<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="3.0" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher">BG</journal-id>
<journal-title-group>
<journal-title>Biogeosciences</journal-title>
<abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1726-4189</issn>
<publisher><publisher-name>Copernicus Publications</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.5194/bg-10-2945-2013</article-id>
<title-group>
<article-title>Trophic state of benthic deep-sea ecosystems from two different continental margins off Iberia</article-title>
</title-group>
<contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Dell'Anno</surname>
<given-names>A.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Pusceddu</surname>
<given-names>A.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Corinaldesi</surname>
<given-names>C.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Canals</surname>
<given-names>M.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Heussner</surname>
<given-names>S.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Thomsen</surname>
<given-names>L.</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Danovaro</surname>
<given-names>R.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
</contrib-group><aff id="aff1">
<label>1</label>
<addr-line>Department of Life and Environmental Science, Polytechnic University of Marche, Via Brecce Bianche, 60131 Ancona, Italy</addr-line>
</aff>
<aff id="aff2">
<label>2</label>
<addr-line>GRC Geociències Marines, Departament d&apos;Estratigrafia, Paleontologia i Geociències Marines, Facultat de Geologia, Universitat de Barcelona, Campus de Pedralbes, Martí i Franquès s/n, 08028 Barcelona, Spain</addr-line>
</aff>
<aff id="aff3">
<label>3</label>
<addr-line>CEFREM, UMR 5110 CNRS-University of Perpignan, 66860 Perpignan Cedex, France</addr-line>
</aff>
<aff id="aff4">
<label>4</label>
<addr-line>School of Engineering and Science – SES, Earth and Space Sciences, Jacobs University, Campus Ring 1, 28759 Bremen, Germany</addr-line>
</aff>
<pub-date pub-type="epub">
<day>02</day>
<month>05</month>
<year>2013</year>
</pub-date>
<volume>10</volume>
<issue>5</issue>
<fpage>2945</fpage>
<lpage>2957</lpage>
<permissions>
<copyright-statement>Copyright: &#x000a9; 2013 A. Dell'Anno et al.</copyright-statement>
<copyright-year>2013</copyright-year>
<license license-type="open-access">
<license-p>This work is licensed under the Creative Commons Attribution 3.0 Unported License. To view a copy of this licence, visit <ext-link ext-link-type="uri"  xlink:href="https://creativecommons.org/licenses/by/3.0/">https://creativecommons.org/licenses/by/3.0/</ext-link></license-p>
</license>
</permissions>
<self-uri xlink:href="https://bg.copernicus.org/articles/10/2945/2013/bg-10-2945-2013.html">This article is available from https://bg.copernicus.org/articles/10/2945/2013/bg-10-2945-2013.html</self-uri>
<self-uri xlink:href="https://bg.copernicus.org/articles/10/2945/2013/bg-10-2945-2013.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/10/2945/2013/bg-10-2945-2013.pdf</self-uri>
<abstract>
<p>The bioavailability of organic matter in benthic deep-sea ecosystems,
commonly used to define their trophic state, can greatly influence key
ecological processes such as biomass production and nutrient cycling. Here,
we assess the trophic state of deep-sea sediments from open slopes and
canyons of the Catalan (NW Mediterranean) and Portuguese (NE Atlantic)
continental margins, offshore east and west Iberia, respectively, by using a
biomimetic approach based on enzymatic digestion of protein and carbohydrate
pools. Patterns of sediment trophic state were analyzed in relation to
increasing water depth, including repeated samplings over a 3 yr period
in the Catalan margin. Two out of the three sampling periods occurred a few
months after dense shelf water cascading events. The benthic deep-sea
ecosystems investigated in this study were characterized by high amounts of
bioavailable organic matter when compared to other deep-sea sediments.
Bioavailable organic matter and its nutritional value were significantly
higher in the Portuguese margin than in the Catalan margin, thus reflecting
differences in primary productivity of surface waters reported for the two
regions. Similarly, sediments of the Catalan margin were characterized by
significantly higher food quantity and quality in spring, when the
phytoplankton bloom occurs in surface waters, than in summer and autumn.
