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Comparative analyses of oceanic ecosystems require an objective framework to define coherent study regions and scale the patterns and processes observed within them. We applied the hierarchical patch mosaic paradigm of landscape ecology to the study of the seasonal variability of the North Pacific to facilitate comparative analysis between pelagic ecosystems and provide spatiotemporal context for Eulerian time-series studies. Using 13-year climatologies of sea surface temperature (SST), photosynthetically active radiation (PAR), and chlorophyll a (chl-a), we classified seascapes in environmental space that were monthly-resolved, dynamic and nested in space and time. To test the assumption that seascapes represent coherent regions with unique biogeochemical function and to determine the hierarchical scale that best characterized variance in biogeochemical parameters, independent data sets were analyzed across seascapes using analysis of variance (ANOVA), nested-ANOVA and multiple linear regression (MLR) analyses. We also compared the classification efficiency (as defined by the ANOVA F-statistic) of resultant dynamic seascapes to a commonly-used static classification system. Variance of nutrients and net primary productivity (NPP) were well characterized in the first two levels of hierarchy of eight seascapes nested within three superseascapes (R2=0.5-0.7). Dynamic boundaries at this level resulted in a nearly 2-fold increase in classification efficiency over static boundaries. MLR analyses revealed differential forcing on pCO2 across seascapes and hierarchical levels and a 33% reduction in mean model error with increased partitioning (from 18.5μatm to 12.0μatm pCO2). Importantly, the empirical influence of seasonality was minor across seascapes at all hierarchical levels, suggesting that seascape partitioning minimizes the effect of non-hydrographic variables. As part of the emerging field of pelagic seascape ecology, this effort provides an improved means of monitoring and comparing oceanographic biophysical dynamics and an objective, quantitative basis by which to scale data from local experiments and observations to regional and global biogeochemical cycles. © 2013.


Documento: Artículo
Título:Hierarchical and dynamic seascapes: A quantitative framework for scaling pelagic biogeochemistry and ecology
Autor:Kavanaugh, M.T.; Hales, B.; Saraceno, M.; Spitz, Y.H.; White, A.E.; Letelier, R.M.
Filiación:Department of Marine Chemistry and Geochemistry, Woods Hole Oceanographic Institution, Woods Hole, MA, United States
College of Earth, Ocean, and Atmospheric Sciences, Oregon State University, 104 CEOAS Administration Building, Corvallis, OR, United States
Departamento de Ciencias de las Atmosfera y los Oceanos, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina
Centro de Investigaciones del Mar y la Atmósfera (CIMA/CONICET-UBA), UMI IFAECI/CNRS, Buenos Aires, Argentina
Idioma: Inglés
Palabras clave:Biogeochemistry; Models; North Pacific; Pelagic environment; Seascapes; Seasonal variations; Multiple linear regressions; North Pacific; Pelagic biogeochemistries; Pelagic environment; Photosynthetically active radiation; Sea surface temperature (SST); Seascapes; Seasonal variation; Analysis of variance (ANOVA); Ecosystems; Linear regression; Models; Oceanography; Plants (botany); Biogeochemistry; biogeochemistry; biophysics; carbon dioxide; chlorophyll a; hierarchical system; hypothesis testing; marine ecosystem; net primary production; pelagic ecosystem; photosynthetically active radiation; quantitative analysis; sea surface temperature; seasonal variation; spatiotemporal analysis; variance analysis; Pacific Ocean; Pacific Ocean (North)
Página de inicio:291
Página de fin:304
Título revista:Progress in Oceanography
Título revista abreviado:Prog. Oceanogr.


