Inverse ecosystem models of the deep-sea: an example from ...
Canyon conditions impact carbon flows in food webs of three sections of the Nazaré canyon Dick van Oevelen1,*, Karline Soetaert1, Rosa García Novoa2,3, Henko de Stigter4, Marina da Cunha5, Antonio Pusceddu6, Roberto Danovaro61 Centre for Estuarine and Marine Ecology, Netherlands Institute of Ecology (NIOO-KNAW), POB 140, 4400 AC Yerseke, The Netherlands2 Jacobs University Bremen, Campus Ring 1, 28759 Bremen, Germany3 Department of Global Change Research, IMEDEA (CSIC-UIB) Instituto Mediterráneo de Estudios Avanzados, Miquel Marqués 21, 07190 Esporles, Spain4 Royal Netherlands Institute for Sea Research (NIOZ), POB 59, 1790 AB Den Burg - Texel, The Netherlands 5 Centro de Estudos do Ambiente e do Mar (CESAM) & Departamento de Biologia, Universidade de Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal 6 Department of Marine Science, Polytechnic University of Marche, Via Brecce Bianche, 60131 Ancona, Italy* corresponding author: d.vanoevelen@nioo.knaw.nl AbstractSubmarine canyons directly transport large amounts of sediment and organic matter (OM) from the continental shelf to the abyssal plain. Three carbon-based food web models were constructed for the upper (300 – 750 m water depth), middle (2700 – 3500 m) and lower section (4000 – 5000 m) of the Nazaré canyon (eastern Atlantic Ocean) using linear inverse modeling to examine how the food web is influenced by the characteristics of the respective canyon section. The models were based on an empirical dataset consisting of biomass and carbon processing data, and general physiological data constraints from the literature. Environmental conditions, most notably organic matter (OM) input and hydrodynamic activity, differed between the canyon sections and strongly affected the benthic food web structure. Despite the large difference in depth, the OM inputs into the food webs of the upper and middle sections were of similar magnitude (7.98±0.84 and 9.30±0.71 mmol C m-2 d-1, respectively). OM input to the lower section was however almost 6-7 times lower (1.26±0.03 mmol C m-2 d-1). Canyon conditions greatly influenced OM processing within the food web. Carbon processing in the upper section was dominated by prokaryotes (70% of total respiration), though there was a significant meiofaunal (21%) and smaller macrofaunal (9%) contribution. The high total faunal contribution to carbon processing resembles that found in shallower continental shelves and upper slopes, although the meiofaunal contribution is surprisingly high and suggest that high current speeds and sediment resuspension in the upper canyon favor the role of the meiofauna. The high OC input and conditions in the accreting sediments of the middle canyon section were more beneficial for megafauna (holothurians), than for the other food web compartments. The high megafaunal biomass (516 mmol C m-2), their large contribution to respiration (56% of total respiration) and secondary production (0.08 mmol C m-2 d-1) shows that these accreting sediments in canyons are megafaunal hotspots in the deep-sea. Conversely, carbon cycling in the lower canyon section was strongly dominated by prokaryotes (86% of respiration) and the food web structure therefore resembled that of lower slope and abyssal plain sediments. This study shows that elevated OM input in canyons may favor the faunal contribution to carbon processing and create hotspots of faunal biomass and carbon processing along the continental shelf.IntroductionSubmarine canyons are incisions of the continental margin and directly link the continental shelf with deep-sea plains by transporting large amounts of sediment PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5kZSBTdGlndGVyPC9BdXRob3I+PFllYXI+MjAwNzwvWWVh
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ADDIN EN.CITE.DATA (Epping et al., 2002; Vetter and Dayton, 1999). The comparatively rapid transport in active canyons results in the sedimentary OM being also of higher quality as compared to slope sediments at similar water depth PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HYXJjaWE8L0F1dGhvcj48WWVhcj4yMDA3PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Garcia et al., 2007; Pusceddu et al., 2010; Vetter and Dayton, 1999). The high quantity and quality of the OM in canyon sediments results in carbon oxidation rates PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5FcHBpbmc8L0F1dGhvcj48WWVhcj4yMDAyPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Epping et al., 2002; Rabouille et al., 2009) and benthic standing stocks of nematodes ADDIN EN.CITE <EndNote><Cite><Author>Ingels</Author><Year>2009</Year><RecNum>1896</RecNum><record><rec-number>1896</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1896</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Wolff, G. A.</author><author>Vanreusel, A.</author></authors></contributors><titles><title>Nematode diversity and its relation to the quantity and quality of sedimentary organic matter in the deep Nazare Canyon, Western Iberian Margin</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>1521-1539</pages><volume>56</volume><number>9</number><dates><year>2009</year></dates><isbn>0967-0637</isbn><accession-num>WOS:000268611100009</accession-num><urls><related-urls><url><Go to ISI>://WOS:000268611100009</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr.2009.04.010</electronic-resource-num></record></Cite></EndNote>(Ingels et al., 2009) and deposit feeding holothurians PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BbWFybzwvQXV0aG9yPjxZZWFyPjIwMDk8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Amaro et al., 2009; De Leo et al., 2010; Vetter and Dayton, 1999) that are higher as compared to adjacent open slopes and indicate extensive carbon cycling in the benthic food web.These latter studies focus on individual components of the benthic food web and suggest that different benthic components may benefit from the enhanced influx of OM into canyons. These comparisons are, however, based on single biomass-to-biomass or process-by-process comparisons. It is unclear how the structure of the whole food web and carbon partitioning within the food web is affected by canyon conditions. Moreover, it is unclear whether and how emerging properties at the whole food web level are impacted by canyon conditions. Network analysis has been developed to condense information contained in complex networks, such as food webs, into interpretable indices ADDIN EN.CITE <EndNote><Cite><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite><Cite><Author>Fath</Author><Year>1999</Year><RecNum>1877</RecNum><record><rec-number>1877</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1877</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fath, B. D.</author><author>Patten, B. C.</author></authors></contributors><titles><title>Review of the foundations of network environ analysis</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>167-179</pages><volume>2</volume><number>2</number><dates><year>1999</year><pub-dates><date>Mar-Apr</date></pub-dates></dates><isbn>1432-9840</isbn><accession-num>ISI:000080173500006</accession-num><label>Fath99</label><urls><related-urls><url><Go to ISI>://000080173500006 </url></related-urls></urls></record></Cite></EndNote>(Fath and Patten, 1999; Ulanowicz, 2004). The index total system throughput ( QUOTE ) sums carbon flows in the food web to obtain a measure of total food web activity. The Finn cycling index summarizes the fraction of total carbon cycling that is generated by recycling processes ADDIN EN.CITE <EndNote><Cite><Author>Allesina</Author><Year>2004</Year><RecNum>1874</RecNum><record><rec-number>1874</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1874</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Allesina, S.</author><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Cycling in ecological networks: Finn's index revisited</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>227-233</pages><volume>28</volume><number>3</number><dates><year>2004</year><pub-dates><date>Jul</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000223122100008</accession-num><label>Allesina04</label><urls><related-urls><url><Go to ISI>://000223122100008 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.04.002</electronic-resource-num></record></Cite></EndNote>(Allesina and Ulanowicz, 2004). Another index that is claimed to be related to food web maturity is average mutual information (AMI), that gauges how orderly and coherently ?ows are inter-connected ADDIN EN.CITE <EndNote><Cite><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><Suffix> and references therein</Suffix><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite></EndNote>(Ulanowicz, 2004 and references therein). It is claimed that AMI is indicative of the developmental status of an ecosystem and that while a food web develops specialization results in higher values of AMI.The Nazaré canyon intersects the Portuguese continental shelf and extends from a water depth of 50 m near the coast down to 5000 m at the abyssal plain and presents an interesting case study because of the varying conditions within the canyon. The upper canyon section (50 – 2700 m water depth) is characterized by a V-shaped valley that is deeply incised in the continental shelf. The middle canyon (2700 – 4000 m) is a broad meandering valley with terraced slopes that may experience high rates of particle and organic matter sedimentation ADDIN EN.CITE <EndNote><Cite><Author>Masson</Author><Year>Submitted, this issue</Year><RecNum>2067</RecNum><record><rec-number>2067</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">2067</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Masson, D. G.</author><author>Huvenne, V. A. I.</author><author>Smeed, D. A.</author><author>Arzola, R. G.</author></authors></contributors><titles><title>Sedimentary processes in the middle Nazaré Canyon: the importance of small-scale heterogeneity in defining the large-scale canyon environment</title></titles><dates><year>Submitted, this issue</year></dates><urls></urls></record></Cite></EndNote>(Masson et al., this issue). 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ADDIN EN.CITE.DATA (Aller and Aller, 2004). Carbon recycling, quantified with the Finn cycling index, may therefore be lower because fewer food web components give rise to more limited recycling in the food web. Also food web maturity, as measured with the network index AMI, is expected to be lower as compared to the middle and lower canyon sections.The terraced slopes of the middle canyon section experience high rates of sedimentation and associated organic matter input. Transport of (semi)-labile OM to these greater depths in the canyon may imply a deviation from the archetypical relation between water depth and sediment oxygen consumption (SOC). The SOC and the network index “total system throughput” is expected to be comparatively elevated in the middle section of the canyon due to the enhanced OM input as compared to open slope sediments at similar water depth. The enhanced input OM may not be partitioned equally among the food web compartments and may be influenced by the environmental conditions in the respective canyon. De Leo et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>De Leo</Author><Year>2010</Year><RecNum>2065</RecNum><record><rec-number>2065</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">2065</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>De Leo, F. C.</author><author>Smith, C. R.</author><author>Rowden, A. A.</author><author>Bowden, D. A.</author><author>Clark, M. R.</author></authors></contributors><titles><title>Submarine canyons: hotspots of benthic biomass and productivity in the deep sea</title><secondary-title>Proceedings of the Royal Society B-Biological Sciences</secondary-title></titles><periodical><full-title>Proceedings of the Royal Society B-Biological Sciences</full-title></periodical><pages>2783-2792</pages><volume>277</volume><number>1695</number><dates><year>2010</year></dates><isbn>0962-8452</isbn><accession-num>WOS:000280779700006</accession-num><urls><related-urls><url><Go to ISI>://WOS:000280779700006</url></related-urls></urls><electronic-resource-num>10.1098/rspb.2010.0462</electronic-resource-num></record></Cite></EndNote>(2010) for example, reported extremely high biomass levels of particularly deposit-feeding holothurians in a low relief muddy sediment at 900 – 1100 m in the Kaikoura Canyon (New Zealand). The conditions in the Kaikoura canyon are reported to be similar to the middle section of the Nazaré canyon and indeed high holothurian abundances are found there too ADDIN EN.CITE <EndNote><Cite><Author>Amaro</Author><Year>2009</Year><RecNum>1897</RecNum><record><rec-number>1897</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1897</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Amaro, T.</author><author>Witte, H.</author><author>Herndl, G. J.</author><author>Cunha, M. R.</author><author>Billett, D. S. M.</author></authors></contributors><titles><title>Deep-sea bacterial communities in sediments and guts of deposit-feeding holothurians in Portuguese canyons (NE Atlantic)</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>1834-1843</pages><volume>56</volume><number>10</number><dates><year>2009</year></dates><isbn>0967-0637</isbn><accession-num>WOS:000269469800016</accession-num><urls><related-urls><url><Go to ISI>://WOS:000269469800016</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr.2009.05.014</electronic-resource-num></record></Cite></EndNote>(Amaro et al., 2009). With a whole food web approach as followed here it will be possible to study quantitatively whether different food web compartments take proportional advantage of the enhanced OM input in this section of the Nazaré canyon.The deeper canyon section is where the canyon widens into a kilometres-broad channel in the abyssal plain PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5kZSBTdGlndGVyPC9BdXRob3I+PFllYXI+MjAwNzwvWWVh
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ADDIN EN.CITE.DATA (de Stigter et al., 2007). This deep canyon section, which only intermittently receives material derived from up-canyon sections via sediment gravity flows, better resembles regular abyssal plain conditions with an associated lower OM input. Under these lower OM inputs, lower faunal contributions to carbon cycling are expected and the more steady conditions may imply a higher food web maturity and higher recycling within the food web.Verifying how specific conditions in the three canyon sections impose on the benthic food web requires an analysis of the trophic structure of the complete benthic food web. The quantification of complete food webs is however a data-demanding effort and canyon data sets are typically incomplete and limited in scope. To overcome these limitations and maximize the amount of information gained from the available data, so-called linear inverse models (LIM) have been developed. LIM allow quantifying biological interactions in a complex food web from an incomplete and uncertain data set such as encountered in the deep-sea ADDIN EN.CITE <EndNote><Cite><Author>Soetaert</Author><Year>2009</Year><RecNum>1831</RecNum><record><rec-number>1831</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1831</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Soetaert, K.</author><author>Van Oevelen, D.</author></authors></contributors><titles><title>Modeling food web interactions in benthic deep-sea ecosystems: a practical guide</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>130-145</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><label>Soetaert09</label><urls></urls></record></Cite></EndNote>(Soetaert and Van Oevelen, 2009). For example, Van Oevelen et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Van Oevelen</Author><Year>2009</Year><RecNum>1832</RecNum><record><rec-number>1832</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1832</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Duineveld, G. C. A.</author><author>Lavaleye, M. S. S.</author><author>Mienis, F.</author><author>Soetaert, K.</author><author>Heip, C. H. R.</author></authors></contributors><titles><title>The cold-water coral community as hotspot of carbon cycling on continental margins: a food web analysis from Rockall Bank (northeast Atlantic)</title><secondary-title>Limnology and Oceanography</secondary-title></titles><periodical><full-title>Limnology and Oceanography</full-title><abbr-1>Limnol. Oceanogr.</abbr-1></periodical><pages>1829–1844</pages><volume>54</volume><number>6</number><dates><year>2009</year></dates><label>VanOevelen2009b</label><urls></urls></record></Cite></EndNote>(2009) using linear inverse modeling to quantify the interactions in the complex food web of a cold-water coral community at Rockall Bank and provided evidence that coral communities are hot-spots of biomass and carbon cycling along continental margins.Here we develop linear inverse models (LIM) to quantify carbon flows in the complex food webs characterizing upper, middle and lower sections of the Nazaré canyon. The observed food web structures and selected network indices are examined as a function of the characteristics of the respective canyon section.Methods2.1 Nazaré canyon characteristicsThe Nazaré canyon, one of the largest submarine canyons in Europe, intersects the Portuguese continental shelf and has been intensively studied in the framework of different European projects such as OMEX-II, EUROSTRATAFORM and HERMES. Expeditions carried out within these projects have resulted in comparatively high data availability on different physical, chemical and biological aspects of the canyon system. De Stigter et al. PEVuZE5vdGU+PENpdGUgRXhjbHVkZUF1dGg9IjEiPjxBdXRob3I+ZGUgU3RpZ3RlcjwvQXV0aG9y