Differences in the benthic trophic state of canyons against open slopes were
more evident in the Portuguese than in the Catalan margin. In both
continental margins, bioavailable organic C concentrations did not vary or
increase with increasing water depth. Overall, our findings suggest that the
intensity of primary production processes along with the lateral transfer of
organic particles, even amplified by episodic events, can have a role in
controlling the quantity and distribution of bioavailable organic detritus
and its nutritional value along these continental margin ecosystems.</p>
</abstract>
<counts><page-count count="13"/></counts>
</article-meta>
</front>
<body/>
<back>
<ref-list>
<title>References</title>
<ref id="ref1">
<label>1</label><mixed-citation publication-type="other" xlink:type="simple">Alvarez-Salgado, X. A., Figueiras, F. G., Perez, F. F., Groom, S., Nogueira, E., Borges, A. V., Chou, L., Moncoiffe, G., R\i os, A. F., Miller, A. E. J., Frankignoulle, M., Savidge, G., and Wollast, R.: The Portugal coastal counter current off NW Spain: new insights on its biogeochemical variability, Prog. Oceanogr., 56, 281–321, 2003.</mixed-citation>
</ref>
<ref id="ref2">
<label>2</label><mixed-citation publication-type="other" xlink:type="simple">Anderson, M. J., Gorley, R. N., and Clarke, K. R.: PERMANOVA+ for PRIMER: Guide to software and statistical methods, PRIMER-E: Plymouth, UK, 2008.</mixed-citation>
</ref>
<ref id="ref3">
<label>3</label><mixed-citation publication-type="other" xlink:type="simple">Arzola, R. G., Wynn, R. B., Lastras, G. B., Masson, D. G., and Weaver, P. P. E.: Sedimentary features and processes in the Nazarè and Setubal submarine canyons, west Iberian margin, Mar. Geol., 250, 64–88, 2008.</mixed-citation>
</ref>
<ref id="ref4">
<label>4</label><mixed-citation publication-type="other" xlink:type="simple">Bauer, J. E. and Druffel, E. R. M.: Ocean margins as a significant source of organic matter to the deep open ocean, Nature, 392, 482–485, 1998..</mixed-citation>
</ref>
<ref id="ref5">
<label>5</label><mixed-citation publication-type="other" xlink:type="simple">Bianchelli, S., Gambi, C., Zeppilli, D., and Danovaro, R.: Metazoan meiofauna in deep-sea canyons and adjacent open slopes: A large-scale comparison with focus on the rare taxa, Deep-Sea Res. I, 57, 420–433, 2010.</mixed-citation>
</ref>
<ref id="ref6">
<label>6</label><mixed-citation publication-type="other" xlink:type="simple">Billett, D. S. M., Bett, B. J., Reid, W. D. K., Boorman, B., and Priede, M.: Long-term change in the abyssal NE Atlantic: The `Amperima Event&apos; revisited, Deep-Sea Res. II, 57, 1406–1417, 2010.</mixed-citation>
</ref>
<ref id="ref7">
<label>7</label><mixed-citation publication-type="other" xlink:type="simple">Bourgeois, S., Pruski, A. M., Sun, M.-Y., Buscail, R., Lantoine, F., Kerhervé, P., Vétion, G., Rivière, B., and Charles, F.: Distribution and lability of land-derived organic matter in the surface sediments of the Rhône prodelta and the adjacent shelf (Mediterranean Sea, France): a multi proxy study, Biogeosciences, 8, 3107–3125, &lt;a href=&quot;http://dx.doi.org/10.5194/bg-8-3107-2011&quot;&gt;https://doi.org/10.5194/bg-8-3107-2011&lt;/a&gt;, 2011.</mixed-citation>
</ref>
<ref id="ref8">
<label>8</label><mixed-citation publication-type="other" xlink:type="simple">Buscail, R. and Germain, C.: Present-day organic matter sedimentation on the NW Mediterranean margin: importance of off-shelf export, Limnol. Oceanogr., 42, 217–229, 1997.</mixed-citation>
</ref>
<ref id="ref9">
<label>9</label><mixed-citation publication-type="other" xlink:type="simple">Buscail, R., Pocklington, R., and Germain, C.: Seasonal variability of the organic matter in a sedimentary coastal environment: sources, degradation and accumulation (continental shelf of the Gulf of Lions-northwestern Mediterranean Sea), Cont. Shelf. Res., 15, 843–869, 1995.</mixed-citation>
</ref>
<ref id="ref10">
