functional morphology of the benthic suspension feeder Phoronopsis viridis (Phoronida). Mar. Biol. 100: 117-126. https://doi.org/10.1007/BF00392961

Josefson A.B., Conley D.J. 1997. Benthic response to a pelagic front. Mar. Ecol. Prog. Ser. 147: 49-62. https://doi.org/10.3354/meps147049

Koski M., Kiorboe T., Takahashi K. 2005. Benthic life in the pelagic: Aggregate encounter and degradation rates by pelagic harpacticoid copepods. Limnol. Ocaenogr. 50: 1254-1263. https://doi.org/10.4319/lo.2005.50.4.1254

Larsena P.S., Riisgård H.U. 1996. Mixing generated by benthic filter-feeders: a diffusion model for near-bottom phytoplankton depletion. J. Sea Res. 37: 81-90. https://doi.org/10.1016/S1385-1101(97)00009-9

Levin L.A., Dayton P.V. 2009. Ecological theory and continental margins: where shallow meets deep. Trends Ecol. Evol. 24: 606-617. https://doi.org/10.1016/j.tree.2009.04.012 PMid:19692143

Levin L.A., Gooday A. 2003. The deep Atlantic Ocean. In: Tyler P.A., Ecosystems of the World, Vol. 28, Ecosystems of the deep oceans. Elsevier, Amsterdam, pp. 111-178.

Longhurst A.R. 1985. Relationship between diversity and the vertical structure of the upper ocean. Deep-Sea Res. Part A 32: 1535-1570. https://doi.org/10.1016/0198-0149(85)90102-5

Lovvorn J.R., Cooper L.W., Brooks M.L., et al. 2005. Organic matter pathways to zooplankton and benthos under pack ice in late winter and open water in late summer in the north-central Bering Sea. Mar. Ecol. Prog. Ser. 291: 135-150. https://doi.org/10.3354/meps291135

MacIntyre S., Alldredge A.L., Gotschalk C.C. 1995. Accumulation of Marine Snow at Density Discontinuities in the Water Column. Limnol. Oceanogr. 40: 449-468. https://doi.org/10.4319/lo.1995.40.3.0449

Magnusson J.J., Harrington C.L., Stewart D.J., et al. 1981. Responses of macrofauna to short-term dynamics of a gulf stream front on the continental shelf. In: Richards F.A., Coastal Upwelling I, Coastal and Estuarine Sciences, American Geophysical Union, pp. 441-448. https://doi.org/10.1029/CO001p0441

Margalef R. 1968. Perspectives in Ecological Theory. The University of Chicago, Chicago.

Margalef R. 1974. Ecología. Ed. Omega, Barcelona.

Margalef R. 1991. Teoria de los Sistemas Ecológicos. Publicacions de la Universitat de Barcelona. Barcelona.

Margalef R. 1997. Our Biosphere. Excellence in Ecology. Ecology Institute. Oldendorf.

Margalef R. 2001. The top layers of water bodies, a most important although relatively neglected piece of the biosphere plumbing. In: Gili J.-M., Pretus J.M., Packard T.T. (eds) A Marine Science Odyssey into the 21st Century. Sci. Mar. 65 (Suppl. 2): 207-213. https://doi.org/10.3989/scimar.2001.65s2207

McGillicuddy Jr. D.J. 2016. Mechanisms of physical-biological-biogeochemical interaction at the oceanic mesoscale. Annu. Rev. Mar. Sci. 8: 125-159. https://doi.org/10.1146/annurev-marine-010814-015606 PMid:26359818

McPhee-Shaw E. 2006. Boundary-interior Exchange: Reviewing the idea that internal-wave mixing enhances lateral dispersal near continental margins. Deep-Sea Res. Part II 53: 42-59. https://doi.org/10.1016/j.dsr2.2005.10.018

Meysman F.J.R., Middelburg J.J., Heip C.H.R. 2006. Bioturbation: a fresh look at Darwin's last idea. Trends Ecol. Evol. 21: 691-694. https://doi.org/10.1016/j.tree.2006.08.002 PMid:16901581

Nilsen M., Pedersen T., Nilssen E.M. 2006. Macrobenthic biomass, productivity (P/B) and production in a high-latitude ecosystem, North Norway. Mar. Ecol. Prog. Ser. 32: 67-77. https://doi.org/10.3354/meps321067

Olli K., Riser C.W., Wassmann P., et al. 2002. Seasonal variation in vertical flux of biogenic matter in the marginal ice zone and the central Barents Sea. J. Mar. Syst. 38: 189-204. https://doi.org/10.1016/S0924-7963(02)00177-X