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ADDIN EN.CITE.DATA (de Stigter et al., 2007). The seabed of the Nazaré canyon is heterogeneous and consists of a highly dynamic thalweg filled with coarse sandy and gravelly deposits, steep sloping canyon walls with rocky outcrops, and terraces with thick accumulations of soft muddy sediments ADDIN EN.CITE <EndNote><Cite><Author>Tyler</Author><Year>2009</Year><RecNum>1909</RecNum><record><rec-number>1909</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1909</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Tyler, P.</author><author>Amaro, T.</author><author>Arzola, R.</author><author>Cunha, M. R.</author><author>de Stigter, H.</author><author>Gooday, A.</author><author>Huvenne, V.</author><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Lastras, G.</author><author>Masson, D.</author><author>Oliveira, A.</author><author>Pattenden, A.</author><author>Vanreusel, A.</author><author>Van Weering, T.</author><author>Vitorino, J.</author><author>Witte, U.</author><author>Wolff, G.</author></authors></contributors><titles><title>Europe's Grand Canyon Nazare Submarine Canyon</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>46-57</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><isbn>1042-8275</isbn><accession-num>WOS:000263776000008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000263776000008</url></related-urls></urls></record></Cite></EndNote>(Tyler et al., 2009). The hard substrata in the thalweg and on steep walls and outcrops are covered in places with a thin, centimeter-thick drape of soft mud, where it is impossible to sample with box- or multicorer to estimate biomass. Moreover, to avoid large heterogeneity in the data set due to seabed differences, the focus of this manuscript is on soft-sediments outside the thalweg, which were split into the three sections as described above. The depth range of the upper section was here limited to 300 – 700 m. Chemical and biological data were available on the concentration of total carbohydrates, lipids and proteins in the sediment ADDIN EN.CITE <EndNote><Cite><Author>Pusceddu</Author><Year>2010</Year><RecNum>1907</RecNum><record><rec-number>1907</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1907</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Pusceddu, Antonio</author><author>Bianchelli, Silvia</author><author>Canals, Miquel</author><author>Sanchez-Vidal, Anna</author><author>Durrieu De Madron, Xavier</author><author>Heussner, Serge</author><author>Lykousis, Vasilios</author><author>de Stigter, Henko</author><author>Trincardi, Fabio</author><author>Danovaro, Roberto</author></authors></contributors><titles><title>Organic matter in sediments of canyons and open slopes of the Portuguese, Catalan, Southern Adriatic and Cretan Sea margins</title><secondary-title>Deep Sea Research Part I: Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep Sea Research Part I: Oceanographic Research Papers</full-title></periodical><pages>441-457</pages><volume>27</volume><keywords><keyword>Submarine canyons</keyword><keyword>Open slopes</keyword><keyword>Sediment organic matter</keyword><keyword>Trophic conditions</keyword><keyword>Mediterranean Sea</keyword><keyword>Atlantic Ocean</keyword></keywords><dates><year>2010</year></dates><isbn>0967-0637</isbn><work-type>doi: DOI: 10.1016/j.dsr.2009.11.008</work-type><urls><related-urls><url>;(Pusceddu et al., 2010), sedimentary chl a content ADDIN EN.CITE <EndNote><Cite><Author>Garcia</Author><Year>2008</Year><RecNum>1910</RecNum><record><rec-number>1910</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1910</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garcia, R.</author><author>Thomsen, L.</author></authors></contributors><titles><title>Bioavailable organic matter in surface sediments of the Nazare canyon and adjacent slope (Western Iberian Margin)</title><secondary-title>Journal of Marine Systems</secondary-title></titles><periodical><full-title>Journal of Marine Systems</full-title><abbr-1>J. Mar. Syst.</abbr-1></periodical><pages>44-59</pages><volume>74</volume><number>1-2</number><dates><year>2008</year></dates><isbn>0924-7963</isbn><accession-num>WOS:000261256100002</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261256100002</url></related-urls></urls><electronic-resource-num>10.1016/j.jmarsys.2007.11.004</electronic-resource-num></record></Cite></EndNote>(Garcia and Thomsen, 2008), sediment diagenesis ADDIN EN.CITE <EndNote><Cite><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(Epping et al., 2002), prokaryotic heterotrophic carbon production (Danovaro, unpub. data), nematode trophic structure ADDIN EN.CITE <EndNote><Cite><Author>Danovaro</Author><Year>2009</Year><RecNum>1908</RecNum><record><rec-number>1908</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1908</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Danovaro, R.</author><author>Bianchelli, S.</author><author>Gambi, C.</author><author>Mea, M.</author><author>Zeppilli, D.</author></authors></contributors><titles><title>alpha-, beta-, gamma-, delta- and epsilon-diversity of deep-sea nematodes in canyons and open slopes of Northeast Atlantic and Mediterranean margins</title><secondary-title>Marine Ecology-Progress Series</secondary-title></titles><periodical><full-title>Marine Ecology-Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><pages>197-209</pages><volume>396</volume><dates><year>2009</year></dates><isbn>0171-8630</isbn><accession-num>WOS:000273549400019</accession-num><urls><related-urls><url><Go to ISI>://WOS:000273549400019</url></related-urls></urls><electronic-resource-num>10.3354/meps08269</electronic-resource-num></record></Cite></EndNote>(Danovaro et al., 2009) and the macro- and megafaunal community structure (Cunha et al., this issue and unpub. data). Such data on biotic and abiotic carbon stocks and transformation rates are perfectly suited to quantify food webs of the three sections of the Nazaré canyon using linear inverse modeling.2.2 Linear inverse modelsThe food web models developed for the Nazaré canyon are constructed using linear inverse modeling ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2010). In an inverse model, the food web compartments and flows between them are fixed a priori (see ‘Food web structure’ below). The flow magnitudes are constrained within the boundaries that are defined by the inclusion of empirical data on standing stocks, flux data and physiology into the model. The food web topology and empirical data are included in a matrix equation with equalities and in a matrix equation with inequalities. These matrix equations are solved simultaneously to recover quantitative values for the flow values, such that the flow values in a model solution are within the boundaries defined by the matrix equations. The model was run 10,000 times and each time a different solution is generated to allow estimating the mean and standard deviation of each unknown flow. It is important to note that by running the model 10,000 times, the uncertainty in the empirical data (see ‘Data availability’ below) is propagated onto an uncertainty estimate of the carbon flows as indicated by its standard deviation. Convergence of the mean and standard deviation of the flows was used to verify whether the set of 10,000 model solutions was sufficiently large.Several reviews on the technical and methodological aspects of linear inverse modeling have been published and will therefore not be repeated here ADDIN EN.CITE <EndNote><Cite><Author>Soetaert</Author><Year>2009</Year><RecNum>1831</RecNum><record><rec-number>1831</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1831</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Soetaert, K.</author><author>Van Oevelen, D.</author></authors></contributors><titles><title>Modeling food web interactions in benthic deep-sea ecosystems: a practical guide</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>130-145</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><label>Soetaert09</label><urls></urls></record></Cite><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Soetaert and Van Oevelen, 2009; Van Oevelen et al., 2010). These reviews contain simple models to exemplify the setup and solution of linear inverse food web models using the software packages LIM ADDIN EN.CITE <EndNote><Cite><Author>Soetaert</Author><Year>2008</Year><RecNum>1830</RecNum><record><rec-number>1830</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1830</key></foreign-keys><ref-type name="Computer Program">9</ref-type><contributors><authors><author>Soetaert, K.</author><author>Van Oevelen, D</author></authors></contributors><titles><title>LIM: Linear Inverse Model examples and solution methods. R package version 1.2</title></titles><dates><year>2008</year></dates><label>Soetaert08</label><urls><related-urls><url> Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Soetaert and Van Oevelen, 2008; Van Oevelen et al., 2010) and limSolve ADDIN EN.CITE <EndNote><Cite><Author>Soetaert</Author><Year>2008</Year><RecNum>1835</RecNum><record><rec-number>1835</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1835</key></foreign-keys><ref-type name="Computer Program">9</ref-type><contributors><authors><author>Soetaert, K.</author><author>Van den Meersche, K.</author><author>Van Oevelen, D.</author></authors></contributors><titles><title>limSolve: Solving linear inverse models. R package version 1.3</title></titles><dates><year>2008</year></dates><label>Soetaert08b</label><urls><related-urls><url>;(Soetaert et al., 2008) that run in the R software ADDIN EN.CITE <EndNote><Cite><Author>R Development Core Team</Author><Year>2008</Year><RecNum>1751</RecNum><record><rec-number>1751</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1751</key></foreign-keys><ref-type name="Computer Program">9</ref-type><contributors><authors><author>R Development Core Team,</author></authors></contributors><titles><title>R: A language and environment for statistical computing</title></titles><dates><year>2008</year></dates><pub-location>Vienna, Austria</pub-location><publisher>R Foundation for Statistical Computing</publisher><isbn>3-900051-07-0</isbn><label>R08</label><urls><related-urls><url>;(R Development Core Team, 2008). The Nazaré food web models are made publically available in the LIM package.2.3 Food web structureThe compartments in the food web models were chosen based on the classical size distribution of prokaryotes (Pro), meiofauna (Mei), macrofauna (Mac) and megafauna (Meg). The faunal compartments were further subdivided based on the feeding classification for nematodes ADDIN EN.CITE <EndNote><Cite><Author>Wieser</Author><Year>1953</Year><RecNum>1931</RecNum><record><rec-number>1931</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1931</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Wieser, W.</author></authors></contributors><titles><title>Die Beziehung zwischen Mundh?hlengestalt, Ern?hrungsweise und Vorkommen bei freilebenden marinen Nematoden. Eine skologisen-morphologische studie</title><secondary-title><style face="normal" font="default" size="100%">Arkiv f</style><style face="normal" font="Symbol" charset="2" size="100%">ü</style><style face="normal" font="default" size="100%">r Zoologie</style></secondary-title></titles><periodical><full-title>Arkiv für Zoologie</full-title></periodical><pages>439-484</pages><volume>4</volume><section>439</section><dates><year>1953</year></dates><urls></urls></record></Cite></EndNote>(Wieser, 1953) and feeding types for macro- and megafauna were surface deposit-feeder (SDF), deposit-feeder (DF), suspension feeder (SF) and predator+scavenger (PS) (see below). The sedimentary organic matter was divided into dissolved organic carbon (DOC) and labile (lDet), semi-labile (sDet) and refractory detritus (rDet).Inputs to the food web are deposition and/or suspension feeding of suspended labile (lDet_w), semi-labile (sDet_w) and refractory detritus (rDet_w). Outputs from the food web are respiration to dissolved inorganic carbon (DIC), burial of rDet, DOC efflux to the water column and export by the macro- and megafaunal compartments (e.g. consumption by fish).The detritus pools in the sediment can be hydrolyzed to DOC and the labile and semi-labile detritus pools are grazed upon by meiofauna and MacSDF, MacDF, MacPS, MegSDF and MegDF. DOC is taken up by prokaryotes or fluxes out of the sediment to the water column. Predatory feeding links are primarily defined based on size class; prokaryotes are consumed by all meiofaunal and non-suspension feeding macro- and megafaunal compartments, meiofaunal compartments are consumed by non-suspension feeding macro- and megafaunal compartments, the macrofaunal compartments MacSDF, MacDF and MacSF are preyed upon by MacPS.Part of the ingested matter by the faunal compartments is not assimilated but instead expelled as feces, the non-assimilated labile (e.g. labile detritus, prokaryotes and faunal compartments) and semi-labile (semi-labile detritus) carbon, flows into semi-labile and refractory detritus, respectively. Respiration by faunal compartments is defined as the sum of maintenance respiration (biomass-specific respiration) and growth respiration (overhead on new biomass production). Prokaryotic mortality is represented here as a flux to DOC and faunal mortality is defined as a flux to labile detritus.2.4 Data availabilityThe Nazaré canyon is one of the best studied canyons in Europe, with studies on sediment transport and/or fate of organic matter PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5kZSBTdGlndGVyPC9BdXRob3I+PFllYXI+MjAwNzwvWWVh
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ADDIN EN.CITE.DATA (e.g. de Stigter et al., 2007; Epping et al., 2002; García et al., 2008), concentration of total carbohydrates, lipids and proteins in the sediment ADDIN EN.CITE <EndNote><Cite><Author>Pusceddu</Author><Year>2010</Year><RecNum>1907</RecNum><record><rec-number>1907</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1907</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Pusceddu, Antonio</author><author>Bianchelli, Silvia</author><author>Canals, Miquel</author><author>Sanchez-Vidal, Anna</author><author>Durrieu De Madron, Xavier</author><author>Heussner, Serge</author><author>Lykousis, Vasilios</author><author>de Stigter, Henko</author><author>Trincardi, Fabio</author><author>Danovaro, Roberto</author></authors></contributors><titles><title>Organic matter in sediments of canyons and open slopes of the Portuguese, Catalan, Southern Adriatic and Cretan Sea margins</title><secondary-title>Deep Sea Research Part I: Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep Sea Research Part I: Oceanographic Research Papers</full-title></periodical><pages>441-457</pages><volume>27</volume><keywords><keyword>Submarine canyons</keyword><keyword>Open slopes</keyword><keyword>Sediment organic matter</keyword><keyword>Trophic conditions</keyword><keyword>Mediterranean Sea</keyword><keyword>Atlantic Ocean</keyword></keywords><dates><year>2010</year></dates><isbn>0967-0637</isbn><work-type>doi: DOI: 10.1016/j.dsr.2009.11.008</work-type><urls><related-urls><url>;(Pusceddu et al., 2010) heterotrophic prokaryotic C production (Danovaro unpub. data), nematode community structure PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HYXJjaWE8L0F1dGhvcj48WWVhcj4yMDA3PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Garcia et al., 2007; Danovaro et al., 2009; Ingels et al., 2009), meiofaunal abundance ADDIN EN.CITE <EndNote><Cite><Author>Bianchelli</Author><Year>2010</Year><RecNum>1948</RecNum><record><rec-number>1948</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1948</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Bianchelli, S.</author><author>Gambi, C.</author><author>Zeppilli, D.</author><author>Danovaro, R.</author></authors></contributors><titles><title>Metazoan meiofauna in deep-sea canyons and adjacent open slopes: A large-scale comparison with focus on the rare taxa</title><secondary-title>Deep Sea Research Part I: Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep Sea Research Part I: Oceanographic Research Papers</full-title></periodical><pages>420-433</pages><volume>57</volume><number>3</number><keywords><keyword>Deep-sea meiofaunal biodiversity</keyword><keyword>Mediterranean Sea</keyword><keyword>Atlantic Ocean</keyword><keyword>Continental margins</keyword></keywords><dates><year>2010</year></dates><isbn>0967-0637</isbn><work-type>doi: DOI: 10.1016/j.dsr.2009.12.001</work-type><urls><related-urls><url>;(Bianchelli et al., 2010), macro- and megafaunal community structure ADDIN EN.CITE <EndNote><Cite><Author>Tyler</Author><Year>2009</Year><RecNum>1909</RecNum><Suffix>`, Cunha et al.`, this issue and unpub. data</Suffix><record><rec-number>1909</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1909</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Tyler, P.</author><author>Amaro, T.</author><author>Arzola, R.</author><author>Cunha, M. R.</author><author>de Stigter, H.</author><author>Gooday, A.</author><author>Huvenne, V.</author><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Lastras, G.</author><author>Masson, D.</author><author>Oliveira, A.</author><author>Pattenden, A.</author><author>Vanreusel, A.</author><author>Van Weering, T.</author><author>Vitorino, J.</author><author>Witte, U.</author><author>Wolff, G.</author></authors></contributors><titles><title>Europe's Grand Canyon Nazare Submarine Canyon</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>46-57</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><isbn>1042-8275</isbn><accession-num>WOS:000263776000008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000263776000008</url></related-urls></urls></record></Cite></EndNote>(Tyler et al., 2009, Cunha et al., this issue and unpub. data). As stated above, empirical data were only included if they were collected from the soft-sediments of the upper, middle or lower section of the canyon.Detritus stocks were delineated as follows (Table 1): the stock of labile detritus was defined as all carbon associated with chlorophyll a. Chlorophyll a concentrations were taken from the top 5 cm in sediments of the off-thalweg stations ADDIN EN.CITE <EndNote><Cite><Author>Garcia</Author><Year>2008</Year><RecNum>1910</RecNum><record><rec-number>1910</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1910</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garcia, R.</author><author>Thomsen, L.</author></authors></contributors><titles><title>Bioavailable organic matter in surface sediments of the Nazare canyon and adjacent slope (Western Iberian Margin)</title><secondary-title>Journal of Marine Systems</secondary-title></titles><periodical><full-title>Journal of Marine Systems</full-title><abbr-1>J. Mar. Syst.