<label>10</label><mixed-citation publication-type="other" xlink:type="simple">Canals, M., Puig, P., Durrieu de Madron, X., Heussner, S., Palanques, A., and Fabres J.: Flushing submarine canyons, Nature, 444, 354–357, 2006.</mixed-citation>
</ref>
<ref id="ref11">
<label>11</label><mixed-citation publication-type="other" xlink:type="simple">Canals, M., Danovaro, R.,. Heussner, S., Lykousis, V., Puig, P., Trincardi, F., Calafat, A. M., Durrieu de Madron, X., Palanques, A., and Sànchez-Vidal, A.: Cascades in Mediterranean submarine grand canyons, Oceanography, 22, 26–43, 2009.</mixed-citation>
</ref>
<ref id="ref12">
<label>12</label><mixed-citation publication-type="other" xlink:type="simple">Cunha, M. R., Paterson, G. L. J., Amaro, T., Blackbird, S., de Stigter, H. C., Ferreira, C., Glover, A., Hilário, A.,&amp;nbsp;Kiriakoulakis, K.,&amp;nbsp;Neal, L.,&amp;nbsp;Ravara, A.,&amp;nbsp;Rodrigues, C. F.,&amp;nbsp;Tiago, Á., and Billett, D. S. M.: Biodiversity of macrofaunal assemblages from three Portuguese submarine canyons (NE Atlantic), Deep-Sea Res. II, 58, 2433–2447, 2011.</mixed-citation>
</ref>
<ref id="ref13">
<label>13</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Dell&apos;Anno, A., and Fabiano, M.: Bioavailability of organic matter in the sediments of the Porcupine Abyssal Plain, northeastern Atlantic, Mar. Ecol. Prog. Ser., 220, 25–32, 2001a.</mixed-citation>
</ref>
<ref id="ref14">
<label>14</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Dell&apos;Anno, A., Fabiano, M., Pusceddu, A., and Tselepides, A.: &lt;br/&gt; Deep-sea ecosystem response to climate changes: the eastern Mediterranean case study, Trends Ecol. Evol., 16, 505–510, 2001b.</mixed-citation>
</ref>
<ref id="ref15">
<label>15</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Della Croce, N., Dell&apos;Anno, A., and Pusceddu, A.: A depocenter of organic matter at 7800 m depth in the SE Pacific Ocean, Deep-Sea Res. I, 50, 1411–1420, 2003.</mixed-citation>
</ref>
<ref id="ref16">
<label>16</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Dell&apos;Anno, A., Corinaldesi, C., Magagnini, M., Noble, R., Tamburini, C., and Weinbauer, M.: Major viral impact on the functioning of benthic deep-sea ecosystems, Nature, 454, 1084–1087, 2008a.</mixed-citation>
</ref>
<ref id="ref17">
<label>17</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Gambi, C., Dell&apos;Anno, A., Corinaldesi, C., Fraschetti, S., Vanreusel, A., Vincx, M., and Gooday, A. J.: Exponential decline of deep-sea ecosystem functioning linked to benthic biodiversity loss, Cur. Biol., 18, 1–8, 2008b.</mixed-citation>
</ref>
<ref id="ref18">
<label>18</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R.: Methods for the study of deep-sea sediments, their functioning and biodiversity, CRC press, Taylor &amp; Francis Group, USA, 2010.</mixed-citation>
</ref>
<ref id="ref19">
<label>19</label><mixed-citation publication-type="other" xlink:type="simple">Danovaro, R., Canals, M., Gambi, C., Heussner, S., Lampadariou, N., and Vanreusel, A.: Exploring benthic biodiversity patterns and hotspots on European margin slopes, Oceanography, 22, 16–25, 2010.</mixed-citation>
</ref>
<ref id="ref20">
<label>20</label><mixed-citation publication-type="other" xlink:type="simple">Dauwe, B., Middelburg, J. J., Van-Rijswijk, P., Sinke, J., Herman, P. M. J., and Heip, C. H. R.: Enzymatically hydrolyzable amino acids in North Sea sediments and their possible implication for sediment nutritional values, J. Mar. Res., 57, 109–134, 1999a.</mixed-citation>
</ref>
<ref id="ref21">
<label>21</label><mixed-citation publication-type="other" xlink:type="simple">Dauwe, B., Middelburg, J. J., Herman, P. M. J., and Heip, C. H. R.: Linking diagenetic alteration of amino acids and bulk organic matter reactivity, Limnol. Oceanogr., 44, 1809–1814, 1999b.</mixed-citation>
</ref>
<ref id="ref22">
<label>22</label><mixed-citation publication-type="other" xlink:type="simple">De Leo, F. C., Smith, C. R., Rowden, A. A., Bowden, D. A., and Clark, M. R.: Submarine canyons: hot spots of benthic biomass and productivity in the deep sea, P. Roy. Soc B-Biol Sci., 277, 2783–2792, 2010.</mixed-citation>