  • Anouar, F., Badran, F., Thiria, S., Probabilistic self-organizing map and radial basis function networks (1998) Neurocomputing, 20, pp. 83-96
  • Belgrano, A., Lima, M., Stenseth, N.C., Non-linear dynamics in marine-phytoplankton population systems: emergent properties of complex marine systems: a macroecological perspective (2004) Marine Ecology Progress Series, 273, pp. 281-289
  • Beaugrand, G., Reid, P.C., Ibañez, F., Biodiversity of North Atlantic and North Sea calanoid copepods (2000) Marine Ecology Progress Series, 204, pp. 299-303
  • Bograd, S.J., Foley, D.G., Schwing, F.B., Wilson, C., Laurs, R.M., Polovina, J.J., Howell, E.A., Brainard, R.E., On the seasonal and interannual migrations of the transition zone chlorophyll front (2004) Geophysical Research Letters, 31, pp. L17204
  • Brander, K., Impacts of climate change on fisheries (2010) Journal of Marine Systems, 79 (3), pp. 389-402
  • Cole, J.J., Communication between terrestrial and marine ecologists: loud, sometimes abrasive, but healthy and occasionally useful: bridging the gap between aquatic and terrestrial ecology (2005) Marine Ecology Progress Series, 304, pp. 272-274
  • Deutsch, C., Gruber, N., Key, R.M., Sarmiento, J.L., Ganachaud, A., Denitrification and N2 fixation in the Pacific Ocean (2001) Global Biogeochemical Cycles, 15, pp. 483-506
  • Devred, E., Sathyendranath, S., Platt, T., Delineation of ecological provinces using ocean colour radiometry (2007) Marine Ecology Progress Series, 346, pp. 1-13
  • Devred, E., Sathyendranath, S., Platt, T., Decadal changes in ecological provinces of the Northwest Atlantic Ocean revealed by satellite observations (2009) Geophysical Research Letters, 36, pp. L19607
  • Doney, S.C., Ruckelshaus, M., Emmett Duffy, J., Barry, J.P., Chan, F., English, C.A., Galindo, H.M., Talley, L.D., Climate change impacts on marine ecosystems (2012) Annual Review of Marine Science, 4, pp. 11-37
  • Dutkiewicz, S., Ward, B.A., Monteiro, F., Follows, M.J., Interconnection between nitrogen fixers and iron in the Pacfic Ocean: theory and numerical model (2012) Global Biogeochemical Cycles, 26, pp. GB1012
  • Evans, W., Strutton, P.G., Chavez, F.P., Impact of tropical instability waves on nutrient and chlorophyll distributions in the equatorial Pacific (2009) Deep Sea Research Part I: Oceanographic Research Papers, 56, pp. 178-188
  • Fortin, M.-J., Dale, M.R.T., (2005) Spatial Analysis: A Guide for Ecologists, , Cambridge University Press
  • Friedrich, T., Oschlies, A., Neural network-based estimates of North Atlantic surface pCO 2 from satellite data: a methodological study (2009) Journal of Geophysical Research, 114, pp. C03020
  • Gillson, L., Landscapes in time and space (2009) Landscape Ecology, 24 (2), pp. 149-155
  • Gruber, N., Warming up, turning sour, losing breath: ocean biogeochemistry under global change (2011) Philosophical Transactions of the Royal Society A, 369, pp. 1980-1996
  • Hales, B., Strutton, P.G., Saraceno, M., Letelier, R., Takahashi, T., Feely, R., Sabine, C., Chavez, F., Satellite-based prediction of pCO2 in coastal waters of the eastern North Pacific (2012) Progress in Oceanography, 103, pp. 1-15
  • Hardman-Mountford, N.J., Hirata, T., Richardson, K.A., Aiken, J., An objective methodology for the classification of ecological pattern into biomes and provinces for the pelagic ocean (2008) Remote Sensing of Environment, 112, pp. 3341-3352
  • Howard, E., Emerson, S., Bushinsky, S., Stump, C., The role of net community production in air-sea carbon fluxes at the North Pacific subarctic-subtropical boundary region (2010) Limnology and Oceanography, 55 (6), pp. 2585-2596
  • Hsieh, C., Glaser, S.M., Lucas, A.J., Sugihara, G., Distinguishing random environmental fluctuations from ecological catastrophes for the North Pacific Ocean (2005) Nature, 435, pp. 336-340
  • Irwin, A.J., Oliver, M.J., Are ocean deserts getting larger α (2009) Geophysical Research Letters, 36, pp. 1-5
  • Jain, A.K., Dubes, R.C., Chen, C.C., Bootstrap techniques for error estimation (1987) IEEE Transactions on Pattern Analysis and Machine Intelligence, 9, pp. 628-633
  • Jassby, A.D., Platt, T., Mathematical formulation of the relationship between photosynthesis and light for phytoplankton (1976) Limnology and Oceanography, 21, pp. 540-547
  • Juranek, L.W., Quay, P.D., Feely, R.A., Lockwood, D., Karl, D.M., Church, M.J., Biological production in the NE Pacific and its influence on air-sea CO2 flux: Evidence from dissolved oxygen isotopes and O2/Ar (2012) Journal of Geophysical Research: Oceans (1978-2012), 117 (C5)
  • Kavanaugh, M.T., Holtgrieve, G.W., Baulch, H., Brum, J.R., Cuvelier, M.L., Filstrup, C.T., Nickols, K.J., Small, G.E., A salty divide in ASLOα (2013) Limnology and Oceanography Bulletin, 22 (2), pp. 34-37
  • Karl, D.M., Church, M.J., Dore, J.E., Letelier, R.M., Mahaffey, C., Predictable and efficient carbon sequestration in the North Pacific Ocean supported by symbiotic nitrogen fixation (2012) Proceedings of the National Academy of Sciences of the United States of America, 109, pp. 1842-1849
  • Karl, D.M., Letelier, R.M., Seascape microbial ecology: Habitat structure, biodiversity and ecosystem function (2009) Guide to Ecology, pp. 488-500. , Princeton University Press, Princeton, New Jersey, S.A. Levin (Ed.)