</abbr-1></periodical><pages>44-59</pages><volume>74</volume><number>1-2</number><dates><year>2008</year></dates><isbn>0924-7963</isbn><accession-num>WOS:000261256100002</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261256100002</url></related-urls></urls><electronic-resource-num>10.1016/j.jmarsys.2007.11.004</electronic-resource-num></record></Cite></EndNote>(Garcia and Thomsen, 2008), which were converted to carbon units by assuming a carbon to chl a ratio of 40. Semi-labile detritus was defined as the sum of the carbohydrates, lipids and proteins (i.e. biopolymeric carbon) that were converted to carbon equivalents ADDIN EN.CITE <EndNote><Cite><Author>Pusceddu</Author><RecNum>1907</RecNum><record><rec-number>1907</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1907</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Pusceddu, Antonio</author><author>Bianchelli, Silvia</author><author>Canals, Miquel</author><author>Sanchez-Vidal, Anna</author><author>Durrieu De Madron, Xavier</author><author>Heussner, Serge</author><author>Lykousis, Vasilios</author><author>de Stigter, Henko</author><author>Trincardi, Fabio</author><author>Danovaro, Roberto</author></authors></contributors><titles><title>Organic matter in sediments of canyons and open slopes of the Portuguese, Catalan, Southern Adriatic and Cretan Sea margins</title><secondary-title>Deep Sea Research Part I: Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep Sea Research Part I: Oceanographic Research Papers</full-title></periodical><pages>441-457</pages><volume>27</volume><keywords><keyword>Submarine canyons</keyword><keyword>Open slopes</keyword><keyword>Sediment organic matter</keyword><keyword>Trophic conditions</keyword><keyword>Mediterranean Sea</keyword><keyword>Atlantic Ocean</keyword></keywords><dates><year>2010</year></dates><isbn>0967-0637</isbn><work-type>doi: DOI: 10.1016/j.dsr.2009.11.008</work-type><urls><related-urls><url>;(Pusceddu et al., 2010). Biopolymeric carbon concentrations were measured only in the top 1 cm and were linearly extrapolated to 5 cm depth under the assumption that all semi-labile detritus is degraded in the top 5 cm. The latter assumption is supported by Epping et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(2002) who showed that carbon degradation occurs primarily in the top 5 cm of the sediment. Refractory detritus was defined as the degradable fraction of the particulate organic carbon in the top 5 cm of the sediment PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5FcHBpbmc8L0F1dGhvcj48WWVhcj4yMDAyPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (derived from organic carbon content profiles in Epping et al., 2002), minus the labile and semi-labile detritus pools.Biomass data were available for prokaryotes and all faunal compartments (i.e., meiofaunal, macrofauna and megafauna; Table 1). Nematodes dominated the metazoan meiofauna (on average 90% of total abundance) and the Wieser feeding classification based on nematode mouth morphology was used to designate biomass to selective feeding (Wieser type 1A + 2A), non-selective feeding (Wieser type 1B) and omnivore/predatory (Wieser type 1B). Polychaetes dominated the macrofaunal compartments and these were grouped into surface-deposit, deposit, suspension and predatory+scavenging feeding compartment based on standard feeding type classification from Fauchald and Jumars ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Fauchald</Author><Year>1979</Year><RecNum>1370</RecNum><record><rec-number>1370</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1370</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fauchald, K.</author><author>Jumars, P. A.</author></authors></contributors><titles><title>The diet of worms: A study of polychaete feeding guilds</title><secondary-title>Oceanography and Marine Biology: an Annual Review</secondary-title><alt-title>Oceanogr. Mar. Biol. Ann. Rev.</alt-title></titles><periodical><full-title>Oceanography and Marine Biology: an Annual Review</full-title><abbr-1>Oceanogr. Mar. Biol. Ann. Rev.</abbr-1></periodical><alt-periodical><full-title>Oceanography and Marine Biology: an Annual Review</full-title><abbr-1>Oceanogr. Mar. Biol. Ann. Rev.</abbr-1></alt-periodical><pages>193-284</pages><volume>39</volume><dates><year>1979</year></dates><label>Fauchald79</label><urls></urls></record></Cite></EndNote>(1979). Biomass-dominant polychaete families in the upper section are Onuphidae (57%) and Sigalionidae (36%), in the middle section Spionidae (61%), Fauveliopsidae (9%) and Ampharetidae (8%), and in the lower section Spionidae (40%), Goniadidae (15%) and Siboglinidae (12%). Other contributions to the macrofaunal biomass from Mollusca, Bivalvia and Crustacea are low (< 3%) in the upper section, higher in the middle section with 48%, 14% and 19%, and negligible in the lower section (<1%), respectively. Finally, the megafaunal surface-deposit feeding community consists of Ypsilothuria bitentaculata (Holothuroidea) and deposit feeding community of Molpadia musculus (Holothuroidea).Since there were no data available on the temporal variability in benthic biomass, these were neglected and it was assumed that the mass balances of all compartments are in steady-state, i.e., QUOTE . This assumption introduces only limited bias in the model solution ADDIN EN.CITE <EndNote><Cite><Author>Vézina</Author><Year>2003</Year><RecNum>655</RecNum><record><rec-number>655</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">655</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Vézina, A. F.</author><author>Pahlow, M.</author></authors></contributors><titles><title>Reconstruction of ecosystem flows using inverse methods: how well do they work?</title><secondary-title>Journal of Marine Systems</secondary-title><alt-title>J. Mar. Syst.</alt-title></titles><periodical><full-title>Journal of Marine Systems</full-title><abbr-1>J. Mar. Syst.</abbr-1></periodical><alt-periodical><full-title>Journal of Marine Systems</full-title><abbr-1>J. Mar. Syst.</abbr-1></alt-periodical><pages>55-77</pages><volume>40</volume><dates><year>2003</year><pub-dates><date>Apr</date></pub-dates></dates><accession-num>ISI:000182625300004</accession-num><label>Vezina03</label><urls><related-urls><url><Go to ISI>://000182625300004</url></related-urls></urls></record></Cite></EndNote>(Vézina and Pahlow, 2003), primarily because net biomass increases (e.g. for the fauna and bacteria) are small as compared to the other flows in the food web.In addition to the standing stock measurements, a variety of data on process rates were available for the different sections of the Nazaré canyon (Table 2). These data were implemented as inequalities by setting the minimum and maximum value found in each section as lower and upper bounds, respectively. The determination of prokaryotic C production in sediment samples was carried out according to the procedure described for marine sediments by Danovaro et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Danovaro</Author><Year>2002</Year><RecNum>1316</RecNum><record><rec-number>1316</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1316</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Danovaro, R.</author><author>Manini, E.</author><author>Dell'Anno, A.</author></authors></contributors><titles><title>Higher abundance of bacteria than of viruses in deep Mediterranean sediments</title><secondary-title>Applied and Environmental Microbiology</secondary-title></titles><periodical><full-title>Applied and Environmental Microbiology</full-title><abbr-1>Appl. Environ. Microbiol.</abbr-1></periodical><pages>1468-1472</pages><volume>68</volume><number>3</number><dates><year>2002</year><pub-dates><date>Mar</date></pub-dates></dates><accession-num>ISI:000174206200060</accession-num><label>Danovaro02a</label><urls><related-urls><url><Go to ISI>://000174206200060</url></related-urls></urls></record></Cite></EndNote>(2002). Sediment subsamples from the top 1 cm were mixed with a solution of 3H-leucine (final concentration 0.2 mmol L-1), were incubated at in situ temperature for 1 hour in the dark. After incubation, samples were supplemented with ethanol (80%) and processed according to Van Duyl and Kop ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Van Duyl</Author><Year>1994</Year><RecNum>308</RecNum><record><rec-number>308</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">308</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Duyl,F.C.</author><author>Kop,A.J.</author></authors></contributors><titles><title>Bacterial production in North Sea sediments: Clues to seasonal and spatial variations</title><secondary-title>Marine Biology</secondary-title><alt-title>Mar. Biol.</alt-title></titles><periodical><full-title>Marine Biology</full-title><abbr-1>Mar. Biol.</abbr-1></periodical><alt-periodical><full-title>Marine Biology</full-title><abbr-1>Mar. Biol.</abbr-1></alt-periodical><pages>323-337</pages><volume>120</volume><keywords><keyword>bacterial production</keyword><keyword>benthic</keyword><keyword>organic matter</keyword><keyword>sediment</keyword></keywords><dates><year>1994</year><pub-dates><date>1994</date></pub-dates></dates><label>VanDuyl94</label><urls></urls></record></Cite></EndNote>(1994) before scintillation counting. Sediment blanks were made adding ethanol immediately after 3H-leucine addition. The incorporated radioactivity in all samples was measured by a liquid scintillation counter. The following equation was used for calculating prokaryotic C production:PCP ~ LI · 131.2 · (%Leu) – 1 · (C: protein) · IDwhere PCP is prokaryotic C production, LI is the leucine incorporation rate (mol ml-1 h-1), 131.2 is the molecular weight of leucine, %Leu is the fraction of leucine in protein (0.073), C:protein is the ratio of cellular carbon to protein (0.86), and ID is the isotope dilution assuming a value of 2.The prokaryotic C production was determined in the top 1 cm and this value was taken as lower bound on prokaryotic production (Table 2). Prokaryote production typically decreases with depth in the sediment due to reduced availability of degradable detritus and electron acceptors ADDIN EN.CITE <EndNote><Cite><Author>Nodder</Author><Year>2003</Year><RecNum>726</RecNum><Prefix>e.g. </Prefix><record><rec-number>726</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">726</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Nodder, S.D.</author><author>Pilditch, C.A.</author><author>Keith Probert, P.</author><author>Hall, J.A.</author></authors></contributors><titles><title>Variability in benthic biomass and activity beneath the Subtropical Front, Chatman Rise, SW Pacific Ocean</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>959-985</pages><volume>50</volume><dates><year>2003</year></dates><label>Nodder03</label><urls></urls></record></Cite><Cite><Author>Glud</Author><Year>2004</Year><RecNum>1299</RecNum><record><rec-number>1299</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1299</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Glud, R.N.</author><author>Middelboe, M.</author></authors></contributors><titles><title>Virus and bacteria dynamics of a coastal sediment: Implications for benthic carbon cycling</title><secondary-title>Limnology and Oceanography</secondary-title></titles><periodical><full-title>Limnology and Oceanography</full-title><abbr-1>Limnol. Oceanogr.</abbr-1></periodical><pages>2073-2081</pages><volume>49</volume><number>6</number><dates><year>2004</year></dates><label>Glud04</label><urls></urls></record></Cite></EndNote>(e.g. Nodder et al., 2003; Glud and Middelboe, 2004). The upper bound on prokaryotic C production for the top 5 cm was set to five times the prokaryotic C production of the top 1 cm. As such, we impose that the integrated prokaryotic C production does not increase within the top 5 cm of the sediment, because the model solution is found between the lower bound (production in top 1 cm layer) and the upper bound (5 times the production in the top 1 cm layer). Carbon burial rates, total respiration rates, total carbon deposition and burial efficiencies for each section were taken from the diagenetic modeling work of Epping et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(2002) (Table 2). We imposed that total respiration and carbon deposition in Epping et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(2002) did not include the respiration and uptake by megafauna, respectively, because the activity of these large burrowing or surface-dwelling organisms is missed in a diagenetic modeling approach that is based on small cores incubations and oxygen profiles in the sediment. An additional number of general inequality constraints were taken from the literature to constrain degradation rates of the labile, semi-labile and refractory detritus pools, prokaryote growth efficiency, release of DOC from the sediment, assimilation efficiency of all faunal compartments, net growth efficiency of all faunal compartments, production and mortality rates of all faunal compartments (Table 2). Since measurements of assimilation and growth efficiencies of deep-sea benthos are very rare, we decided to use an extensive literature review ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2006</Year><RecNum>1496</RecNum><record><rec-number>1496</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1496</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Herman, P. M. J.</author><author>Moodley, L.</author><author>Hamels, I.</author><author>Moens, T.</author><author>Heip, C. H. R.</author></authors></contributors><titles><title>Carbon flows through a benthic food web: Integrating biomass, isotope and tracer data</title><secondary-title>Journal of Marine Research</secondary-title><alt-title>J. Mar. Res.</alt-title></titles><periodical><full-title>Journal of Marine Research</full-title><abbr-1>J. Mar. Res.</abbr-1></periodical><alt-periodical><full-title>Journal of Marine Research</full-title><abbr-1>J. Mar. Res.</abbr-1></alt-periodical><pages>1-30</pages><volume>64</volume><number>3</number><dates><year>2006</year></dates><label>VanOevelen06c</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2006b) of temperate benthos as basis for these constraints. Biomass-specific maintenance respiration of all faunal compartments was defined as 0.01 d-1 at 20°C ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2006</Year><RecNum>1496</RecNum><IDText>VanOevelen06c</IDText><Prefix>see references in </Prefix><record><rec-number>1496</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1496</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Herman, P. M. J.</author><author>Moodley, L.</author><author>Hamels, I.</author><author>Moens, T.</author><author>Heip, C. H. R.</author></authors></contributors><titles><title>Carbon flows through a benthic food web: Integrating biomass, isotope and tracer data</title><secondary-title>Journal of Marine Research</secondary-title><alt-title>J. Mar. Res.</alt-title></titles><periodical><full-title>Journal of Marine Research</full-title><abbr-1>J. Mar. Res.</abbr-1></periodical><alt-periodical><full-title>Journal of Marine Research</full-title><abbr-1>J. Mar. Res.</abbr-1></alt-periodical><pages>1-30</pages><volume>64</volume><number>3</number><dates><year>2006</year></dates><label>VanOevelen06c</label><urls></urls></record></Cite></EndNote>(see references in Van Oevelen et al., 2006b) and is corrected with Q10 of 2, giving a temperature-correction factor (Tlim) for each canyon section (Table 2). Benthic organisms do not feed indiscriminately on the available food sources. Both surface-deposit and deposit-feeding holothurians and echinoderms ingest organic matter with higher than ambient chlorophyll a and total hydrolysable amino acid concentrations PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HaW5nZXI8L0F1dGhvcj48WWVhcj4yMDAxPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Ginger et al., 2001; Witbaard et al., 2001; Amaro et al., 2010), though selectivity differs between feeding modes with surface-deposit feeders typically exhibiting stronger selectivity than deposit feeders ADDIN EN.CITE <EndNote><Cite><Author>Wigham</Author><Year>2003</Year><RecNum>1507</RecNum><IDText>Wigham03</IDText><record><rec-number>1507</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1507</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Wigham, B. D.</author><author>Hudson, I. R.</author><author>Billett, D. S. M.</author><author>Wolff, G. A.</author></authors></contributors><titles><title>Is long-term change in the abyssal Northeast Atlantic driven by qualitative changes in export flux? Evidence from selective feeding in deep-sea holothurians</title><secondary-title>Progress in Oceanography</secondary-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><pages>409-441</pages><volume>59</volume><number>4</number><dates><year>2003</year></dates><accession-num>ISI:000220421900004</accession-num><label>Wigham03</label><urls><related-urls><url><Go to ISI>://000220421900004</url></related-urls></urls></record></Cite></EndNote>(Wigham et al., 2003). Selectivity between labile detritus and semi-labile detritus for megafauna was defined as the ratio of chlorophyll a concentrations in the gut with respect to the ambient surface sediment. The level of selectivity varies from 1 to 10 for deposit feeding holothurians to >500 for the surface deposit feeding holothurians Amperima rosea ADDIN EN.CITE <EndNote><Cite><Author>Wigham</Author><Year>2003</Year><RecNum>1507</RecNum><Prefix>Porcupine Abyssal Plain`, </Prefix><record><rec-number>1507</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1507</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Wigham, B. D.</author><author>Hudson, I. R.</author><author>Billett, D. S. M.</author><author>Wolff, G. A.</author></authors></contributors><titles><title>Is long-term change in the abyssal Northeast Atlantic driven by qualitative changes in export flux? Evidence from selective feeding in deep-sea holothurians</title><secondary-title>Progress in Oceanography</secondary-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><pages>409-441</pages><volume>59</volume><number>4</number><dates><year>2003</year></dates><accession-num>ISI:000220421900004</accession-num><label>Wigham03</label><urls><related-urls><url><Go to ISI>://000220421900004</url></related-urls></urls></record></Cite></EndNote>(Porcupine Abyssal Plain, Wigham et al., 2003). Selectivity at the Antarctic Peninsula was less evident (selectivity of 2 to 7), possibly because of the existence of a food bank, but there was a clear separation between deposit and surface deposit feeders ADDIN EN.CITE <EndNote><Cite><Author>Wigham</Author><Year>2008</Year><RecNum>1939</RecNum><record><rec-number>1939</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1939</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Wigham, B. D.</author><author>Galley, E. A.</author><author>Smith, C. R.</author><author>Tyler, P. A.</author></authors></contributors><titles><title>Inter-annual variability and potential for selectivity in the diets of deep-water Antarctic echinoderms</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>2478-2490</pages><volume>55</volume><number>22-23</number><dates><year>2008</year></dates><isbn>0967-0645</isbn><accession-num>WOS:000261911500008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261911500008</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.06.007</electronic-resource-num></record></Cite></EndNote>(Wigham et al., 2008). Therefore, no to moderate selectivity of 1 to 10 for deposit feeders and strong selectivity (50 to 100) for surface-deposit feeders was assumed in the model (Table 2). Since no comparable data are available for macrofauna, similar selectivity ranges were defined for these compartments (Table 2). Finally, few organisms in benthic food webs can be considered as sole predators ADDIN EN.CITE <EndNote><Cite><Author>Fauchald</Author><Year>1979</Year><RecNum>1370</RecNum><record><rec-number>1370</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1370</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fauchald, K.</author><author>Jumars, P. A.</author></authors></contributors><titles><title>The diet of worms: A study of polychaete feeding guilds</title><secondary-title>Oceanography and Marine Biology: an Annual Review</secondary-title><alt-title>Oceanogr. Mar. Biol. Ann. Rev.</alt-title></titles><periodical><full-title>Oceanography and Marine Biology: an Annual Review</full-title><abbr-1>Oceanogr. Mar. Biol. Ann. Rev.</abbr-1></periodical><alt-periodical><full-title>Oceanography and Marine Biology: an Annual Review</full-title><abbr-1>Oceanogr. Mar. Biol. Ann. Rev.