</ref>
<ref id="ref23">
<label>23</label><mixed-citation publication-type="other" xlink:type="simple">de Stigter, H. C., Boer, W., de Jesus Mendes, P.A., Jesus, C. C., Thomsen, L., van den Bergh, G. D., and van Weering, T. C. E.: Recent sediment transport and deposition in the Nazarè Canyon, Portuguese continental margin, Mar. Geol., 246, 144–164, 2007.</mixed-citation>
</ref>
<ref id="ref24">
<label>24</label><mixed-citation publication-type="other" xlink:type="simple">Dell&apos;Anno, A., and Danovaro, R.: Extracellular DNA plays a key role in deep-sea ecosystem functioning, Science, 309, p. 2179, 2005.</mixed-citation>
</ref>
<ref id="ref25">
<label>25</label><mixed-citation publication-type="other" xlink:type="simple">Dell&apos;Anno, A., Fabiano, M., Mei, M. L., and Danovaro, R.: Enzymatically hydrolised protein and carbohydrate pools in deep-sea sediments: estimates of the potential bioavailable fraction and methodological considerations, Mar. Ecol. Prog. Ser., 196, 15–23, 2000.</mixed-citation>
</ref>
<ref id="ref26">
<label>26</label><mixed-citation publication-type="other" xlink:type="simple">Dell&apos;Anno, A., Mei, M. L., Pusceddu, A., and Danovaro, R.: Assessing the trophic state and eutrophication of coastal marine systems: a new approach based on the biochemical composition of sediment organic matter, Mar. Poll. Bull., 44, 611–622, 2002.</mixed-citation>
</ref>
<ref id="ref27">
<label>27</label><mixed-citation publication-type="other" xlink:type="simple">Dell&apos;Anno, A., Corinaldesi, C., Stavrakakis, S., Lykousis, V., and Danovaro, R.: Pelagic-benthic coupling and diagenesis of nucleic acids in a deep-sea continental margin and an open-slope system of the Eastern Mediterranean, Appl. Environ. Microbiol., 71, 6070–6076, 2005.</mixed-citation>
</ref>
<ref id="ref28">
<label>28</label><mixed-citation publication-type="other" xlink:type="simple">Dunne, J. P., Sarmiento, J. L., and Gnanadesikan, A.: A synthesis of global particle export from the surface ocean and cycling through the ocean interior and on the seafloor, Global Biogeochem. Cycles, 21, GB4006, &lt;a href=&quot;http://dx.doi.org/10.1029/2006GB002907&quot;&gt;https://doi.org/10.1029/2006GB002907&lt;/a&gt;, 2007.</mixed-citation>
</ref>
<ref id="ref29">
<label>29</label><mixed-citation publication-type="other" xlink:type="simple">Estrada, M.: Primary production in the northwestern Mediterranean. Sci. Mar., 60, 55–64, 1996.</mixed-citation>
</ref>
<ref id="ref30">
<label>30</label><mixed-citation publication-type="other" xlink:type="simple">Epping, E., van der Zee, C., Soetaert, K., and Helder, W.: On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazarè Canyon (NE Atlantic), Prog. Oceanogr., 52, 399–431, 2002.</mixed-citation>
</ref>
<ref id="ref31">
<label>31</label><mixed-citation publication-type="other" xlink:type="simple">Fabiano, M., Danovaro, R., and Fraschetti, S.: A three-year time series of elemental and biochemical composition of organic matter in subtidal sandy sediments of the Ligurian Sea (north western Mediterranean), Cont. Shelf Res., 15, 1453–1469, 1995.</mixed-citation>
</ref>
<ref id="ref32">
<label>32</label><mixed-citation publication-type="other" xlink:type="simple">Fabiano, M., Pusceddu, A., Dell&apos;Anno, A., Armeni, M., Vanucci, S., Lampitt, R., Wolff, G., and Danovaro, R.: Fluxes of phytopigments and labile organic matter to the deep ocean in the NE Atlantic Ocean, Prog. Oceanogr., 50, 89–104, 2001.</mixed-citation>
</ref>
<ref id="ref33">
<label>33</label><mixed-citation publication-type="other" xlink:type="simple">García, R., and Thomsen, L.: Bioavailable organic matter in surface sediments of the Nazarè canyon and adjacent slope (Western Iberian Margin), J. Mar. Syst., 74, 44–59, 2008.</mixed-citation>
</ref>
<ref id="ref34">