Orejas C., Gili J.-M., López-González P.J., et al. 2001. Feeding strategies and diet composition of four species of Antarctic cnidarian species. Polar Biol. 24: 620-627. https://doi.org/10.1007/s003000100272

Parrish C.C., Thompson R.J., Deibel D. 2005. Lipid classes and fatty acids in plankton and settling matter during the spring bloom in a cold ocean coastal environment. Mar. Ecol. Prog. Ser. 286: 57-68. https://doi.org/10.3354/meps286057

Prat N. 2015. Ramon Margalef 1919-2004. Una obra ingente no totalmente bien reconocida. Ecosistemas 24: 97-103. https://doi.org/10.7818/ECOS.2015.24-1.17

Prat N., Ros J. D., Peters F. 2015. Biografía científica del profesor Ramon Margalef López. In: Prat N., Ros, J. D., Peters F. (eds) Ramon Margalef, ecólogo de la biosfera. Una biografía científica, Universitat de Barcelona, Fundació Agbar, pp. 19-99.

Puig P., Palanques A., Martín J. 2014. Contemporary sediment-transport processes in submarine canyons. Ann. Rev. Mar. Sci. 6: 53-77. https://doi.org/10.1146/annurev-marine-010213-135037 PMid:23937169

Reise K. 2002. Sediment mediated species interactions in coastal waters. J. Sea Res. 48: 127-141. https://doi.org/10.1016/S1385-1101(02)00150-8

Richter C., Wunsch M., Rasheed M., et al. 2001. Endoscopic exploration of Red Sea coral reefs reveals dense populations of cavity-dwelling sponges. Nature 413: 726-730. https://doi.org/10.1038/35099547 PMid:11607030

Ritzrau W., Graf G. 1992. Increase of microbial biomass in the benthic turbidity zone of Kiel Bight after resuspension by a storm event. Limnol. Oceanogr. 37: 1081-1086. https://doi.org/10.4319/lo.1992.37.5.1081

Ritzrau W., Thomsen L. 1997. Spatial distribution of particle composition and microbial activity in the benthic boundary layer (BBL) of the Northeast Water Polynya. J. Mar Syst. 10: 415-428. https://doi.org/10.1016/S0924-7963(96)00073-5

Ritzrau W., Thomsen L., Lara R.J., et al. 1997. Enhanced microbial utilisation of dissolved aminoacids indicates rapid modification of organic matter in the benthic boundary layer. Mar. Ecol. Prog. Ser. 156: 43-50. https://doi.org/10.3354/meps156043

Sabatés A., Masó M. 1990. Effect of a shelf-slope front on the spatial distribution of mesopelagic fish larvae in the western Mediterranean. Deep-Sea Res. Part A 37: 1085-1098. https://doi.org/10.1016/0198-0149(90)90052-W

Sheridan C.C., Lee C., Wakeham S.G., et al. 2002. Suspended particle organic composition and cycling in surface and midwaters of the ecuatorial Pacific Ocean. Deep-Sea Res. Part I 49: 1983-2008. https://doi.org/10.1016/S0967-0637(02)00118-8

Schewe I., Soltwedel T. 2003. Benthic response to ice-edge-induced particle flux in the Arctic Ocean. Polar Biol. 26: 610-620. https://doi.org/10.1007/s00300-003-0526-8

Schonberg S.V., Clarke J.T., Dunton K.H. 2014. Distribution, abundance, biomass and diversity of benthic infauna in the Northeast Chukchi Sea, Alaska: Relation to environmental variables and marine mammals. Deep Sea Research Part II 102: 144-163. https://doi.org/10.1016/j.dsr2.2013.11.004

Sheridan C.C., Lee C., Wakeham S.G., et al. 2002. Suspended particle organic composition and cycling in surface and midwaters of the equatorial Pacific Ocean. Deep-Sea Res. Part I 49: 1983-2008. https://doi.org/10.1016/S0967-0637(02)00118-8

Sherrell R.M., Field M.P., Gao Y. 1998. Temporal variability of suspended mass and composition in the Northeast Pacific water column: relationships to sinking flux and lateral advection. Deep-Sea Res. Part II 45: 733-761. https://doi.org/10.1016/S0967-0645(97)00100-8

Sims D.W., Quayle V.A. 1998. Selective foraging behaviour of basking sharks on zooplancton in a small-scale front. Nature 393: 460-464. https://doi.org/10.1038/30959

Slobodkin L.B. 1961. Growth and Regulation of animal Populations. Holt, Rinehart & Winston, New York.