</abbr-1></alt-periodical><pages>193-284</pages><volume>39</volume><dates><year>1979</year></dates><label>Fauchald79</label><urls></urls></record></Cite></EndNote>(Fauchald and Jumars, 1979), therefore the predatory meio-, macro- and megafaunal compartments were assumed rely between 75% and 100% through predatory feeding to account for this (Table 2).2.5 Network indicesThe network indices QUOTE , QUOTE and QUOTE were directly calculated from the output of the sampling algorithm in R using the newly developed R-package NetIndices ADDIN EN.CITE <EndNote><Cite><Author>Kones</Author><Year>2009</Year><RecNum>1787</RecNum><record><rec-number>1787</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1787</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kones, J. K.</author><author>Soetaert, K.</author><author>van Oevelen, D</author><author>Owino, J. O.</author></authors></contributors><titles><title>Are network indices robust indicators of food web functioning? a Monte Carlo approach</title><secondary-title>Ecological Modelling</secondary-title></titles><periodical><full-title>Ecological Modelling</full-title><abbr-1>Ecol. Model.</abbr-1></periodical><pages>370–382</pages><volume>220</volume><dates><year>2009</year></dates><label>Kones09</label><urls></urls></record></Cite></EndNote>(Kones et al., 2009). Details on the calculation of the indices can be found in Ulanowicz ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite></EndNote>(2004) and Kones et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Kones</Author><Year>2009</Year><RecNum>1787</RecNum><record><rec-number>1787</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1787</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kones, J. K.</author><author>Soetaert, K.</author><author>van Oevelen, D</author><author>Owino, J. O.</author></authors></contributors><titles><title>Are network indices robust indicators of food web functioning? a Monte Carlo approach</title><secondary-title>Ecological Modelling</secondary-title></titles><periodical><full-title>Ecological Modelling</full-title><abbr-1>Ecol. Model.</abbr-1></periodical><pages>370–382</pages><volume>220</volume><dates><year>2009</year></dates><label>Kones09</label><urls></urls></record></Cite></EndNote>(2009), but a summary of the nomenclature (Table 3) and calculation algorithms (Table 4) are included in this manuscript. Network indices were calculated for the complete set of food web solutions (10,000 for each section). The network indices were compared between canyon sections by calculating the fraction of which the randomized set of indices of one canyon section is larger than that of another section. For example, when this fraction is 0.90, this implies that 90% of the values of section 1 are larger than the ones of section 2 (and consequently 10% of the values are lower). We define differences of >90% and <10% as significant difference and >95% and <5% as highly significant difference.Results 3.1 Food web structureThe models of the upper and middle canyon could be solved with the default equality and inequality constraints. However, the first attempt to solve the model of the lower section with the default set of constraints was unsuccessful, which indicates that some of the data embedded in the linear inverse model are in conflict with each other. Subsequent analysis showed that the minimum degradation of semi-labile detritus (4761 · 8.21·10-4 = 3.9 mmol C m-2 d-1, Table 1 & 2) was higher than the maximum rates of total carbon oxidation and carbon deposition (0.90 and 1.3 mmol C m-2 d-1, respectively). Since the latter two data are site-specific field data, it was decided to modify the literature bound on the minimum rate of semi-labile degradation through pre-multiplication with the temperature limitation factor (Tlim = 0.30, Table 2). This allowed the model to be solved and its implications will be discussed below.The mean flow values and standard deviations for the three sections of the Nazaré canyon are reported in Web appendix 1.The quality of the model solutions was evaluated with the Coefficient of Variation (CoV), which is the standard deviation of a flow divided by the mean flow value. As such, the CoV provides an indication for the residual uncertainty in the solution, where flows with a relatively large residual uncertainty have a comparatively high CoV and flows with a relatively small residual uncertainty have a comparatively low CoV. All flows in all three canyon sections had a CoV that was smaller than 1. Maximum CoV were 0.86, 0.90 and 0.86 for the upper, middle and lower canyon section, respectively and were associated with transfer of one the nematode compartments to the (surface) deposit-feeding macrobenthos. The CoV was smaller than 0.75 for 81%, 73% and 82% of the flows of the upper, middle and lower canyon section, respectively, and the CoV was smaller than 0.50 for 40%, 40% and 45% of the flows. Total carbon input (mmol C m-2 d-1) to the different food webs was 7.98±0.84 (5% labile, 75% semi-labile and 20% refractory detritus), 9.30±0.71 (9% labile, 89% semi-labile and 2% refractory detritus) and 1.26±0.03 (6% labile, 90% semi-labile and 4% refractory detritus) for the upper, middle and lower canyon section, respectively. Total respiration was 4.52±0.28, 5.06±0.30 and 0.86±0.02 mmol C m-2 d-1 and organic carbon burial was 3.05±0.80, 3.85±0.35 and 0.34±0.04 mmol C m-2 d-1 for the upper, middle and lower canyon section, respectively. Prokaryotes dominated carbon respiration in the upper (70%) and lower (82%) section, but their contribution to total respiration is lower (38%) than the total megafaunal respiration in the middle section (57%) (Table 5). Summed meiofaunal respiration contributes 21% tot total respiration in the upper, 3% in the middle and 13% in the lower canyon section, whereas summed macrofaunal respiration contributes 8% in the upper, 1% in the middle and 5% in the lower section. Summed export fluxes (i.e. secondary production not consumed within the food web) differed between the sections with 0.18±0.08, 0.10±0.05 and 0.02±0.006 mmol C m-2 d-1 for the upper, middle and lower section, respectively.The structural differences between the food webs become apparent when flows are plotted as mean net values in a circular food web structure (Fig. 1). The main differences between the upper and lower section are the more important role of the non-selective feeding meiofauna compartment (Fig. 1A vs. 1C) and MacPS compartment (Fig. 1D vs 1F) in carbon cycling in the upper canyon section. Of similar importance, however, is the pathway of deposition of semi-labile, dissolution to dissolved organic carbon, prokaryotic uptake of this DOC and prokaryotic respiration in the upper and lower sections (Fig. 1A vs. 1C). Consistent with their comparatively low contribution to total respiration, the carbon flows related to the macrofaunal compartments are small, except for the MacPS compartment in the upper canyon section that show up mostly in the lower row of Fig.1. The food web structure of the middle canyon section stands out primarily because of the dominant role of the MegDF and, to a lesser extent, MegSDF compartments (Fig. 1B and 1H). Moreover, carbon cycling by the macrobenthic compartments, especially MacPS, is less important as compared to the upper and lower canyon section. There is a dominance of semi-labile detritus in the diets of most faunal compartments in the upper section of the Nazaré canyon, with semi-labile detritus supplying between 53% and 95% of carbon of the non-predatory compartments and 11-12% of the predatory compartments MeiPS and MacPS, respectively (Fig. 2A). Labile detritus (2 – 15%) and prokaryotes (2 – 22%) supply a comparable lower fraction of carbon to the non-predatory compartments and 4 – 5% to the predatory compartments. Non-predatory meiofaunal compartments fuels the meiofaunal and macrofaunal predatory compartments in similar amounts (21 – 50%). Faunal diets of the non-predatory compartments in the middle section are comparable to the upper section, with a dominance of semi-labile detritus (42 – 93%) and labile (2 – 21%) detritus (Fig. 2B). The diet contribution of prokaryotes to non-predatory faunal compartments varies between 2 and 21%. Dependence on selective and non-selective feeding meiofaunal compartments is highest for predatory meiofauna (80%), followed by predatory macrofauna (48%) and <10% for the other macrofaunal and megafaunal compartments. The diet of the predatory/scavenging macrofaunal compartment is diverse, with no clear dominance of any resource (3 – 25%).The diet compositions in the lower section of the Nazaré canyon resemble overall those of the upper section (Fig. 2A vs. 2C). Again, semi-labile detritus is most important (between 76 – 98%) in the diets of non-predatory faunal compartments. Diet contributions of labile detritus and prokaryotes are similar for selective feeding meiofauna (9-10%), non-selective meiofauna (each 1%), predatory/omnivore meiofaunal (each 5%), surface-deposit feeding macrofauna (each 5%), deposit-feeding macrofauna (each 1%) and predatory/scavenging macrofauna (4-5%) (Fig. 2C). The meiofaunal compartments MeiSF + MeiNF are important resources for the meiofaunal predators/omnivores (together 80% of the diet) and predatory (69%) macrofauna, but are of lesser importance for surface-deposit (10%), deposit feeding (1%). The diet composition of predatory/scavenging macrofauna is diverse though with a high importance of selective feeding meiofauna (54%) and lower contributions ranging from 1 - 11% from other resources.The diet of suspension-feeding macrofauna is similar among the canyon sections and is partitioned among labile (32 – 36%) and semi-labile (64 – 68%) detritus from the water column.The dominant fate of prokaryotic production in all three sections is mortality (52 – 88%) and grazing by meiofauna in the upper canyon section (31%) and by megafauna in the middle section (36%) (Fig. 3A-C). The majority of the meiofaunal secondary production is grazed by macrofauna in the upper (56%) and lower (47%) canyon section, while megafaunal grazing is important in the middle section (36%) and grazing by meiofauna (MeiPO) is important with a consistent contribution of 18 – 23% in the three sections (Fig. 3D-F). The fate of macrofaunal production is partitioned similarly in all three canyon sections with maintenance representing 22 – 24%, mortality 29 – 34%, predation by macrofauna (MacPS) 2 – 20% and export 29 – 42% (Fig. 3G-I). The fate of megafauna is dominated by maintenance respiration (91%) and with limited contributions of mortality (5%) and export (4%) (Fig. 3J).3.2 Network indicesThe network indices total system throughput ( QUOTE ), Finn cycling index ( QUOTE ) and average mutual information ( QUOTE ) were calculated for the three sections (Fig. 4) and compared (Table 6). The QUOTE does not differ significantly between the upper and middle sections with median values of 41.1 and 39.7 mmol C m-2 d-1, respectively, but QUOTE is significantly lower in the lower section with a median of 6.7 mmol C m-2 d-1 (Table 6). Differences in QUOTE are highly significant between canyon sections (Table 6) and median values are 0.13, 0.06 and 0.17 for the upper, middle and lower section, respectively. QUOTE is not significantly different between the upper (median of 2.21) and middle (2.22) canyon section, but significantly lower for the lower section (2.12). DiscussionIn this paper, we present the first quantitative analysis of carbon flows within food webs of different sections of a submarine canyon. This provides a unique opportunity to study how different characteristics within a canyon influence food web structure and attributes such as total system throughput, recycling within the food web and food web maturity. The modeled food webs of the upper, mid and lower canyon sections are based on a large variety of site-specific biological and biogeochemical data and are combined with physiological constraints and empirical relations from the literature. Despite the large amount of data that are implemented, this is insufficient to uniquely quantify all carbon flows ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2010). This implies that a “solution space” exists, within which an infinite number of solutions are present that are consistent with the data ADDIN EN.CITE <EndNote><Cite><Author>Soetaert</Author><Year>2009</Year><RecNum>1831</RecNum><record><rec-number>1831</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1831</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Soetaert, K.</author><author>Van Oevelen, D.</author></authors></contributors><titles><title>Modeling food web interactions in benthic deep-sea ecosystems: a practical guide</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>130-145</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><label>Soetaert09</label><urls></urls></record></Cite></EndNote>(Soetaert and Van Oevelen, 2009). Conventional single-solution modeling approaches typically find a final solution at or close to boundaries of the solution space, making the final solution sensitive to the exact boundaries of the solution space PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5WYW4gT2V2ZWxlbjwvQXV0aG9yPjxZZWFyPjIwMTA8L1ll
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ADDIN EN.CITE.DATA ( Vézina et al., 2004; Kones et al., 2006; Van Oevelen et al., 2010). The multi-solution approach followed here, samples the solution space ADDIN EN.CITE <EndNote><Cite><Author>Van den Meersche</Author><Year>2009</Year><RecNum>1693</RecNum><record><rec-number>1693</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1693</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van den Meersche, K.</author><author>Soetaert, K.</author><author>Van Oevelen, D.</author></authors></contributors><titles><title>xsample(): an R function for sampling linear inverse problems</title><secondary-title>Journal of Statistical Software</secondary-title></titles><periodical><full-title>Journal of Statistical Software</full-title><abbr-1>J. Stat. Softw.</abbr-1></periodical><pages>1-15</pages><volume>30</volume><number>1</number><dates><year>2009</year></dates><label>VandenMeersche09</label><urls><related-urls><url>;(Van den Meersche et al., 2009) such that the mean of this sampled set represents the best central flow value that is less sensitive to the boundaries of the solution space ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2010). Moreover, the standard deviation on each carbon flow indicates how the uncertainty in the data set propagates to an uncertainty on its value ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2010). The Coefficient of Variation (CoV) was smaller than 0.75 for 73 – 82% flows in the three sections (Web appendix), which indicates that the residual uncertainty on the flows is comparatively low and that the food web is well-constrained. The lowest CoVs are associated with the respiration flows of the biotic compartments, whereas highest CoVs are predominantly associated with carbon flows that exist between biotic compartments. This directly relates to the data availability. The carbon requirement of faunal compartments is constrained primarily by the available biomass data. There are however few data that constrain the origin of this carbon, such that the residual uncertainty on diet contributions and fates of secondary production are comparatively high. Perhaps even more important than the residual uncertainty on the flows, are the limitations and uncertainties with respect to the assumptions that were needed to setup the model. These sources of uncertainty mainly concern substrate heterogeneity and combining different data sets and will be discussed now.The seafloor in the Nazaré canyon is heterogeneous and consists of rocks, boulders, coarse gravel sediments, steep walls, a highly dynamic thalweg and terraces consisting of soft-sediments. The hard substrata may be draped with a thin soft muddy layer. Not surprisingly, also the associated fauna changes with substratum type and condition. Rocky surfaces for example are dominated by suspension feeders such as hard and soft corals, gorgonians, anemones, sea pens and crinoids ADDIN EN.CITE <EndNote><Cite><Author>Tyler</Author><Year>2009</Year><RecNum>7</RecNum><record><rec-number>7</rec-number><ref-type name='Journal Article'>17</ref-type><contributors><authors><author>Tyler, P.</author><author>Amaro, T.</author><author>Arzola, R.</author><author>Cunha, M. R.</author><author>de Stigter, H.</author><author>Gooday, A.</author><author>Huvenne, V.</author><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Lastras, G.</author><author>Masson, D.</author><author>Oliveira, A.</author><author>Pattenden, A.</author><author>Vanreusel, A.</author><author>Van Weering, T.</author><author>Vitorino, J.</author><author>Witte, U.</author><author>Wolff, G.</author></authors></contributors><titles><title>Europe's Grand Canyon Nazare Submarine Canyon</title><secondary-title>Oceanography</secondary-title></titles><pages>46-57</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><isbn>1042-8275</isbn><accession-num>WOS:000263776000008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000263776000008</url></related-urls></urls></record></Cite></EndNote>(Tyler et al., 2009). In thalweg sediments, the biomass of nematodes ADDIN EN.CITE <EndNote><Cite><Author>Garcia</Author><Year>2007</Year><RecNum>1735</RecNum><record><rec-number>1735</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1735</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garcia, R.</author><author>Koho, K. A.</author><author>De Stigter, H. C.</author><author>Epping, E.</author><author>Koning, E.</author><author>Thomsen, L.</author></authors></contributors><titles><title>Distribution of meiobenthos in the Nazare canyon and adjacent slope (western Iberian Margin) in relation to sedimentary composition</title><secondary-title>Marine Ecology-Progress Series</secondary-title></titles><periodical><full-title>Marine Ecology-Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><pages>207-220</pages><volume>340</volume><dates><year>2007</year></dates><isbn>0171-8630</isbn><accession-num>ISI:000248011500018</accession-num><label>Garcia07</label><urls><related-urls><url><Go to ISI>://000248011500018 </url></related-urls></urls></record></Cite></EndNote>(Garcia et al., 2007) is about one order of magnitude lower than in soft-sediment terraces ADDIN EN.CITE <EndNote><Cite><Author>Ingels</Author><Year>2009</Year><RecNum>3</RecNum><record><rec-number>3</rec-number><ref-type name='Journal Article'>17</ref-type><contributors><authors><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Wolff, G. A.</author><author>Vanreusel, A.</author></authors></contributors><titles><title>Nematode diversity and its relation to the quantity and quality of sedimentary organic matter in the deep Nazare Canyon, Western Iberian Margin</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><pages>1521-1539</pages><volume>56</volume><number>9</number><dates><year>2009</year></dates><isbn>0967-0637</isbn><accession-num>WOS:000268611100009</accession-num><urls><related-urls><url><Go to ISI>://WOS:000268611100009</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr.2009.04.010</electronic-resource-num></record></Cite></EndNote>(Ingels et al., 2009), which is attributed to repeated sediment disturbance of thalweg sediments that prevents the development of a mature nematode community ADDIN EN.CITE <EndNote><Cite><Author>Garcia</Author><Year>2007</Year><RecNum>1735</RecNum><record><rec-number>1735</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1735</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garcia, R.