<label>34</label><mixed-citation publication-type="other" xlink:type="simple">George, J. D.: Organic matter available to the polychaete &lt;i&gt;Cirriformia tentaculata&lt;/i&gt; (Montagu) living in an intertidal mud flat, Limnol. Oceanogr., 9, 453–455, 1964.</mixed-citation>
</ref>
<ref id="ref35">
<label>35</label><mixed-citation publication-type="other" xlink:type="simple">Gooday, A. J.: Biological responses to seasonally varying fluxes of organic matter to the ocean floor: a review, J. Oceanogr., 58, 305–332, 2002.</mixed-citation>
</ref>
<ref id="ref36">
<label>36</label><mixed-citation publication-type="other" xlink:type="simple">Grémare, A., Gutiérrez, D., Anschutz, P., Amouroux, J. M., Deflandre, B., and Vétion, G.: Spatio-temporal changes in totally and enzymatically hydrolyzable amino acids of superficial sediments from three contrasted areas, Prog. Oceanogr., 65, 89–111, 2005.</mixed-citation>
</ref>
<ref id="ref37">
<label>37</label><mixed-citation publication-type="other" xlink:type="simple">Heussner, S., Durrieu de Madron X., Calafat A., Canals, M., Carbonne, J., Delsaut, N., and Saragoni, G.: Spatial and temporal variability of downward particle fluxes on a continental slope: Lessons from an 8-yr experiment in the Gulf of Lions (NW Mediterranean), Mar. Geol. 234: 63–92, 2006.</mixed-citation>
</ref>
<ref id="ref38">
<label>38</label><mixed-citation publication-type="other" xlink:type="simple">Ingels, J., Kiriakoulakis, K., Wolff, G. A., and Vanreusel, A.: Nematode diversity and its relation to the quantity and quality of sedimentary organic matter in the deep Nazarè Canyon, Western Iberian Margin. Deep-Sea Res., 56, 1521–1539, 2009.</mixed-citation>
</ref>
<ref id="ref39">
<label>39</label><mixed-citation publication-type="other" xlink:type="simple">Ivanov, V. V., Shapiro, G. I., Huthnance, J. M., Aleynik, D. L., and Golovin, P. N.: Cascades of dense water around the world ocean, Progr. Oceanogr., 60, 47–98, 2004.</mixed-citation>
</ref>
<ref id="ref40">
<label>40</label><mixed-citation publication-type="other" xlink:type="simple">Kiriakoulakis, K., Blackbird, S., Ingels, J., Vanreusel, A., and Wolff, G. A.: Organic geochemistry of submarine canyons: The Portuguese Margin, Deep-Sea Res. II, 58, 2477–2488, 2011.</mixed-citation>
</ref>
<ref id="ref41">
<label>41</label><mixed-citation publication-type="other" xlink:type="simple">Lampitt, R. S., Bett, B. J., Kiriakoulakis, K., Popova, E. E., Ragueneau, O., Vangriesheim, A., and Wolff, G. A.: Material supply to the abyssal seafloor in the Northeast Atlantic, Prog. Oceanogr., 50, 27–63, 2001.</mixed-citation>
</ref>
<ref id="ref42">
<label>42</label><mixed-citation publication-type="other" xlink:type="simple">Lefevre, D., Minas, H. J., Minas, M., Robinson, C., Williams, P. J. L. E. B., and Woodward, E. M. S.: Review of gross community production, primary production, net community production and dark community respiration in the Gulf of Lions, Deep-Sea Res., 44, 801–832, 1997.</mixed-citation>
</ref>
<ref id="ref43">
<label>43</label><mixed-citation publication-type="other" xlink:type="simple">Liu, K-K., Atkinson, L., Quiñones, R., and Talaue-McManus, L.: Carbon and nutrient fluxes in continental margins: a global synthesis, Springer-Verlag, Berlin Heidelberg, 2010.</mixed-citation>
</ref>
<ref id="ref44">
<label>44</label><mixed-citation publication-type="other" xlink:type="simple">López-Fernández, P., Bianchelli, S., Pusceddu, A., Calafat, A., Danovaro, R., and Canals, M.: Bioavailable compounds in sinking particulate organic matter, Blanes Canyon, NW Mediterranean Sea: effects of a large storm and sea surface biological processes, Progr. Oceanogr., in press, 2013.</mixed-citation>
</ref>
<ref id="ref45">
<label>45</label><mixed-citation publication-type="other" xlink:type="simple">Lorenzen, C. and Jeffrey, J.: Determination of chlorophyll in seawater. Technical Paper in Marine Science (UNESCO), 35, 1–20, 1980.</mixed-citation>
</ref>
<ref id="ref46">