Smetacek V. 1984. The supply of food to the benthos. In: Fasham M.J.R. (ed.) Flows of energy and materials in marine ecosystems. Theory and practice. Plenum Press, pp. 517-547. https://doi.org/10.1007/978-1-4757-0387-0_20

Smith C.R., Mincks S., DeMaster D.J. 2006. A synthesis of bentho-pelagic coupling on the Antarctic shelf: Food banks, ecosystem inertia and global climate change. Deep-Sea Res. Part II 53: 875-894. https://doi.org/10.1016/j.dsr2.2006.02.001

Springer A.M., McRoy C.P., Flint M.V. 1996. The Bering Sea Green Belt: shelf-edge processes and ecosystem production. Fish. Oceanogr., 5(3-4): 205-223. https://doi.org/10.1111/j.1365-2419.1996.tb00118.x

Stemmann L., Jackson G.A., Gorsky G. 2004. A vertical model of particle size distributions and fluxes in the midwater column that includes biological and physical processes - Part II: application to a three year survey in the NW Mediterranean Sea. Deep-Sea Res. Part I 51: 885-908. https://doi.org/10.1016/j.dsr.2004.03.002

Tenore K.R., Zajac R.N., Terwin J., et al. 2006. Characterizing the role benthos plays in large coastal seas and estuaries: A modular approach. J. Exp. Mar. Biol. Ecol. 330: 392-402. https://doi.org/10.1016/j.jembe.2005.12.042

Thiem Ø., Ravagnan E-. Fossa J.H., et al. 2006. Food supply mechanisms for cold-water corals along a continental shelf edge. J. Mar. Syst. 60: 207-219. https://doi.org/10.1016/j.jmarsys.2005.12.004

Thomsen L. 1999. Processes in the benthic boundary layer at continental margins and their implication for benthic carbon cycle. J. Sea Res. 41: 73-86. https://doi.org/10.1016/S1385-1101(98)00039-2

Thomsen L., Van Weering C.E. Tj. 1998. Spatial and temporal variability of particulate matter in the benthic boundary layer at the N.W. European continental margin (Goban Spur). Progr. Oceanogr. 42: 61-76. https://doi.org/10.1016/S0079-6611(98)00028-7

Turley C. 2000. Bacteria in the cold deep-sea benthic boundary layer and sediment-water interface of the NE Atlantic. Microbiol. Ecol. 33: 89-99. https://doi.org/10.1016/S0168-6496(00)00058-1

Valiela I. 1995. Marine Ecological Processes. Springer-Verlag, New-York. https://doi.org/10.1007/978-1-4757-4125-4

Van der Loeff M.M.R., Meyer R., Rudels B., et al. 2002. Resuspension and particle transport in the benthic nepheloid layer in and near Fram Strait in relation to faunal abundance and 234Th depletion. Deep-Sea Res. Part I 49: 1941-1958. https://doi.org/10.1016/S0967-0637(02)00113-9

Van Leeuwen C.G. 1966. A theoretical approach to pattern and process in vegetation. Wentia 15: 25-46. https://doi.org/10.1111/j.1438-8677.1966.tb00019.x

Vendrell B., Gili J.M., Gasol J.M., et al. 2005. How do benthic antarctic communities impact near-bottom water layer properties? ASLO Summer Meeting 2005, 18-24 Junio, Santiago de Compostela.

Vogel S. 1994. Life in moving fluids: The physical biology of flow. University Press Princeton.

Wainwright S.C. 1990. Sediment-to-water fluxes of particulate material and microbes by resuspension and their contribution to the planktonic food web. Mar. Ecol. Prog. Ser. 62: 271-281. https://doi.org/10.3354/meps062271

Wildish D., Kristmanson D. 1997. Benthic suspension feeders and flow. Cambridge University Press, Cambridge. https://doi.org/10.1017/CBO9780511529894

Witte U. 1996. Seasonal reproduction in deep-sea sponges - triggered by vertical particle flux? Mar. Biol. 124: 571-581. https://doi.org/10.1007/BF00351038

Witte U., Brattegard T., Graf G., et al. 1997. Particle capture and deposition by deep-sea sponges from the Norwegian-Greenland Sea. Mar. Ecol. Prog. Ser. 154: 241-252. https://doi.org/10.3354/meps154241

Witte U., Aberle N., Sand M., et al. 2003. Rapid response of a deep-sea benthic community to POM enrichment: an in situ experimental study. Mar. Ecol. Prog. Ser. 251: 27-36. https://doi.org/10.3354/meps251027

Wotton R.S. 1994. Particulate and disolved organic matter as food. In: Wotton R.S. (eds), The biology of Particles in aquatic systems, Lewis Publishers, Boca Raton, pp. 235-288. https://doi.org/10.1201/9781003070146-10

Zabala M., Ballesteros E. 1989. Surface-dependent strategies and energy flux in benthic marine communities or, why corals do not exist in the Mediterranean. Sci. Mar. 53: 3-17.