</author><author>Koho, K. A.</author><author>De Stigter, H. C.</author><author>Epping, E.</author><author>Koning, E.</author><author>Thomsen, L.</author></authors></contributors><titles><title>Distribution of meiobenthos in the Nazare canyon and adjacent slope (western Iberian Margin) in relation to sedimentary composition</title><secondary-title>Marine Ecology-Progress Series</secondary-title></titles><periodical><full-title>Marine Ecology-Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><pages>207-220</pages><volume>340</volume><dates><year>2007</year></dates><isbn>0171-8630</isbn><accession-num>ISI:000248011500018</accession-num><label>Garcia07</label><urls><related-urls><url><Go to ISI>://000248011500018 </url></related-urls></urls></record></Cite></EndNote>(Garcia et al., 2007). In addition, megafauna and the giant epifaunal protozoans (xenophyophores) were not observed in the thalweg ADDIN EN.CITE <EndNote><Cite><Author>Tyler</Author><Year>2009</Year><RecNum>7</RecNum><record><rec-number>7</rec-number><ref-type name='Journal Article'>17</ref-type><contributors><authors><author>Tyler, P.</author><author>Amaro, T.</author><author>Arzola, R.</author><author>Cunha, M. R.</author><author>de Stigter, H.</author><author>Gooday, A.</author><author>Huvenne, V.</author><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Lastras, G.</author><author>Masson, D.</author><author>Oliveira, A.</author><author>Pattenden, A.</author><author>Vanreusel, A.</author><author>Van Weering, T.</author><author>Vitorino, J.</author><author>Witte, U.</author><author>Wolff, G.</author></authors></contributors><titles><title>Europe's Grand Canyon Nazare Submarine Canyon</title><secondary-title>Oceanography</secondary-title></titles><pages>46-57</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><isbn>1042-8275</isbn><accession-num>WOS:000263776000008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000263776000008</url></related-urls></urls></record></Cite></EndNote>(Tyler et al., 2009) but are found outside the thalweg. Up to now, there are no quantitative data available on the biomass and activity of the filter-feeding community in the Nazaré canyon on rocky substrata. Moreover, quantitative data on the faunal community in the thalweg is only sparsely available and its food web structure is not representative for that of large sections of the canyon. Hence, in this study we restricted our analysis to the soft-sediments of the terraces adjacent to the thalweg and excluded other substrate types. This implies for example that we may miss the potentially high carbon processing activity associated with the canyon walls. In terms of areal coverage however, these soft-sediments with net mud deposition represent an appreciable ~70% of the total surface area of the canyon ADDIN EN.CITE <EndNote><Cite><Author>Masson</Author><Year>2010</Year><RecNum>1949</RecNum><record><rec-number>1949</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1949</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Masson, D. G.</author><author>Huvenne, V. A.</author><author>De Stigter, H.</author><author>Wolff, G. A.</author><author>Kiriakoulakis, K.</author><author>Arzola, R. G.</author><author>Blackbird, S. J.</author></authors></contributors><titles><title>Efficient burial of carbon in a submarine canyon</title><secondary-title>Geology</secondary-title></titles><periodical><full-title>Geology</full-title></periodical><pages>831-834 </pages><volume>38</volume><number>9</number><dates><year>2010</year></dates><urls></urls></record></Cite></EndNote>(Masson et al., 2010), such that a significantly large part of the Nazaré canyon is addressed here.One compartment that is not included in the food web is Foraminifera, which are protozoans that are typically of meiofaunal size but can occur as giant epifauna (xenophyophores). Meiofaunal foraminifera ADDIN EN.CITE <EndNote><Cite><Author>Koho</Author><Year>2008</Year><RecNum>1899</RecNum><record><rec-number>1899</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1899</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Koho, K. A.</author><author>Garcia, R.</author><author>de Stigter, H. C.</author><author>Epping, E.</author><author>Koning, E.</author><author>Kouwenhoven, T. J.</author><author>van Der Zwaan, G. J.</author></authors></contributors><titles><title>Sedimentary labile organic carbon and pore water redox control on species distribution of benthic foraminifera: A case study from Lisbon-Setubal Canyon (southern Portugal)</title><secondary-title>Progress in Oceanography</secondary-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><pages>55-82</pages><volume>79</volume><number>1</number><dates><year>2008</year></dates><isbn>0079-6611</isbn><accession-num>WOS:000261023400004</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261023400004</url></related-urls></urls><electronic-resource-num>10.1016/j.pocean.2008.07.004</electronic-resource-num></record></Cite></EndNote>(Koho et al., 2008) and epifaunal xenophyophores ADDIN EN.CITE <EndNote><Cite><Author>Tyler</Author><Year>2009</Year><RecNum>1909</RecNum><record><rec-number>1909</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1909</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Tyler, P.</author><author>Amaro, T.</author><author>Arzola, R.</author><author>Cunha, M. R.</author><author>de Stigter, H.</author><author>Gooday, A.</author><author>Huvenne, V.</author><author>Ingels, J.</author><author>Kiriakoulakis, K.</author><author>Lastras, G.</author><author>Masson, D.</author><author>Oliveira, A.</author><author>Pattenden, A.</author><author>Vanreusel, A.</author><author>Van Weering, T.</author><author>Vitorino, J.</author><author>Witte, U.</author><author>Wolff, G.</author></authors></contributors><titles><title>Europe's Grand Canyon Nazare Submarine Canyon</title><secondary-title>Oceanography</secondary-title></titles><periodical><full-title>Oceanography</full-title></periodical><pages>46-57</pages><volume>22</volume><number>1</number><dates><year>2009</year></dates><isbn>1042-8275</isbn><accession-num>WOS:000263776000008</accession-num><urls><related-urls><url><Go to ISI>://WOS:000263776000008</url></related-urls></urls></record></Cite></EndNote>(Tyler et al., 2009) have a high abundance in especially the muddy terraces with stable redox conditions and low disturbance. Foraminifera have been shown to play an important role in the initial processing of fresh phytodetritus under deep-sea conditions ADDIN EN.CITE <EndNote><Cite><Author>Moodley</Author><Year>2002</Year><RecNum>723</RecNum><record><rec-number>723</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">723</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Moodley, L.</author><author>Middelburg, J. J.</author><author>Boschker, H. T. S.</author><author>Duineveld, G. C. A.</author><author>Pel, R.</author><author>Herman, P. M. J.</author><author>Heip, C. H. R.</author></authors></contributors><titles><title>Bacteria and Foraminifera: Key players in a short-term deep-sea benthic response to phytodetritus</title><secondary-title>Marine Ecology Progress Series</secondary-title><alt-title>Mar. Ecol. Prog. Ser.</alt-title></titles><periodical><full-title>Marine Ecology Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><alt-periodical><full-title>Marine Ecology Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></alt-periodical><pages>23-29</pages><volume>236</volume><dates><year>2002</year></dates><accession-num>ISI:000177007100003</accession-num><label>Moodley02</label><urls><related-urls><url><Go to ISI>://000177007100003</url></related-urls></urls></record></Cite></EndNote>(Moodley et al., 2002) although their contribution may also be more limited PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Xb3VsZHM8L0F1dGhvcj48WWVhcj4yMDA3PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Woulds et al., 2007). Moreover, their contribution to total respiration in continental shelf sediments was recently found to be limited to <3% ADDIN EN.CITE <EndNote><Cite><Author>Geslin</Author><Year>2010</Year><RecNum>2051</RecNum><record><rec-number>2051</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">2051</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Geslin, E.</author><author>Risgaard-Petersen, N.</author><author>Lombard, F.</author><author>Metzger, E.</author><author>Langlet, D.</author><author>Jorissen, F.</author></authors></contributors><titles><title>Oxygen respiration rates of benthic foraminifera as measured with oxygen microsensors</title><secondary-title>Journal of Experimental Marine Biology and Ecology</secondary-title></titles><periodical><full-title>Journal of Experimental Marine Biology and Ecology</full-title><abbr-1>J. Exp. Mar. Biol. Ecol.</abbr-1></periodical><volume>doi:10.1016/j.jembe.2010.10.011</volume><dates><year>2010</year></dates><urls></urls></record></Cite></EndNote>(Geslin et al., 2010). Unfortunately, the available abundance data could not be converted to biomass with reasonable accuracy, and since biomass is essential to constrain their activity in the food web we therefore decided to omit this compartment in this analysis.The site-specific data that we include in this study were lumped into the three canyon sections (Table 1 and 2). However, since deep-sea research is time consuming, conducted over large spatial areas and depends on ship time availability and meteorological/sea conditions, the data were not collected synoptically. Inevitably, this data ‘lumping’ into canyon sections will introduce errors in the food web analysis linked to the spatial and temporal variability of the data collected. Nevertheless, the Nazaré canyon is comparatively well-studied and one of the strengths of linear inverse modeling is that datasets are merged and tested for internal consistency ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2010</Year><RecNum>1752</RecNum><record><rec-number>1752</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1752</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Van den Meersche, K.</author><author>Meysman, F.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vézina, A. F.</author></authors></contributors><titles><title>Quantitative reconstruction of food webs using linear inverse models</title><secondary-title>Ecosystems</secondary-title></titles><periodical><full-title>Ecosystems</full-title><abbr-1>Ecosystems</abbr-1></periodical><pages>32–45</pages><volume>13</volume><dates><year>2010</year></dates><label>VanOevelen10 Ecosystems LIM</label><urls></urls></record></Cite></EndNote>(Van Oevelen et al., 2010). Given the amount of data in the models (Table 1 and 2), the inverse model analysis at least showed that the different data sets are consistent. The only exception was that the minimum degradation rate of semi-labile detritus in the lower canyon section was higher than the maximum rates of carbon oxidation and total carbon deposition. The carbon oxidation and deposition data are site-specific data and were therefore maintained. Instead, the minimum bound on semi-labile degradation was reduced by multiplication with the temperature limitation factor, which allowed solving the food web model. Several explanations may apply here. First, water temperature in the deep canyon section is about 2.5°C and lowest of the three sections. This low temperature may cause degradation to proceed slower than in the higher sections of the canyon with comparatively higher water temperatures. Moreover, the quality of the semi-labile detritus may have decreased during transport through the canyon and this may also lower the degradation rates further. Despite this minor adaptation that was needed, the results from the present analysis serve as a significant first step in gaining insight in the food web structure of submarine canyons.4.1 Upper canyon sectionThe dynamic upper canyon receives about 8±0.84 mmol C m-2 d-1, which is lower than the 15 – 23 mmol C m-2 d-1 that is predicted using an empirical relation for continental shelf sediments ADDIN EN.CITE <EndNote><Cite><Author>Middelburg</Author><Year>1997</Year><RecNum>1846</RecNum><Prefix>i.e. summed burial and mineralization rates at 700 and 300 m`, respectively`, </Prefix><record><rec-number>1846</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1846</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Middelburg, J. J.</author><author>Soetaert, K.</author><author>Herman, P. M. J.</author></authors></contributors><titles><title>Empirical relationships for use in global diagenetic models</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>327-344</pages><volume>44</volume><number>2</number><dates><year>1997</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0967-0637</isbn><accession-num>ISI:A1997WQ10500008</accession-num><label>Middelburg97</label><urls><related-urls><url><Go to ISI>://A1997WQ10500008 </url></related-urls></urls></record></Cite></EndNote>(i.e. summed burial and mineralization rates at 700 and 300 m, respectively, Middelburg et al., 1997). However, carbon inputs at the open slope sediments of the adjacent Iberian margin are substantially lower than predicted by the empirical relation by Middelburg et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Middelburg</Author><Year>1997</Year><RecNum>1846</RecNum><record><rec-number>1846</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1846</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Middelburg, J. J.</author><author>Soetaert, K.</author><author>Herman, P. M. J.</author></authors></contributors><titles><title>Empirical relationships for use in global diagenetic models</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>327-344</pages><volume>44</volume><number>2</number><dates><year>1997</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>0967-0637</isbn><accession-num>ISI:A1997WQ10500008</accession-num><label>Middelburg97</label><urls><related-urls><url><Go to ISI>://A1997WQ10500008 </url></related-urls></urls></record></Cite></EndNote>(1997) and are between 2.3 and 4.3 mmol C m-2 d-1 ADDIN EN.CITE <EndNote><Cite><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(Epping et al., 2002). Thus, carbon inputs to the upper canyon section is higher those of adjacent slopes, but not extremely high as compared to other slope sediments. Burial rates in the upper and middle canyon are substantial flows in the food web (Fig. 1A, B), but burial efficiencies are comparable to Iberian open slopes and relate to sediment accumulations rates ADDIN EN.CITE <EndNote><Cite><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(Epping et al., 2002). Hence, the efficiency with which the food web processes organic carbon is similar to open slope sediments.The model results allow detailed deciphering of the biotic compartments that are responsible for carbon processing within the canyon. Woulds et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Woulds</Author><Year>2009</Year><RecNum>1922</RecNum><record><rec-number>1922</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1922</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Woulds, C.</author><author>Andersson, J. H.</author><author>Cowie, G. L.</author><author>Middelburg, J. J.</author><author>Levin, L. A.</author></authors></contributors><titles><title>The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>393-402</pages><volume>56</volume><number>6-7</number><dates><year>2009</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000267780700011</accession-num><label>Woulds09</label><urls><related-urls><url><Go to ISI>://000267780700011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.10.008</electronic-resource-num></record></Cite></EndNote>(2009) used the results of isotope tracer experiments from different slope sediments to define different categories of biological C-processing. In this categorization, the “active-faunal-uptake” category contains mostly shallow (<300 m) slope sediments and is characterized by 10 – 25% metazoan uptake. This category matches best with the upper canyon section that has a faunal contribution of ~40% and bacterial contribution of 60% to total carbon assimilation. The faunal contribution to total respiration and carbon processing typically decreases with increasing water depth and associated decrease in carbon input PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IZWlwPC9BdXRob3I+PFllYXI+MjAwMTwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (Heip et al., 2001; Rowe et al., 2008; Woulds et al., 2009). Henceforth, the high faunal contribution in the upper canyon section is probably related to the higher OM content and quality as compared to slope sediments at comparable water depth PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HYXJjaWE8L0F1dGhvcj48WWVhcj4yMDA3PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Garcia et al., 2007; Garcia and Thomsen, 2008; Pusceddu et al., 2010). One striking difference however is that meiofauna dominated faunal processing and contributed around 33% of the total carbon assimilation in the upper canyon section, which is much higher than in open slopes sediments included in the overview of Woulds et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Woulds</Author><Year>2009</Year><RecNum>1922</RecNum><record><rec-number>1922</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1922</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Woulds, C.</author><author>Andersson, J. H.</author><author>Cowie, G. L.</author><author>Middelburg, J. J.</author><author>Levin, L. A.</author></authors></contributors><titles><title>The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>393-402</pages><volume>56</volume><number>6-7</number><dates><year>2009</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000267780700011</accession-num><label>Woulds09</label><urls><related-urls><url><Go to ISI>://000267780700011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.10.008</electronic-resource-num></record></Cite></EndNote>(2009). This high contribution also translates into a much higher meiofaunal respiration at 21% of total respiration in the upper section of the Nazaré canyon as compared to other open slopes that vary from 4 – 8% PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IZWlwPC9BdXRob3I+PFllYXI+MjAwMTwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (Piepenburg et al., 1995; Heip et al., 2001; Soetaert et al., 2009). Rowe et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Rowe</Author><Year>2008</Year><RecNum>1915</RecNum><record><rec-number>1915</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1915</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rowe, G. T.</author><author>Wei, C. L.</author><author>Nunnally, C.