  • Kotliar, N.B., Wiens, J.A., Multiple scales of patchiness and patch structure: a hierarchical framework for the study of heterogeneity (1990) Oikos, 59, pp. 253-260
  • Lachkar, Z., Gruber, N., A comparative study of biological production in eastern boundary upwelling systems using an artificial neural network (2012) Biogeosciences, 9, pp. 293-308
  • Landschützer, P., Gruber, N., Bakker, D.C.E., Schuster, U., Nakaoka, S., Payne, M.R., Sasse, T., Zeng, J., A neural network-based estimate of the seasonal to inter-annual variability of the Atlantic Ocean carbon sink (2013) Biogeosciences Discussion, 10, pp. 8799-8849
  • Letelier, R.M., Bidigare, R.R., Hebel, D.V., Ondrusek, M., Winn, C.D., Karl, D.M., Temporal variability of phytoplankton community structure based on pigment analysis (1993) Limnology and Oceanography, 38, pp. 1420-1437
  • Levin, S.A., Whitfield, M., Patchiness in marine and terrestrial systems: from individuals to populations [and Discussion] (1994) Philosophical Transactions of the Royal Society B: Biological Sciences, 343, pp. 99-103
  • Litzow, M.A., Ciannelli, L., Oscillating trophic control induces community reorganization in a marine ecosystem (2007) Ecology Letters, 10, pp. 1124-1134
  • Longhurst, A.R., (1998) Ecological Geography of the Sea, , Elsevier Press, London UK, 2007 (second ed.)
  • Lockwood, D., Quay, P.D., Kavanaugh, M.T., Juranek, L.W., Feely, R., Influence of net community production on air-sea CO2 flux in the Northeast Pacific (2012) Global Biogeochemical Cycles, 26, pp. GB4010
  • Lubchenco, J., Petes, L.E., The interconnected biosphere: science at the ocean's tipping points (2010) Oceanography, 23, pp. 115-129
  • Luo, Y.-W., Doney, S.C., Anderson, L.A., Benavides, M., Berman-Frank, I., Bode, A., Bonnet, S., Zehr, J.P., Database for diazotrophs in global ocean: abundances, biomass and nitrogen fixation rates (2012) Earth System Science Data, 4, pp. 47-73
  • Luo, Y.-W., Lima, I.D., Karl, D.M., Doney, S.C., Feely, R., Data-based assessment of environmental controls on global marine nitrogen fixation (2013) Biogeosciences Discussion, 10, pp. 7367-7412
  • McCune, B., Grace, J.B., Urban, D.L., (2002) Analysis of Ecological Communities, 28. , MjM Software Design, Gleneden Beach, Oregon
  • Mitchell, J.G., Yamazaki, H., Seuront, L., Wolk, F., Hua, L., Phytoplankton patch patterns: seascape anatomy in a turbulent ocean (2008) Journal of Marine Systems, 69, pp. 247-253
  • Murawski, S.A., Steele, J.H., Taylor, P., Fogarty, M.J., Sissenwine, M.P., Ford, M., Suchman, C., Why compare marine ecosystemsα (2010) ICES Journal of Marine Science, 67, pp. 1-9
  • Oliver, M.J., Irwin, A.J., Objective global ocean biogeographic provinces (2008) Geophysical Research Letters, 35 (15), pp. L15601
  • O'Neill, R.V., Gardner, R.H., Turner, M.G., A hierarchical neutral model for landscape analysis (1992) Landscape Ecology, 7, pp. 55-61
  • Park, G.-H., Wannikhof, R., Doney, S.C., Takahashi, T., Lee, K., Feely, R.A., Sabine, C.L., Lima, I.D., Variability of global net sea-air CO2 fluxes over the last three decades using empirical relationships (2010) Tellus B, 62, pp. 352-368
  • Platt, T., Sathyendranath, S., Spatial structure of pelagic ecosystem processes in the global ocean (1999) Ecosystems, 2, pp. 384-394
  • Platt, T., Sathyendranath, S., Ecological indicators for the pelagic zone of the ocean from remote sensing (2008) Remote Sensing of Environment, 112, pp. 3426-3436