<label>46</label><mixed-citation publication-type="other" xlink:type="simple">Martìn, J., Palanques, A., Vitorino, J., Oliveira, A., and de Stigter, H. C.: Near-bottom particulate matter dynamics in the Nazarè submarine canyon under calm and stormy conditions, Deep-Sea Res. II, 58, 2388–2400, 2011.</mixed-citation>
</ref>
<ref id="ref47">
<label>47</label><mixed-citation publication-type="other" xlink:type="simple">Masson, D. G., Huvenne, V. A. I., de Stigter, H. C., Wolff, G. A., Kiriakoulakis, K., Arzola, R. G., and Blackbird, S.: Efficient burial of carbon in a submarine canyon, Geology, 38, 831–834, 2010.</mixed-citation>
</ref>
<ref id="ref48">
<label>48</label><mixed-citation publication-type="other" xlink:type="simple">Mayer, L. M.: Sedimentary organic matter preservation: an assessment and speculative synthesis – a comment, Mar. Chem., 49, 123–126, 1995.</mixed-citation>
</ref>
<ref id="ref49">
<label>49</label><mixed-citation publication-type="other" xlink:type="simple">Mayer, L. M., Schick, L. L., Sawyer, T., Plante, C. J., Jumars, P. A., and Self, R. L.: Bioavailable amino acids in sediments: a biomimetic, kinetics-based approach, Limnol. Oceanogr., 40, 511–520, 1995.</mixed-citation>
</ref>
<ref id="ref50">
<label>50</label><mixed-citation publication-type="other" xlink:type="simple">Mayer, L. M., Schick, L. L., and Allison, M. A.: Input of nutritionally rich organic matter from the Mississippi river to the Louisiana coastal Zone, Estuar. Coast., 31, 1052–1062, 2008.</mixed-citation>
</ref>
<ref id="ref51">
<label>51</label><mixed-citation publication-type="other" xlink:type="simple">MacArthur, R. H. and Pianka, E. R.: On the optimal use of a patchy environment, Am. Nat., 100, 603–609, 1966.</mixed-citation>
</ref>
<ref id="ref52">
<label>52</label><mixed-citation publication-type="other" xlink:type="simple">McClain, C. R. and Barry, J. P.: Habitat heterogeneity, disturbance, and productivity work in concert to regulate biodiversity in deep submarine canyons, Ecology, 91, 964–976, 2010.</mixed-citation>
</ref>
<ref id="ref53">
<label>53</label><mixed-citation publication-type="other" xlink:type="simple">Middelburg, J. J. and Meysman, F. J. R.: Burial at sea, Science, 317, 1294–1295, 2007.</mixed-citation>
</ref>
<ref id="ref54">
<label>54</label><mixed-citation publication-type="other" xlink:type="simple">Middelburg, J. J., Nieuwenhuize, J., and van Breugel, P.: Black carbon in marine sediments, Mar. Chem., 65, 245–252, 1999.</mixed-citation>
</ref>
<ref id="ref55">
<label>55</label><mixed-citation publication-type="other" xlink:type="simple">Millot, C.: The Gulf of Lions&apos; hydrodynamics, Cont. Shelf Res., 10, 885–894, 1990.</mixed-citation>
</ref>
<ref id="ref56">
<label>56</label><mixed-citation publication-type="other" xlink:type="simple">Palanques, A., Durrieu de Madron, X., Puig, P., Fabres, J., Guilleñ, J., Calafat, A., Canals, M., Heussner, S., and Bonnin, J.: Suspended sediment fluxes and transport processes in the Gulf of Lions submarine canyons. The role of storms and dense water cascading, Mar. Geol., 236, 43–61, 2006.</mixed-citation>
</ref>
<ref id="ref57">
<label>57</label><mixed-citation publication-type="other" xlink:type="simple">Palanques, A., Puig, P., Durrieu de Madron, X., Sanchez-Vidal, A., Pasqual, C., Martín, J., Calafat, A., Heussner, S., and Canals, M.: Sediment transport to the deep canyons and open-slope of the western Gulf of Lions during the 2006 intense cascading and open-sea convection period, Progr. Oceanogr., 106, 1–15, 2012.</mixed-citation>
</ref>
<ref id="ref58">
<label>58</label><mixed-citation publication-type="other" xlink:type="simple">Pasqual, C., Sanchez-Vidal, A., Zúñiga, D., Calafat, A., Canals, M., Durrieu de Madron, X., Puig, P., Heussner, S., Palanques, A., and Delsaut, N.: Flux and composition of settling particles across the continental margin of the Gulf of Lion: the role of dense shelf water cascading, Biogeosciences, 7, 217–231, &lt;a href=&quot;http://dx.doi.org/10.5194/bg-7-217-2010&quot;&gt;https://doi.org/10.5194/bg-7-217-2010&lt;/a&gt;, 2010.</mixed-citation>