</author><author>Haedrich, R.</author><author>Montagna, P.</author><author>Baguley, J. G.</author><author>Bernhard, J. M.</author><author>Wicksten, M.</author><author>Ammons, A.</author><author>Briones, E. E.</author><author>Soliman, Y.</author><author>Deming, J. W.</author></authors></contributors><titles><title>Comparative biomass structure and estimated carbon flow in food webs in the deep Gulf of Mexico</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>2699-2711</pages><volume>55</volume><number>24-26</number><dates><year>2008</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000261969100021</accession-num><urls><related-urls><url><Go to ISI>://000261969100021</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.07.020</electronic-resource-num></record></Cite></EndNote>(2008) and Bagulay et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Baguley</Author><Year>2008</Year><RecNum>1918</RecNum><record><rec-number>1918</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1918</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Baguley, J. G.</author><author>Montagna, P. A.</author><author>Hyde, L. J.</author><author>Rowe, G. T.</author></authors></contributors><titles><title>Metazoan meiofauna biomass, grazing, and weight-dependent respiration in the Northern Gulf of Mexico deep sea</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>2607-2616</pages><volume>55</volume><number>24-26</number><dates><year>2008</year></dates><isbn>0967-0645</isbn><accession-num>WOS:000261969100011</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261969100011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.07.010</electronic-resource-num></record></Cite></EndNote>(2008) report even substantially higher contributions ranging from ~20 up to 51% for the Northern Gulf of Mexico. Their estimates are based on biomass-specific respiration rates of 0.04 to 0.11 d-1 at a temperature of 4 – 5°C. Moodley et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Moodley</Author><Year>2008</Year><RecNum>1861</RecNum><record><rec-number>1861</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1861</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Moodley, L.</author><author>Steyaert, M.</author><author>Epping, E.</author><author>Middelburg, J. J.</author><author>Vincx, M.</author><author>van Avesaath, P.</author><author>Moens, T.</author><author>Soetaert, K.</author></authors></contributors><titles><title>Biomass-specific respiration rates of benthic meiofauna: Demonstrating a novel oxygen micro-respiration system</title><secondary-title>Journal of Experimental Marine Biology and Ecology</secondary-title></titles><periodical><full-title>Journal of Experimental Marine Biology and Ecology</full-title><abbr-1>J. Exp. Mar. Biol. Ecol.</abbr-1></periodical><pages>41-47</pages><volume>357</volume><number>1</number><dates><year>2008</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0022-0981</isbn><accession-num>ISI:000255310400005</accession-num><label>Moodley08</label><urls><related-urls><url><Go to ISI>://000255310400005 </url></related-urls></urls><electronic-resource-num>10.1016/j.jembe.2007.12.025</electronic-resource-num></record></Cite></EndNote>(2008) used a novel micro-respiration system and reported specific rates of 0.021 to 0.032 d-1 for intertidal (20°C) Nematoda, Ostracoda and Foraminifera over a biomass range of 0.7 to 5.2 μC ind-1. Nematodes from the Gulf of Mexico are smaller ADDIN EN.CITE <EndNote><Cite><Author>Baguley</Author><Year>2008</Year><RecNum>1918</RecNum><Prefix>~0.1μC ind-1`, </Prefix><record><rec-number>1918</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1918</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Baguley, J. G.</author><author>Montagna, P. A.</author><author>Hyde, L. J.</author><author>Rowe, G. T.</author></authors></contributors><titles><title>Metazoan meiofauna biomass, grazing, and weight-dependent respiration in the Northern Gulf of Mexico deep sea</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>2607-2616</pages><volume>55</volume><number>24-26</number><dates><year>2008</year></dates><isbn>0967-0645</isbn><accession-num>WOS:000261969100011</accession-num><urls><related-urls><url><Go to ISI>://WOS:000261969100011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.07.010</electronic-resource-num></record></Cite></EndNote>(~0.1μC ind-1, Baguley et al., 2008), but specific respiration rates are still fairly high as compared to these intertidal meiofauna. The high meiofaunal contribution to total community respiration is therefore probably also related to the comparatively high biomass-specific respiration rates that are estimated for the Gulf of Mexico. Clearly more experimental work for especially small nematodes at lower temperatures is needed to better constrain these respiration rates.The carbon sources that are consumed by meiofauna to fuel these respiration rates are detritus and prokaryotes ADDIN EN.CITE <EndNote><Cite><Author>Rowe</Author><Year>2008</Year><RecNum>1915</RecNum><Prefix>e.g.`, </Prefix><Suffix>`, this study</Suffix><record><rec-number>1915</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1915</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rowe, G. T.</author><author>Wei, C. L.</author><author>Nunnally, C.</author><author>Haedrich, R.</author><author>Montagna, P.</author><author>Baguley, J. G.</author><author>Bernhard, J. M.</author><author>Wicksten, M.</author><author>Ammons, A.</author><author>Briones, E. E.</author><author>Soliman, Y.</author><author>Deming, J. W.</author></authors></contributors><titles><title>Comparative biomass structure and estimated carbon flow in food webs in the deep Gulf of Mexico</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>2699-2711</pages><volume>55</volume><number>24-26</number><dates><year>2008</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000261969100021</accession-num><urls><related-urls><url><Go to ISI>://000261969100021</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.07.020</electronic-resource-num></record></Cite></EndNote>(e.g., Rowe et al., 2008, this study). Stable isotope tracer experiments allow direct quantification of labile food assimilation rates of amongst others meiofauna. 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ADDIN EN.CITE.DATA ( Moens et al., 2007; Franco et al., 2008; Ingels et al., 2011;), a limited (<5%) contribution to 13C uptake by metazoan meiofauna on open slope ADDIN EN.CITE <EndNote><Cite><Author>Moodley</Author><Year>2002</Year><RecNum>723</RecNum><record><rec-number>723</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">723</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Moodley, L.</author><author>Middelburg, J. J.</author><author>Boschker, H. T. S.</author><author>Duineveld, G. C. A.</author><author>Pel, R.</author><author>Herman, P. M. J.</author><author>Heip, C. H. R.</author></authors></contributors><titles><title>Bacteria and Foraminifera: Key players in a short-term deep-sea benthic response to phytodetritus</title><secondary-title>Marine Ecology Progress Series</secondary-title><alt-title>Mar. Ecol. Prog. Ser.</alt-title></titles><periodical><full-title>Marine Ecology Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><alt-periodical><full-title>Marine Ecology Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></alt-periodical><pages>23-29</pages><volume>236</volume><dates><year>2002</year></dates><accession-num>ISI:000177007100003</accession-num><label>Moodley02</label><urls><related-urls><url><Go to ISI>://000177007100003</url></related-urls></urls></record></Cite></EndNote>(Moodley et al., 2002) and abyssal plain ADDIN EN.CITE <EndNote><Cite><Author>Witte</Author><Year>2003</Year><RecNum>794</RecNum><record><rec-number>794</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">794</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Witte, U.</author><author>Wenzhofer, F.</author><author>Sommer, S.</author><author>Boetius, A.</author><author>Heinz, P.</author><author>Aberle, N.</author><author>Sand, M.</author><author>Cremer, A.</author><author>Abraham, W. R.</author><author>Jorgensen, B. B.</author><author>Pfannkuche, O.</author></authors></contributors><titles><title>In situ experimental evidence of the fate of a phytodetritus pulse at the abyssal sea floor</title><secondary-title>Nature</secondary-title><alt-title>Nature</alt-title></titles><periodical><full-title>Nature</full-title><abbr-1>Nature</abbr-1></periodical><alt-periodical><full-title>Nature</full-title><abbr-1>Nature</abbr-1></alt-periodical><pages>763-766</pages><volume>424</volume><number>6950</number><dates><year>2003</year><pub-dates><date>Aug 14</date></pub-dates></dates><accession-num>ISI:000184733900036</accession-num><label>Witte03b</label><urls><related-urls><url><Go to ISI>://000184733900036</url></related-urls></urls></record></Cite></EndNote>(Witte et al., 2003) sediments and negligible bacterivory by nematodes in a slope sediment ADDIN EN.CITE <EndNote><Cite><Author>Guilini</Author><Year>2010</Year><RecNum>1952</RecNum><record><rec-number>1952</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1952</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Guilini, K.</author><author>Van Oevelen, D.</author><author>Soetaert, K.</author><author>Middelburg, J. J.</author><author>Vanreusel, A.</author></authors></contributors><titles><title>Nutritional importance of benthic bacteria for deep-sea nematodes from the Arctic ice margin: Results of an isotope tracer experiment</title><secondary-title>Limnology and Oceanography</secondary-title></titles><periodical><full-title>Limnology and Oceanography</full-title><abbr-1>Limnol. Oceanogr.</abbr-1></periodical><pages>1977-1989</pages><volume>55</volume><number>5</number><section>1977</section><dates><year>2010</year></dates><urls></urls></record></Cite></EndNote>(Guilini et al., 2010). Irrespective of the labeled substrate or setting, meiofauna consistently show an uptake of labile 13C carbon that seems to be in imbalance with carbon requirements as estimated from biomass-specific respiration rates. This is not in contrast with the meiofaunal diet composition as inferred for the Nazaré canyon (Fig. 2), where semi-labile detritus (a carbon source not used in isotope tracer studies) is the dominant component. This dominance of semi-labile detritus in their diet would explain the low labeling of metazoan meiofauna (dominated by nematodes) in isotope tracer studies. It also agrees with Soetaert et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Soetaert</Author><Year>1997</Year><RecNum>1932</RecNum><record><rec-number>1932</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1932</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Soetaert, K.</author><author>Vanaverbeke, J.</author><author>Heip, C.</author><author>Herman, P. M. J.</author><author>Middelburg, J. J.</author><author>Sandee, A.</author><author>Duineveld, G.</author></authors></contributors><titles><title>Nematode distribution in ocean margin sediments of the Goban Spur (northeast Atlantic) in relation to sediment geochemistry</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>1671-1683</pages><volume>44</volume><number>9-10</number><dates><year>1997</year></dates><isbn>0967-0637</isbn><accession-num>WOS:A1997YF35200010</accession-num><urls><related-urls><url><Go to ISI>://WOS:A1997YF35200010</url></related-urls></urls></record></Cite></EndNote>(1997), who found a strong positive correlation between depth profiles of nematodes and organic N content and suggested that the concentration of lower quality food primarily determines nematode depth distribution. The elevated OM input in the upper canyon section combined with hydrodynamic conditions with current speeds of up to 30 – 40 cm s-1 appear to particularly favor meiofauna, whereas macro- and megafauna have a lower contribution to carbon processing as compared to open slope sediments. As a result, meiofaunal biomass in the upper canyon section rank among the highest reported in marine sediments ADDIN EN.CITE <EndNote><Cite><Author>Rex</Author><Year>2006</Year><RecNum>1839</RecNum><record><rec-number>1839</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1839</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rex, M. A.</author><author>Etter, R. J.</author><author>Morris, J. S.</author><author>Crouse, J.</author><author>McClain, C. R.</author><author>Johnson, N. A.</author><author>Stuart, C. T.</author><author>Deming, J. W.</author><author>Thies, R.</author><author>Avery, R.</author></authors></contributors><titles><title>Global bathymetric patterns of standing stock and body size in the deep-sea benthos</title><secondary-title>Marine Ecology-Progress Series</secondary-title><alt-title>Mar. Ecol.-Prog. Ser.</alt-title></titles><periodical><full-title>Marine Ecology-Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><pages>1-8</pages><volume>317</volume><dates><year>2006</year></dates><isbn>0171-8630</isbn><accession-num>ISI:000239896600001</accession-num><label>Rex06</label><urls><related-urls><url><Go to ISI>://000239896600001 </url></related-urls></urls></record></Cite></EndNote>(Rex et al., 2006), whereas macrofaunal biomass is comparatively low. Prokaryotes are responsible for the dominant part of carbon cycling and respiration in the upper canyon section (Fig. 1 and Table 5). An important pathway, also seen in the middle and lower canyon section, is deposition of semi-labile detritus, dissolution to dissolved organic carbon, to prokaryotic uptake of this DOC and subsequent prokaryote respiration. A dominance of prokaryotes in carbon cycling and respiration is commonly found in continental shelf sediments PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DYW5maWVsZDwvQXV0aG9yPjxZZWFyPjE5OTM8L1llYXI+
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ADDIN EN.CITE.DATA (Canfield et al., 1993; Piepenburg et al., 1995; Heip et al., 2001; Rowe et al., 2008). Hence, it appears that hydrodynamic conditions in the upper canyon act predominantly on carbon partitioning between faunal compartments rather than on the partitioning between pro- and eukaryotes.4.2 Middle canyon sectionSoft-sediment terraces in the middle section of the canyon experience high sedimentation rates PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5kZSBTdGlndGVyPC9BdXRob3I+PFllYXI+MjAwNzwvWWVh
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ADDIN EN.CITE.DATA (de Stigter et al., 2007; Tyler et al., 2009; Masson et al., 2010), which is accompanied by an input of organic matter of 9.30±0.71 mmol C m-2 d-1 that is comparable to the upper canyon section. These high OM inputs clearly show that the archetypical picture seen in open slope sediments that biomass, respiration and carbon processing decreases with increasing water depth does not necessarily hold for submarine canyons.With respect to the carbon partitioning within the food web, the middle canyon section seems to fall in the “metazoan-macrofaunal-uptake-dominated” category, a category that is typically found in shelf and upper slopes, with a comparatively high macrofaunal biomass ADDIN EN.CITE <EndNote><Cite><Author>Woulds</Author><Year>2009</Year><RecNum>1922</RecNum><record><rec-number>1922</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1922</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Woulds, C.</author><author>Andersson, J. H.</author><author>Cowie, G. L.</author><author>Middelburg, J. J.</author><author>Levin, L. A.</author></authors></contributors><titles><title>The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>393-402</pages><volume>56</volume><number>6-7</number><dates><year>2009</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000267780700011</accession-num><label>Woulds09</label><urls><related-urls><url><Go to ISI>://000267780700011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.10.008</electronic-resource-num></record></Cite></EndNote>(Woulds et al., 2009). An importanct discrepancy with the categorization by Woulds et al. is that faunal carbon processing in the middle canyon is not dominated by macrofauna, but by surface deposit-feeding and deposit-feeding megafauna (i.e. the holothurians Ypsilothuria bitentaculata and Molpadia musculus, respectively). The megafaunal importance is also apparent in community respiration (57%) and export of secondary production from the food web (79%). De Leo et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>De Leo</Author><Year>2010</Year><RecNum>2065</RecNum><record><rec-number>2065</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">2065</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>De Leo, F. C.</author><author>Smith, C. R.</author><author>Rowden, A. A.</author><author>Bowden, D. A.</author><author>Clark, M. R.</author></authors></contributors><titles><title>Submarine canyons: hotspots of benthic biomass and productivity in the deep sea</title><secondary-title>Proceedings of the Royal Society B-Biological Sciences</secondary-title></titles><periodical><full-title>Proceedings of the Royal Society B-Biological Sciences</full-title></periodical><pages>2783-2792</pages><volume>277</volume><number>1695</number><dates><year>2010</year></dates><isbn>0962-8452</isbn><accession-num>WOS:000280779700006</accession-num><urls><related-urls><url><Go to ISI>://WOS:000280779700006</url></related-urls></urls><electronic-resource-num>10.1098/rspb.2010.0462</electronic-resource-num></record></Cite></EndNote>(2010) reported recently for the Kaikoura Canyon (New Zealand) an extremely high biomass of 89±18 g C m-2 of megafauna (dominated by M. musculus) in low relief, muddy and accreting sediments at 900 – 1100 m of water depth. Megafaunal biomass in the middle section of the Nazaré canyon is about an order of magnitude lower (6.2 g C m-2), but still 2 – 3 orders of magnitude higher than found in open slopes at comparable depth ADDIN EN.CITE <EndNote><Cite><Author>Rex</Author><Year>2006</Year><RecNum>1839</RecNum><record><rec-number>1839</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1839</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rex, M. A.</author><author>Etter, R. J.</author><author>Morris, J. S.</author><author>Crouse, J.</author><author>McClain, C. R.</author><author>Johnson, N. A.</author><author>Stuart, C. T.</author><author>Deming, J. W.</author><author>Thies, R.</author><author>Avery, R.</author></authors></contributors><titles><title>Global bathymetric patterns of standing stock and body size in the deep-sea benthos</title><secondary-title>Marine Ecology-Progress Series</secondary-title><alt-title>Mar. Ecol.-Prog. Ser.</alt-title></titles><periodical><full-title>Marine Ecology-Progress Series</full-title><abbr-1>Mar. Ecol. Prog. Ser.</abbr-1></periodical><pages>1-8</pages><volume>317</volume><dates><year>2006</year></dates><isbn>0171-8630</isbn><accession-num>ISI:000239896600001</accession-num><label>Rex06</label><urls><related-urls><url><Go to ISI>://000239896600001 </url></related-urls></urls></record></Cite></EndNote>(Rex et al., 2006). Amaro et al. PEVuZE5vdGU+PENpdGUgRXhjbHVkZUF1dGg9IjEiPjxBdXRob3I+QW1hcm88L0F1dGhvcj48WWVh