  • Polovina, J.J., Howell, E., Kobayashi, D.R., Seki, M.P., The transition zone chlorophyll front, a dynamic global feature defining migration and forage habitat for marine resources (2001) Progress in Oceanography, 49, pp. 469-483
  • Polovina, J.J., Dunne, J.P., Woodworth, P.A., Howell, E.A., Projected expansion of the subtropical biome and contraction of the temperate and equatorial upwelling biomes in the North Pacific under global warming (2011) ICES Journal of Marine Science: Journal du Conseil, 68 (6), pp. 986-995
  • Richardson, A., Risien, C., Shillington, F., Using self-organizing maps to identify patterns in satellite imagery (2003) Progress in Oceanography, 59, pp. 223-239
  • Saraceno, M., Provost, C., Lebbah, M., Biophysical regions identification using an artificial neuronal network: a case study in the South Western Atlantic (2006) Advances in Space Research, 37, pp. 793-805
  • Siegel, D.A., Westberry, T.K., O'Brien, M.C., Nelson, N.B., Michaels, A.F., Morrison, J.R., Scott, A., Hammer, M.A., Bio-optical modeling of primary production on regional scales: the Bermuda BioOptics project (2001) Deep-Sea Research Part IIi-Topical Studies in Oceanography, 48, pp. 1865-1896
  • Siegel, D.A., Behrenfeld, M.J., Maritorena, S., McClain, C.R., Antoine, D., Bailey, S.W., Bontempi, P.S., Regional to global assessments of phytoplankton dynamics from the SeaWiFS mission (2013) Remote Sensing of Environment, 135, pp. 77-91
  • Somerville, M., (1853), Physical Geography. Lea & Blanchard. Michigan Historical Reprint Series, University of Michigan, Ann Arbor, MISteele, J.H., The ocean landscapeα (1989) Landscape Ecology, 3, pp. 185-192
  • Steele, J.H., Can ecological theory cross the land-sea boundaryα (1991) Journal of Theoretical Biology, 153, pp. 425-436
  • Steele, J.H., Henderson, E.W., A simple model for plankton patchiness (1992) Journal of Plankton Research, 14, pp. 1397-1403
  • Takahashi, T., Sutherland, S.C., Sweeney, C., Poisson, A., Metzl, N., Tilbrook, B., Bates, N., Nojiri, Y., Global sea-air CO2 flux based on climatological surface ocean pCO(2), and seasonal biological and temperature effects (2002) Deep-Sea Research Part II-Topical Studies in Oceanography, 49, pp. 1601-1622
  • Takahashi, T., Sutherland, S.C., Wanninkhof, R., Sweeney, C., Feely, R.A., Chipman, D.W., Hales, B., de Baar, H.J.W., Climatological mean and decadal change in surface ocean pCO(2), and net sea-air CO2 flux over the global oceans (2009) Deep-Sea Research Part II: Topical Studies in Oceanography, 56, pp. 554-577
  • Telszewski, M., Chazottes, A., Schuster, U., Watson, A.J., Moulin, C., Bakker, D.C.E., González-Dávila, M., Wanninkhof, R., Estimating the monthlypCO2distribution in the North Atlantic using a self-organizing neural network (2009) Biogeosciences, 6, pp. 1405-1421
  • Turner, M.G., Landscape ecology: what is the state of the scienceα (2005) Annual Review of Ecology Evolution and Systematics, 36, pp. 319-344
  • Turner, M.G., Gardner, R.H., O'Neill, R.V., (2001) Landscape Ecology in Theory and Practice: Pattern and Process, , Springer-Verlag, New York
  • Troll, C., Die geographische Landschaft und ihre Erforschung. Studium Generale 3(4/5):163-181 (1950) Foundation Papers in Landscape Ecology, , Columbia University Press, New York, J.A. Wiens, M.R. Moss, M.G. Turner, D.J. Mladenoff (Eds.)