</ref>
<ref id="ref59">
<label>59</label><mixed-citation publication-type="other" xlink:type="simple">Paterson, G. L. J., Glover, A. G., Cunha, M. R., Neal, L., de Stigter, H. C., Kiriakoulakis, K., Billett, D. S. M., Wolff, G. A., Tiago, A., Ravara, A., Lamont, P., and Tyler P.: Disturbance, productivity and diversity in deep-sea canyons: A worm&apos;s eye view, Deep-Sea Res. II, 58, 2448–2460, 2011.</mixed-citation>
</ref>
<ref id="ref60">
<label>60</label><mixed-citation publication-type="other" xlink:type="simple">Pusceddu, A., Dell&apos;Anno, A., Danovaro, R., Manini, E., Sarà, G., and Fabiano, M.: Enzymatically hydrolyzable protein and carbohydrate sedimentary pools as indicators of the trophic state of &apos;detritus sink&apos; systems: a case study in a Mediterranean coastal lagoon, Estuaries, 26, 641–650, 2003.</mixed-citation>
</ref>
<ref id="ref61">
<label>61</label><mixed-citation publication-type="other" xlink:type="simple">Pusceddu, A., Dell&apos;Anno, A., Fabiano, M., and Danovaro, R.: Quantity, biochemical composition and bioavailability of sediment organic matter as complementary signatures of benthic trophic status, Mar. Ecol. Prog. Ser. 375, 41–52, 2009.</mixed-citation>
</ref>
<ref id="ref62">
<label>62</label><mixed-citation publication-type="other" xlink:type="simple">Pusceddu, A., Bianchelli, S., Sanchez Vidal, A., Canals, M., Durrieu De Madron, X., Heussner, S., Lykousis, V., de Stigter, H., Trincardi, F., and Danovaro, R.: Organic matter in sediments of canyons and open slopes in the Eastern Atlantic and Mediterranean continental margins, Deep-Sea Res. I, 57, 441–457, 2010.</mixed-citation>
</ref>
<ref id="ref63">
<label>63</label><mixed-citation publication-type="other" xlink:type="simple">Rex, M. A., Etter, R. J., Morris, J. S., Crouse, J., McClain, C. R., Johnson, N. A., Stuart, C. T., Thies, R., and Avery, R.: Global bathymetric patterns of standing stock and body size in the deep-sea benthos, Mar. Ecol. Progr. Ser. 317, 1–8, 2006.</mixed-citation>
</ref>
<ref id="ref64">
<label>64</label><mixed-citation publication-type="other" xlink:type="simple">Ruhl, H. A. and Smith, K. L.: Shifts in deep-sea community structure linked to climate and food supply, Science, 305, 513–515, 2004.</mixed-citation>
</ref>
<ref id="ref65">
<label>65</label><mixed-citation publication-type="other" xlink:type="simple">Ruhl, H. A., Ellena, J. A., and Smith, Jr., K. L.: Connections between climate, food limitation, and carbon cycling in abyssal sediment communities: a long time-series perspective, P. Natl. Acad. Sci. USA, 105, 17006–17011, 2008.</mixed-citation>
</ref>
<ref id="ref66">
<label>66</label><mixed-citation publication-type="other" xlink:type="simple">Sampéré, T. P., Bianchi, T. S., Wakeham, S. G., and Allison, M. A.: Sources of organic matter in surface sediments of the Louisiana Continental margin: effects of major depositional/transport pathways and Hurricane Ivan, Cont. Shelf Res., 28, 2472–2487, 2008.</mixed-citation>
</ref>
<ref id="ref67">
<label>67</label><mixed-citation publication-type="other" xlink:type="simple">Sanchez-García, L., de Andres, J. R., Martin-Rubi, J. A., and Louchouarn, P.: Diagenetic state and source characterization of marine sediments from the inner continental shelf of the Gulf of Cadiz (SW Spain), constrained by terrigenous biomarkers, Org. Geochem., 40, 184–194, 2009.</mixed-citation>
</ref>
<ref id="ref68">
<label>68</label><mixed-citation publication-type="other" xlink:type="simple">Sanchez-Vidal, A., Pasqual, C., Kerhervé, P., Calafat, A., Heussner, S., Palanques, A., Durrieu de Madron, X., Canals, M., and Puig, P.: Impact of dense shelf water cascading on the transfer of organic matter to the deep western Mediterranean basin, Geophys. Res. Let., 35, 1–5, 2008.</mixed-citation>
</ref>
<ref id="ref69">