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ADDIN EN.CITE.DATA (2010) also inferred that prokaryotes delivered <0.1% of the assimilated proteins and it was concluded that holothurians do not appear to rely on microbes for direct nutrition. This is also supported by our diet reconstruction of deposit-feeding megafauna (i.e., M. musculus), where prokaryotes play only a marginal role (Fig. 2B). Carbon partitioning with the food web of the middle canyon section at 2700 – 4000 m is comparable to much shallower shelf and upper-slope sediments, where also an important faunal contribution is typically found. The large faunal contribution in the middle canyon section is due to the comparatively high input of OM, which is quantitatively comparable to the upper canyon section. It is however unclear why canyon-specific conditions in the middle section are particularly beneficial for (surface) deposit-feeding holothurians as compared to for example macrofaunal polychaetes. The deposit-feeding megafauna consist predominantly of the holothurian head-down feeder M. musculus and there was no evidence for a specialized prokaryotic community in the guts of M. musculus that may aid in the hydrolyzation of organic matter ADDIN EN.CITE <EndNote><Cite><Author>Amaro</Author><Year>2009</Year><RecNum>1897</RecNum><record><rec-number>1897</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1897</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Amaro, T.</author><author>Witte, H.</author><author>Herndl, G. J.</author><author>Cunha, M. R.</author><author>Billett, D. S. M.</author></authors></contributors><titles><title>Deep-sea bacterial communities in sediments and guts of deposit-feeding holothurians in Portuguese canyons (NE Atlantic)</title><secondary-title>Deep-Sea Research Part I-Oceanographic Research Papers</secondary-title></titles><periodical><full-title>Deep-Sea Research Part I-Oceanographic Research Papers</full-title><abbr-1>Deep-Sea Res. I</abbr-1></periodical><pages>1834-1843</pages><volume>56</volume><number>10</number><dates><year>2009</year></dates><isbn>0967-0637</isbn><accession-num>WOS:000269469800016</accession-num><urls><related-urls><url><Go to ISI>://WOS:000269469800016</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr.2009.05.014</electronic-resource-num></record></Cite></EndNote>(Amaro et al., 2009). Other possible explanations for a strong proliferation of M. musculus in soft accreting sediments within canyons may involve a better adaptation to high sediment rates, enhanced trapping of the depositing organic matter in their feeding pits and negative feedbacks on macrofauna through, for example, predation or sediment disturbance.4.3 Lower canyon sectionThe food web structure in the lower canyon section is markedly distinct from the upper and middle sections (Fig. 1). Not only is total carbon input (1.26±0.03 mmol C m-2 d-1) about an order of magnitude lower than in the upper and middle sections, but also its partitioning within the food web differs considerably. OM input in the lower section is lower, because OM delivery from the upper and middle canyon section is less frequent, OM has been degraded during transport through the canyon and the lower canyon begins where the V-shaped valley widens into a kilometers-wide channel thereby lowering the OM input per surface area.Respiration in the lower canyon section is strongly dominated by protozoa (82% of total respiration) whereas the faunal compartments each respire <10%. These characteristics place the lower canyon section in the “respiration-dominated” category, in which most OM is respired by the prokaryotic community and the role of benthic fauna in carbon cycling is low ADDIN EN.CITE <EndNote><Cite><Author>Woulds</Author><Year>2009</Year><RecNum>1922</RecNum><record><rec-number>1922</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1922</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Woulds, C.</author><author>Andersson, J. H.</author><author>Cowie, G. L.</author><author>Middelburg, J. J.</author><author>Levin, L. A.</author></authors></contributors><titles><title>The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>393-402</pages><volume>56</volume><number>6-7</number><dates><year>2009</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000267780700011</accession-num><label>Woulds09</label><urls><related-urls><url><Go to ISI>://000267780700011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.10.008</electronic-resource-num></record></Cite></EndNote>(Woulds et al., 2009). Other sites that fall in this category are lower slope sediments and abyssal plains ADDIN EN.CITE <EndNote><Cite><Author>Woulds</Author><Year>2009</Year><RecNum>1922</RecNum><record><rec-number>1922</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1922</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Woulds, C.</author><author>Andersson, J. H.</author><author>Cowie, G. L.</author><author>Middelburg, J. J.</author><author>Levin, L. A.</author></authors></contributors><titles><title>The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions</title><secondary-title>Deep-Sea Research Part II-Topical Studies in Oceanography</secondary-title></titles><periodical><full-title>Deep-Sea Research Part II-Topical Studies in Oceanography</full-title><abbr-1>Deep-Sea Res. II</abbr-1></periodical><pages>393-402</pages><volume>56</volume><number>6-7</number><dates><year>2009</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0967-0645</isbn><accession-num>ISI:000267780700011</accession-num><label>Woulds09</label><urls><related-urls><url><Go to ISI>://000267780700011</url></related-urls></urls><electronic-resource-num>10.1016/j.dsr2.2008.10.008</electronic-resource-num></record></Cite></EndNote>(Woulds et al., 2009), suggesting that the benthic food of the lower canyon section resembles others sites at similar depth . The lower canyon section seems to be less influenced by canyon conditions as compared to the upper and middle section of the canyon.4.4 Comparison of canyon sections with network indicesThe lower carbon processing in the lower canyon is also evident in the index total system throughput ( QUOTE ), in which carbon flows are summed to obtain a measure of total food web activity ADDIN EN.CITE <EndNote><Cite><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite></EndNote>(Ulanowicz, 2004). Total system throughput does not differ significantly between the upper and middle sections (medians of 41.1 and 39.7 mmol C m-2 d-1, respectively), but is significantly lower in the lower canyon section (median of 6.7 mmol C m-2 d-1) (Table 6). Though community respiration and OM input is higher for the middle canyon section, total system throughput is slightly elevated (not significantly) in the upper canyon section. This reversal in activity measures is probably linked to the low recycling within the food web of the middle canyon as quantified with the Finn cycling index (Fig. 4B). This index summarizes the fraction of total carbon cycling that is generated by recycling processes ADDIN EN.CITE <EndNote><Cite><Author>Allesina</Author><Year>2004</Year><RecNum>1874</RecNum><record><rec-number>1874</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1874</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Allesina, S.</author><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Cycling in ecological networks: Finn's index revisited</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>227-233</pages><volume>28</volume><number>3</number><dates><year>2004</year><pub-dates><date>Jul</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000223122100008</accession-num><label>Allesina04</label><urls><related-urls><url><Go to ISI>://000223122100008 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.04.002</electronic-resource-num></record></Cite></EndNote>(Allesina and Ulanowicz, 2004). Significant differences in recycling are found between the canyon sections, with the most notable difference being low recycling in the middle canyon section. One explanation relates to the viral shunt ADDIN EN.CITE <EndNote><Cite><Author>Danovaro</Author><Year>2008</Year><RecNum>1879</RecNum><record><rec-number>1879</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1879</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Danovaro, R.</author><author>Dell'Anno, A.</author><author>Corinaldesi, C.</author><author>Magagnini, M.</author><author>Noble, R.</author><author>Tamburini, C.</author><author>Weinbauer, M.</author></authors></contributors><titles><title>Major viral impact on the functioning of benthic deep-sea ecosystems</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title><abbr-1>Nature</abbr-1></periodical><pages>1084-U27</pages><volume>454</volume><number>7208</number><dates><year>2008</year><pub-dates><date>Aug</date></pub-dates></dates><isbn>0028-0836</isbn><accession-num>ISI:000258719600030</accession-num><label>Danovaro08</label><urls><related-urls><url><Go to ISI>://000258719600030 </url></related-urls></urls><electronic-resource-num>10.1038/nature07268</electronic-resource-num></record></Cite></EndNote>(Danovaro et al., 2008), in which viral infection cause lysis of prokaryotes and the subsequent release of dissolved organic matter that is again recycled by other heterotrophic prokaryotes ADDIN EN.CITE <EndNote><Cite><Author>Van Oevelen</Author><Year>2006</Year><RecNum>1403</RecNum><Prefix>e.g.`, </Prefix><record><rec-number>1403</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1403</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Van Oevelen, D.</author><author>Middelburg, J. J.</author><author>Soetaert, K.</author><author>Moodley, L.</author></authors></contributors><titles><title>The fate of bacterial carbon in an intertidal sediment: Modeling an in situ isotope tracer experiment</title><secondary-title>Limnology and Oceanography</secondary-title><alt-title>Limnol. Oceanogr.</alt-title></titles><periodical><full-title>Limnology and Oceanography</full-title><abbr-1>Limnol. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Limnology and Oceanography</full-title><abbr-1>Limnol. Oceanogr.</abbr-1></alt-periodical><pages>1302-1314</pages><volume>51</volume><number>3</number><dates><year>2006</year></dates><label>VanOevelen06a</label><urls></urls></record></Cite></EndNote>(e.g., Van Oevelen et al., 2006a). Prokaryotes dominate carbon flows in the lower section, but this dominance is reduced in the upper and particularly the middle canyon section. If the viral-mediated shunt significantly influences the FCI, this would explain the decreasing FCI when going from the lower, upper to the middle canyon section. To examine the impact of the viral shunt on the FCI, the viral shunt was eliminated from the food web by only including the net flow from DOC to prokaryotes in the FCI calculations. Though differences in FCI remain, the FCI of the upper and lower sections drops to medians of 0.07 and 0.04, respectively, whereas the middle section is much less affected with a drop to 0.03. This exercise clearly shows that the viral shunt increases carbon recycling in benthic food webs rendering recycling to be higher in prokaryote-dominated food webs as compared to faunal-dominated food webs. The index average mutual information (AMI) gauges the developmental status of an ecosystem in the sense that while food webs develop, trophic specialization will result in higher values for AMI ADDIN EN.CITE <EndNote><Cite><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite></EndNote>(Ulanowicz, 2004). The AMI is that part of the flow diversity ADDIN EN.CITE <EndNote><Cite><Author>Ulanowicz</Author><Year>2004</Year><RecNum>1878</RecNum><Prefix>i.e. the Shannon index applied to flow diversity`, </Prefix><record><rec-number>1878</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1878</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ulanowicz, R. E.</author></authors></contributors><titles><title>Quantitative methods for ecological network analysis</title><secondary-title>Computational Biology and Chemistry</secondary-title></titles><periodical><full-title>Computational Biology and Chemistry</full-title></periodical><pages>321-339</pages><volume>28</volume><number>5-6</number><dates><year>2004</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1476-9271</isbn><accession-num>ISI:000226382900002</accession-num><label>Ulanowicz04</label><urls><related-urls><url><Go to ISI>://000226382900002 </url></related-urls></urls><electronic-resource-num>10.1016/pbiolchem.2004.09.001</electronic-resource-num></record></Cite></EndNote>(i.e. the Shannon index applied to flow diversity, Ulanowicz, 2004) that quantifies how orderly and coherently carbon ?ows are inter-connected. Since the AMI is claimed to assess the developmental status of an ecosystems it is interesting to assess whether differences in the food web structures are also reflected in the AMI index. More specifically, we had expected the less-disturbed lower canyon section to have highest AMI values with decreasing values going up-canyon. Differences in AMI between the upper and middle canyon are non-significant (Table 6), though large differences exist in environmental conditions and food web structure. The AMI is significantly lower in the lower canyon section though this section is less impacted by canyon conditions as compared to the other two sections. Tobor-Kaplon et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Tobor-Kaplon</Author><Year>2007</Year><RecNum>2070</RecNum><record><rec-number>2070</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">2070</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Tobor-Kaplon, M. A.</author><author>Holtkamp, R.</author><author>Scharler, U. M.</author><author>Doroszuk, A.</author><author>Kuenen, F. J. A.</author><author>Bloem, J.</author><author>De Ruiter, P. C.</author></authors></contributors><titles><title>Evaluation of information indices as indicators of environmental stress in terrestrial soils</title><secondary-title>Ecological Modelling</secondary-title></titles><periodical><full-title>Ecological Modelling</full-title><abbr-1>Ecol. Model.</abbr-1></periodical><pages>80-90</pages><volume>208</volume><number>1</number><dates><year>2007</year></dates><isbn>0304-3800</isbn><accession-num>WOS:000250472700010</accession-num><urls><related-urls><url><Go to ISI>://WOS:000250472700010</url></related-urls></urls><electronic-resource-num>10.1016/j.ecolmodel.2007.04.022</electronic-resource-num></record></Cite></EndNote>(2007) quantified the AMI of soil food webs that were exposed to different stress levels (i.e. pH and copper) and concluded that AMI appeared useful as an indicator of environmental stress at the ecosystem level. For the benthic food webs analyzed here however, there does not seem to be a straightforward relation between AMI and environmental stress. On the other hand, there is another important factor that influences food web structure when going down-canyon, namely the reduced OM input. To verify the usefulness of AMI as a stress indicator it is therefore necessary to compare the AMI of marine benthic food webs at similar levels of OM input, but different levels of environmental stress. In conclusion, benthic food web structures in the upper, middle and lower sections of the Nazaré canyon were shown to be influenced by the conditions in the particular canyon section. The OM input in the upper and middle canyon sections is elevated as compared to those of the surrounding open slope sediments and this resulted in a higher contribution of fauna in carbon processing as compared to open slope sites at similar water depth. The compartments that were responsible for the faunal processing were strongly influenced by conditions in the particular canyon section. In the upper canyon section, a dominance of meiofauna in faunal carbon processing was evident, whereas a high faunal contribution to carbon processing in open slope sediments is typically dominated by macrofauna. It is proposed that hydrodynamic disturbance and resulting sediment resuspension in the upper canyon shifts the balance towards the meiofauna. In contrast, the food web of the accreting sediments in the middle canyon showed a completely different pattern where carbon processing was dominated by the megafaunal holothurians. Our study confirms that accreting sediments in canyons can be hotspots of megafaunal biomass and production and megafauna can greatly influence carbon processing. The food web structure of the lower canyon section resembled that of lower slope and abyssal plain sediment, where carbon processing is dominated by prokaryotes. The influence of the canyon-specific processes seems to vanish in the deeper sections where the Nazaré canyon widens and enters the abyssal plain. In all canyon sections, a dominance of semi-labile detritus in the diet of (surface) deposit feeders is suggested. These results are supported by stable isotope tracer (for meiofauna) and gut transformation (holothurian M. musculus) studies. This study shows that elevated OM input in canyons may favor the faunal contribution to carbon processing and creating hotspots of faunal biomass and carbon processing along the continental shelf.AcknowledgementsThis research was supported by the HERMES project (contractGOCE-CT-2005-511234), funded by the European Commission’s Sixth Framework Programme under the priority “Sustainable Development, Global Change and Ecosystems”, and HERMIONE project (grant agreement n° 226354") funded by the European Community's Seventh Framework Programme (FP7/2007-2013). This is publication 5018 of the Netherlands Institute of Ecology (NIOO-KNAW), Yerseke.References ADDIN EN.REFLIST Aller, J.Y., Aller, R.C., 2004. Physical disturbance creates bacterial dominance of benthic biological communities in tropical deltaic environments of the Gulf of Papua. 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References are: 1) Garcia and Thomson ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Garcia</Author><Year>2008</Year><RecNum>1910</RecNum><record><rec-number>1910</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1910</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garcia, R.</author><author>Thomsen, L.</author></authors></contributors><titles><title>Bioavailable organic matter in surface sediments of the Nazare canyon and adjacent slope (Western Iberian Margin)</title><secondary-title>Journal of Marine Systems</secondary-title></titles><periodical><full-title>Journal of Marine Systems</full-title><abbr-1>J. Mar. 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ADDIN EN.CITE.DATA , In Press), 3) Epping et al. ADDIN EN.CITE <EndNote><Cite ExcludeAuth="1"><Author>Epping</Author><Year>2002</Year><RecNum>1736</RecNum><record><rec-number>1736</rec-number><foreign-keys><key app="EN" db-id="v5a005r0tpxrpcexfa6vtrs0wxz5wap0zsrx">1736</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Epping, E.</author><author>van der Zee, C.</author><author>Soetaert, K.</author><author>Helder, W.</author></authors></contributors><auth-address>Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, NL-1790 AB Den Burg, Netherlands. Netherlands Inst Ecol, NL-4401 EA Yerseke, Netherlands.