  • Venrick, E.L., The Distribution and Significance of Richelia intracellularis Schmidt in the North Pacific Central Gyre (1974) Limnology and Oceanography, 19, pp. 437-445
  • Vichi, M., Allen, J.I., Masina, S., Hardman-Mountford, N.J., The emergence of ocean biogeochemical provinces: a quantitative assessment and a diagnostic for model evaluation (2011) Global Biogeochemical Cycles, 25, pp. GB2005
  • Villareal, T.A., Nitrogen-fixation by the cyanobacterial symbiont of the diatom genus Hemiaulus (1991) Marine Ecology Progress Series, 76 (2), pp. 201-204
  • Ward, J.H., Hierarchical grouping to optimize an objective function (1963) Journal of the American Statistical Association, 58, pp. 236-244
  • Weber, T.S., Deutsch, C., Ocean nutrient ratios governed by plankton biogeography (2010) Nature, 467, pp. 550-554
  • Westberry, T.K., Behrenfeld, M.J., Siegel, D.A., Boss, E., Carbon-based primary productivity modeling with vertically resolved photoacclimation (2008) Global Biogeochemical Cycles, 22, pp. GB2024
  • White, A.E., Spitz, Y.H., Letelier, R.M., What factors are driving summer phytoplankton blooms in the North Pacific Subtropical Gyreα (2007) Journal of Geophysical Research, 112, pp. 1-11
  • Winn, C.D., Campbell, L., Christian, J.R., Letelier, R.M., Hebel, D.V., Dore, J.E., Fujieki, L., Karl, D.M., Seasonal variability in the phytoplankton community of the North Pacific Subtropical Gyre, Global Biogeochem (1995) Cycles, 9, pp. 605-620
  • Wilson, C., Villareal, T.A., Maximenko, N., Bograd, S.J., Montoya, J.P., Schoenbaechler, C.A., Biological and physical forcings of late summer chlorophyll blooms at 30°N in the oligotrophic Pacific (2008) Journal of Marine Systems, 69, pp. 164-176
  • Wu, J., Loucks, O.L., From balance of nature to hierarchical patch dynamics: a paradigm shift in ecology (1995) The Quarterly Review of Biology, 70, pp. 439-466
  • Wu, J., Hierarchy and scaling: extrapolating information along a scaling ladder (1999) Canadian Journal of Remote Sensing, 25 (4), pp. 367-380


---------- APA ----------
Kavanaugh, M.T., Hales, B., Saraceno, M., Spitz, Y.H., White, A.E. & Letelier, R.M. (2014) . Hierarchical and dynamic seascapes: A quantitative framework for scaling pelagic biogeochemistry and ecology. Progress in Oceanography, 120, 291-304.
---------- CHICAGO ----------
Kavanaugh, M.T., Hales, B., Saraceno, M., Spitz, Y.H., White, A.E., Letelier, R.M. "Hierarchical and dynamic seascapes: A quantitative framework for scaling pelagic biogeochemistry and ecology" . Progress in Oceanography 120 (2014) : 291-304.
---------- MLA ----------
Kavanaugh, M.T., Hales, B., Saraceno, M., Spitz, Y.H., White, A.E., Letelier, R.M. "Hierarchical and dynamic seascapes: A quantitative framework for scaling pelagic biogeochemistry and ecology" . Progress in Oceanography, vol. 120, 2014, pp. 291-304.
---------- VANCOUVER ----------
Kavanaugh, M.T., Hales, B., Saraceno, M., Spitz, Y.H., White, A.E., Letelier, R.M. Hierarchical and dynamic seascapes: A quantitative framework for scaling pelagic biogeochemistry and ecology. Prog. Oceanogr. 2014;120:291-304.