<label>69</label><mixed-citation publication-type="other" xlink:type="simple">Sanchez-Vidal, A., Pasqual, C., Kerhervé, P., Heussner, S., Calafat, A. Palanques, A., Durrieu de Madron, X., Canals, M., and Puig, P.: Across margin export of organic matter by cascading events traced by stable isotopes, northwestern Mediterranean Sea, Limnol. Oceanogr., 54, 1488–1500, 2009.</mixed-citation>
</ref>
<ref id="ref70">
<label>70</label><mixed-citation publication-type="other" xlink:type="simple">Smith, K. L., and Kaufmann, R. S.: Long-term discrepancy between food supply and demand in the deep-eastern North Pacific, Science, 284, 1174–1177, 1999.</mixed-citation>
</ref>
<ref id="ref71">
<label>71</label><mixed-citation publication-type="other" xlink:type="simple">Smith, C. R., De Leo, F. C., Bernardino, A. F., Sweetman, A. K., and Martinez Arbizu, P.: Abyssal food limitation, ecosystem structure and climate change, Trends Ecol. Evol, 23, 518–528, 2008.</mixed-citation>
</ref>
<ref id="ref72">
<label>72</label><mixed-citation publication-type="other" xlink:type="simple">Smith, K. L., Ruhl, H. A., Bett, B. J., Billett, D. S. M., Lampitt, R. S., and Kaufmann, R. S.: Climate, carbon cycling, and deep-ocean ecosystems, P. Natl. Acad. Sci., 106, 19211–19218, 2010.</mixed-citation>
</ref>
<ref id="ref73">
<label>73</label><mixed-citation publication-type="other" xlink:type="simple">Sun, M., Aller, R. C., and Lee, C.: Early diagenesis of chlorophyll a in Long Island Sound sediments: a measure of carbon flux and particle reworking, J. Mar. Res., 49, 279–401, 1991.</mixed-citation>
</ref>
<ref id="ref74">
<label>74</label><mixed-citation publication-type="other" xlink:type="simple">Thiel, H.: Benthos in upwelling regions. In: Boje R, Tomczak M (eds) Upwelling ecosystems. Springer-Verlag, Berlin, 124–138, 1978.</mixed-citation>
</ref>
<ref id="ref75">
<label>75</label><mixed-citation publication-type="other" xlink:type="simple">Tselepides, A., Polychronaki, T., Marrale, D., Akoumianaki, I., Dell&apos;Anno, A., Pusceddu, A., and Danovaro, R.: Organic matter composition of the continental shelf and bathyal sediments of the Cretan Sea (NE Mediterranean), Prog. Oceanogr., 46, 311–344, 2000.</mixed-citation>
</ref>
<ref id="ref76">
<label>76</label><mixed-citation publication-type="other" xlink:type="simple">Underwood, A. J.: Experiments in Ecology: Their logical design and interpolation using analysis of variance, Cambridge University Press, Cambridge, 1997.</mixed-citation>
</ref>
<ref id="ref77">
<label>77</label><mixed-citation publication-type="other" xlink:type="simple">van Oevelen, D., Soetaert, K., García Novoa, R., de Stigter, H., da Cunha, M., Pusceddu, A., and Danovaro, R.: Canyon conditions impact carbon flows in food webs of three sections of the Nazarè canyon, Deep-Sea Res. II, 58, 2461–2476, 2011.</mixed-citation>
</ref>
<ref id="ref78">
<label>78</label><mixed-citation publication-type="other" xlink:type="simple">van Oevelen, D., Soetaert, K., and Heip, C.: Carbon flows in the benthic food web of the Porcupine Abyssal Plain: The (un)importance of labile detritus in supporting microbial and faunal carbon demands, Limnol. Oceanogr., 57, 645–664, 2012.</mixed-citation>
</ref>
<ref id="ref79">
<label>79</label><mixed-citation publication-type="other" xlink:type="simple">Vitorino, J., Oliveira, A., Jouanneau, J. M., and Drago, T.: Winter dynamics on the northern Portuguese shelf. Part 1: physical processes, Prog. Oceanogr., 52, 29–153, 2002.</mixed-citation>
</ref>
<ref id="ref80">
<label>80</label><mixed-citation publication-type="other" xlink:type="simple">Wakeham, S. G. and Canuel, E. A.: Degradation and preservation of organic matter in marine sediments, in: The Handbook of Environmental Chemistry, edited by: Volkman, J. K., 295–321, 2006.</mixed-citation>
</ref>
<ref id="ref81">
<label>81</label><mixed-citation publication-type="other" xlink:type="simple">Weaver, P. E., Billett, D. M., Boetius, A., Danovaro, R., Freiwald, A., and Sibuet, M.: Hotspot ecosystem Research on Europe&apos;s Deep-Ocean Margins, Oceanography, 17, 132–143, 2004.</mixed-citation>
</ref>
</ref-list>
</back>
</article>