Epping, E, Netherlands Inst Sea Res, NIOZ, Dept Marine Chem & Geol, POB 59, NL-1790 AB Den Burg, Netherlands.</auth-address><titles><title>On the oxidation and burial of organic carbon in sediments of the Iberian margin and Nazare Canyon (NE Atlantic)</title><secondary-title>Progress in Oceanography</secondary-title><alt-title>Prog. Oceanogr.</alt-title></titles><periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></periodical><alt-periodical><full-title>Progress in Oceanography</full-title><abbr-1>Prog. Oceanogr.</abbr-1></alt-periodical><pages>399-431</pages><volume>52</volume><number>2-4</number><keywords><keyword>DEEP-SEA SEDIMENTS</keyword><keyword>BACTERIAL SULFATE REDUCTION</keyword><keyword>QUINAULT</keyword><keyword>SUBMARINE-CANYON</keyword><keyword>CONTINENTAL-MARGIN</keyword><keyword>MARINE-SEDIMENTS</keyword><keyword>PARTICULATE</keyword><keyword>MATTER</keyword><keyword>EARLY DIAGENESIS</keyword><keyword>GOBAN SPUR</keyword><keyword>HEMIPELAGIC SEDIMENTS</keyword><keyword>NORTHEAST</keyword><keyword>ATLANTIC</keyword></keywords><dates><year>2002</year></dates><isbn>0079-6611</isbn><accession-num>ISI:000176432000017</accession-num><label>Epping02</label><work-type>Review</work-type><urls><related-urls><url><Go to ISI>://000176432000017 </url></related-urls></urls><language>English</language></record></Cite></EndNote>(2002), 4) Danovaro (unpub. data), 5) biomass is Danovaro et al. (unpub. data), but biodiversity analysis in Danovaro et al. 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(unpub. data).CompartmentUpperMiddleLowerRefLabile detritus (lDet)35.8 ± 19.846.9 ± 16.410.9 ± 6.71Semi-labile detritus (sDet)5393 ± 24195114 ± 26924761 ± 23842Refractory detritus (rDet)6613766661502113Prokaryotes (Pro)4.84 ± 0.083.14 ± 0.112.79 ± 0.094Selective feeding meiofauna (MeiSF)6.80 ± 1.982.32 ± 0.772.34 ± 2.005Non-selective feeding meiofauna (MeiNF)12.42 ± 3.622.46 ± 0.820.96 ± 0.835Predatory+omnivore meiofauna (MeiPO)2.42 ± 0.700.63 ± 0.210.34 ± 0.295Surface deposit feeding macrofauna (MacSDF)0.860.52 ± 0.560.40 ± 0.716, 7Deposit feeding macrofauna (MacDF)0.392.28 ± 0.820.32 ± 0.426, 7Suspension feeding macrofauna (MacSF)0.040.73 ± 0.170.82 ± 1.016, 7Predatory+scavenging macrofauna (MacPS)17.61.02 ± 0.302.00 ± 3.576, 7Surface deposit feeding megafauna (MegSDF)21.35 ± 10.436, 7Deposit feeding megafauna (MegDF)494.7 ± 703.06, 7Table 2. Equality and inequality constraints on processes implemented for the food web models of Nazaré canyon. 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Inequality descriptionUpperMiddleLowerUnitReferenceTemperature limitation (Tlim)0.540.350.30-See textDegradation rate of lDet1 [2.74·10-3,3.29·10-2][2.74·10-3,3.29·10-2][2.74·10-3,3.29·10-2]d-11Degradation rate of sDet1[8.21·10-4, 1.51·10-2][8.21·10-4, 1.51·10-2][8.21·10-4, 1.51·10-2]d-11Degradation rate of rDet1[2.27·10-6, 8.22·10-4][2.27·10-6, 8.22·10-4][2.27·10-6, 8.22·10-4]d-11Prokaryotic C production[1.44, 7.20][0.25, 1.25][0.49, 2.44]mmol C m-2 d-12Prokaryotic growth efficiency2[0.05, 0.45][0.05, 0.45][0.05, 0.45]-3Viral lysis of prokaryotic production[0.40, 1.00][0.40, 1.00][0.40, 1.00]-4, 5Faunal maintenance respirationTlim·0.01·StockTlim·0.01·StockTlim·0.01·Stockmmol C m-2 d-16Assimilation efficiency of labile sources Mei3[0.57, 0.77][0.57, 0.77][0.57, 0.77]-6, 7Assimilation efficiency of semi-labile detritus Mei3[0.29, 0.39][0.29, 0.39][0.29, 0.39]-6, 7Net growth efficiency Mei4[0.60, 0.90][0.60, 0.90][0.60, 0.90]-7Production rate Mei5Tlim·[0.05, 0.20]Tlim·[0.05, 0.20]Tlim·[0.05, 0.20]d-17Mortality rate Mei5Tlim·[0, 0.20]d-17Feeding preference MeiSF, MacSDF and MegSDF6[50, 100][50, 100][50, 100]-See textFeeding preference MeiNSF, MacDF and MegDF6[1, 10][1, 10][1, 10]-See textFeeding preference MeiPO, MacPS and MegPS7[0.75, 1.00][0.75, 1.00][0.75, 1.00]-See textAssimilation efficiency of labile sources of Mac and Meg3[0.40, 0.75][0.40, 0.75][0.40, 0.75]-6, 7Assimilation efficiency of semi-labile detritus of Mac and Meg3[0.20, 0.38][0.20, 0.38][0.20, 0.38]-See textNet growth efficiency Mac and Meg4[0.50, 0.70][0.50, 0.70][0.50, 0.70]-6, 7Production rate Mac5Tlim·[0.01, 0.05]Tlim·[0.01, 0.05]Tlim·[0.01, 0.05]d-17, 8Mortality rate Mac5Tlim·[0.0, 0.05]Tlim·[0.0, 0.05]Tlim·[0.0, 0.05]d-17, 8Production rate Meg5Tlim·[0.0027, 0.0137]Tlim·[0.0027, 0.0137]Tlim·[0.0027, 0.0137]d-19Mortality rate Meg5Tlim·[0.0, 0.0137]Tlim·[0.0, 0.0137]Tlim·[0.0, 0.0137]d-19Prokaryotic respiration as fraction of respiration by Bac, Mei and Mac[0.60, 1.00][0.60, 1.00][0.30, 1.00]1, see TextRespiration of Bac, Mei and Mac[1.02, 4.91][0.75, 2.3][0.36, 0.90]mmol C m-2 d-11Carbon deposition from lDet_w, sDet_w, rDet_w and by MacSF[0.96, 9.4][0.64, 3.9][0.31, 1.3]mmol C m-2 d-11Burial efficiency[0.15, 0.48][0.08, 0.43][0.11, 0.36]-1DOC Efflux from sediment relative to total POC input[0, 0.10][0, 0.10][0, 0.10]-111 Degradation rate is defined as outflows from detritus compartment QUOTE divided its stock: QUOTE .2 Prokaryotic growth efficiency is defined as fraction of prokaryotic carbon uptake used for production: QUOTE .3 Assimilation efficiency is defined as fraction of ingested carbon being assimilated: QUOTE .4 Net growth efficiency is defined as: QUOTE 5 The mortality and production rates are biomass-specific.6 Feeding preference is defined as QUOTE and is 1 when food sources are consumed in their stock proportion. 7 Feeding preference is defined as fraction of total ingested met by predation.Table 3. Nomenclature of symbols used in calculation of network indices.TermDescriptionNumber of internal compartments in the network, excluding 0 (zero), QUOTE and QUOTE External source (i.e. detritus input)Useable export from the food web (i.e. secondary production)Unusable export from the food web (i.e. respiration and DOC efflux)Flow from compartment QUOTE to QUOTE where QUOTE represents the columns of the flow matrix and QUOTE the rowsFlow matrix, excluding flows to and from the externalsTotal inflows to compartment QUOTE Total outflows from compartment QUOTE Total inflows to compartment QUOTE , excluding inflow from external sourcesTotal outflows from compartment QUOTE , excluding outflow to external sourcesA negative state derivative, considered as a gain to the system pool of mobile energyA positive state derivative, considered as a loss from the system pool of mobile energyFlow into compartment QUOTE from outside the networkFlow out of the network for compartment QUOTE to compartments QUOTE and QUOTE , respectivelyThe number of species with which both QUOTE and QUOTE interact divided by the number of species with which either QUOTE or QUOTE interactIdentity matrixTable 4. Algorithms for the calculation of the network indices; see Table 3 for symbols.Index nameCodeFormulaTotal System ThroughputT..Total System ThroughflowTSTTotal System cycled throughflow QUOTE Finn’s Cycling IndexFCIAverage Mutual InformationAMITable 5. Model derived total respiration (mmol C m-2 d-1) and the biotic contributions (%) to total respiration in the food webs of the upper, middle and lower sections of the Nazaré canyon. See Table 1 for partmentUpperMiddleLowerTotal respiration4.52±0.28 5.06±0.300.86±0.02Bac70.037.981.7MeiSF6.11.08.2MeiNF11.81.53.2MeiPO2.60.41.1MacSDF0.50.170.7MacDF0.220.70.5MacSF0.020.21.25MacPS8.230.33.3MegSDF2.89MegDF54.5Table 6. Comparison of network indices calculated for the different sections of the Nazaré canyon. The numbers indicate the fraction of network values that are higher in one section as compared to another section based on a pair-wise comparison. Significant differences are in italic and highly significant differences are in bold. Network indexupper > middleupper > lowermiddle > lower0.621.001.001.000.030.000.430.930.95Figure legendsFig. 1. Food webs picturing scaled carbon flows (mmol C m-2 d-1) in the upper, middle and lower sections of the Nazaré canyon. All carbon flows are depicted in the top row (A-C), carbon flows are truncated at a maximum value of 1.5 mmol C m-2 d-1 in the middle row (D-F) and at 0.15 mmol C m-2 d-1 in the bottom row (G-I). See Table 1 for abbreviations of food web compartments. Other abbreviations are: DOC is dissolved organic carbon in the sediment, lDet_w, sDet_w and rDet_w are labile, semi-labile and refractory detritus in the water column, DOC_w is dissolved organic carbon in the water column and DIC is dissolved inorganic carbon.Fig. 2. Faunal diets in the upper (A), middle (B) and lower (C) sections of the Nazaré canyon. See Table 1 and Fig. 1 for abbreviations.Fig. 3. Fate of secondary production (%) of prokaryotes (A-C), meiofauna (D-F), macrofauna (G-I) and megafauna (J). Absolute production (mmol C m-2 d-1) is plotted above the compartment. The possible fates of this secondary production are maintenance respiration (“maint”), mortality other than predation (“mort”), export (“exp”) and predation by meiofauna (“mei”), macrofauna (“mac”) and megafauna (“meg”).Fig. 4. Box plots of the network indices total system throughput QUOTE (A), Finn cycling index QUOTE (B) and average mutual information QUOTE (C) of the upper, middle and lower sections of the Nazaré canyon.247650447675Figure 11018540436880Figure 2 Figure 33175610235 Figure 4-21780595885 Web appendixMean and standard deviation of the food web flows (mmol C m-2 d-1) of the upper, middle and lower areas of the Nazaré canyon. Empty cells indicate that the flow is not present in the food web of the respective area.Upper areaMiddle areaLower areaFlowMeanSt. dev.MeanSt. dev.MeanSt. dev.lDet_w→lDet3.88E-012.29E-018.10E-013.67E-015.88E-024.36E-02lDet_w→MacSF1.60E-038.30E-041.97E-029.02E-031.80E-029.63E-03sDet_w→sDet5.99E+007.90E-018.23E+008.44E-011.10E+003.48E-02sDet_w→MacSF3.23E-031.66E-033.54E-021.64E-023.81E-022.07E-02rDet_w→rDet1.58E+009.64E-012.02E-011.62E-015.01E-023.68E-02lDet→DOC3.85E-012.47E-015.50E-013.61E-017.93E-025.22E-02sDet→DOC1.16E+006.90E-015.15E-013.99E-015.09E-017.73E-02rDet→DOC2.52E+006.34E-011.64E+003.23E-012.26E-016.99E-02lDet→MeiSF2.83E-011.85E-019.08E-025.09E-024.25E-022.68E-02lDet→MeiNF1.09E-017.60E-022.10E-021.39E-022.50E-031.68E-03lDet→MeiPO2.62E-022.21E-024.71E-033.93E-032.10E-031.76E-03lDet→MacSDF9.97E-038.45E-034.09E-033.40E-031.62E-031.36E-03lDet→MacDF1.19E-031.01E-034.65E-033.93E-032.30E-041.90E-04lDet→MacPS6.17E-025.15E-023.13E-032.67E-035.32E-034.41E-03lDet→MegSDF8.82E-026.08E-02lDet→MegDF2.31E-011.26E-01sDet→MeiSF1.22E+002.22E-012.64E-016.10E-024.01E-014.31E-02sDet→MeiNF4.43E+004.99E-015.89E-017.03E-022.28E-013.00E-02sDet→MeiPO5.83E-023.14E-021.02E-025.40E-034.64E-032.46E-03sDet→MacSDF5.68E-021.16E-022.33E-024.74E-032.51E-024.22E-03sDet→MacDF6.10E-029.98E-032.35E-013.79E-023.36E-024.58E-03sDet→MacPS1.71E-018.80E-026.74E-033.63E-031.33E-026.20E-03sDet→MegSDF1.72E-016.78E-02sDet→MegDF6.90E+006.68E-01rDet→Burial3.05E+007.98E-013.85E+003.47E-013.35E-013.99E-02DOC→DOC_w2.16E-011.23E-012.86E-011.48E-014.71E-022.30E-02DOC→Bac5.14E+004.23E-012.96E+001.85E-011.23E+003.51E-02Bac→DIC3.18E+003.16E-011.91E+001.03E-017.05E-012.56E-02Bac→DOC1.28E+003.45E-015.38E-011.02E-014.65E-013.49E-02Bac→MeiSF4.25E-011.89E-019.33E-025.08E-025.05E-022.44E-02Bac→MeiNF1.42E-018.58E-022.13E-021.40E-022.55E-031.68E-03Bac→MeiPO2.74E-022.26E-024.70E-033.92E-032.10E-031.78E-03Bac→MacSDF1.04E-028.66E-034.12E-033.42E-031.62E-031.38E-03Bac→MacDF1.21E-031.03E-034.63E-033.93E-032.30E-041.90E-04Bac→MacPS6.40E-025.23E-023.04E-032.60E-035.19E-034.23E-03Bac→MegSDF9.06E-026.11E-02Bac→MegDF2.83E-019.85E-02MeiSF→DIC2.77E-019.63E-027.19E-022.49E-027.07E-021.48E-02MeiSF→lDet1.20E-019.33E-022.69E-022.15E-023.46E-021.47E-02MeiSF→sDet2.42E-016.92E-026.38E-021.87E-023.18E-027.75E-03MeiSF→rDet8.12E-011.57E-011.76E-014.34E-022.70E-013.28E-02MeiSF→MeiPO1.71E-011.09E-013.51E-022.14E-022.07E-027.80E-03MeiSF→MacSDF9.87E-038.27E-033.78E-033.21E-031.63E-031.38E-03MeiSF→MacDF1.19E-031.02E-034.21E-033.63E-032.30E-041.90E-04MeiSF→MacPS2.97E-011.65E-011.50E-021.07E-026.35E-021.12E-02MeiSF→MegSDF2.46E-022.01E-02MeiSF→MegDF2.69E-022.11E-02MeiNF→DIC5.34E-011.87E-017.66E-022.21E-022.78E-027.16E-03MeiNF→lDet1.36E-011.00E-012.78E-022.13E-021.62E-029.22E-03MeiNF→sDet8.30E-022.92E-021.41E-024.95E-031.68E-035.90E-04MeiNF→rDet2.93E+003.65E-013.94E-015.37E-021.55E-012.32E-02MeiNF→MeiPO2.68E-011.12E-014.15E-022.19E-021.41E-027.36E-03MeiNF→MacSDF1.01E-028.48E-033.80E-033.23E-031.59E-031.35E-03MeiNF→MacDF1.20E-031.03E-034.25E-033.69E-032.30E-042.00E-04MeiNF→MacPS7.15E-011.45E-011.59E-021.11E-021.71E-029.54E-03MeiNF→MegSDF2.58E-022.05E-02MeiNF→MegDF2.79E-022.15E-02MeiPO→DIC1.18E-013.62E-022.07E-026.21E-039.50E-032.69E-03MeiPO→lDet9.71E-026.21E-027.65E-036.17E-037.60E-034.54E-03MeiPO→sDet1.77E-014.72E-023.07E-028.07E-031.42E-023.64E-03MeiPO→rDet3.85E-022.09E-026.72E-033.58E-033.06E-031.63E-03MeiPO→MacSDF9.68E-038.29E-032.99E-032.68E-031.53E-031.32E-03MeiPO→MacDF1.19E-031.03E-033.15E-032.85E-032.20E-041.90E-04MeiPO→MacPS1.10E-016.36E-026.75E-035.65E-037.54E-034.56E-03MeiPO→MegSDF9.87E-037.07E-03MeiPO→MegDF7.77E-036.32E-03MacSDF→DIC2.21E-023.41E-038.63E-031.35E-035.75E-038.80E-04MacSDF→lDet5.55E-033.97E-032.24E-031.59E-031.46E-031.04E-03MacSDF→sDet2.58E-027.39E-039.61E-032.84E-033.93E-031.21E-03MacSDF→rDet4.21E-021.00E-021.73E-024.08E-031.91E-023.77E-03MacSDF→MacPS5.60E-034.02E-032.13E-031.56E-031.45E-031.04E-03MacSDF→Export5.63E-034.01E-032.21E-031.59E-031.46E-031.05E-03MacDF→DIC9.94E-031.52E-033.77E-025.83E-034.70E-036.70E-04MacDF→lDet2.48E-031.79E-039.77E-036.91E-031.19E-038.40E-04MacDF→sDet2.73E-039.30E-049.46E-033.23E-035.00E-041.60E-04MacDF→rDet4.68E-029.03E-031.80E-013.42E-022.60E-024.13E-03MacDF→MacPS2.57E-031.83E-038.86E-036.47E-031.17E-038.50E-04MacDF→Export2.51E-031.81E-039.99E-036.95E-031.18E-038.50E-04MacSF→DIC9.40E-041.80E-041.11E-022.06E-031.08E-021.97E-03MacSF→lDet2.50E-041.80E-042.94E-032.16E-032.82E-032.04E-03MacSF→sDet7.70E-044.80E-049.38E-035.37E-038.63E-035.52E-03MacSF→rDet2.38E-031.30E-032.59E-021.27E-022.83E-021.63E-02MacSF→MacPS2.50E-041.80E-042.84E-032.10E-032.78E-032.03E-03MacSF→Export2.50E-041.80E-042.94E-032.13E-032.84E-032.06E-03MacPS→DIC3.72E-016.33E-021.64E-022.67E-032.85E-023.83E-03MacPS→lDet1.27E-017.52E-026.36E-033.72E-031.09E-025.78E-03MacPS→sDet6.35E-011.31E-013.07E-028.29E-035.79E-029.04E-03MacPS→rDet1.22E-016.39E-024.82E-032.65E-039.50E-034.53E-03MacPS→Export1.71E-017.71E-026.17E-033.71E-031.05E-025.92E-03MegSDF→DIC1.46E-011.56E-02MegSDF→lDet1.41E-029.82E-03MegSDF→sDet1.13E-014.10E-02MegSDF→rDet1.24E-015.14E-02MegSDF→Export1.42E-029.76E-03MegDF→DIC2.75E+002.47E-01MegDF→lDet9.07E-027.22E-02MegDF→sDet2.07E-015.73E-02MegDF→rDet4.36E+004.22E-01MegDF→Export6.77E-025.22E-02 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