Community structure of the benthic macrofauna along the continental slope of Santos Basin and São Paulo plateau, SW Atlantic.
DOI:
https://doi.org/10.1590/Keywords:
Brazilian margin, Baseline study, Deep-sea, Macrobenthos, Self-Organizing MapAbstract
Continental margins usually have a high degree of environmental heterogeneity, which, in turn, promotes high
benthic biodiversity. The South-Southeast regions concentrate the most well-mapped areas of the Brazilian
continental margin regarding seafloor geomorphology and physical oceanography. However, the structure of
the soft-sediment benthic fauna of the continental slope is still unknown. Characterization and understanding
of the Brazilian continental slope biota are imperative since human activities are increasing in the last decades,
especially after the discovery of the pre-salt reservoir in Santos Basin. In this study, we aimed to establish
a baseline of the spatial distribution of the benthic macrofaunal communities regarding their latitudinal and
bathymetric patterns in the Santos Basin to support future environmental monitoring activities in the region.
As part of the Santos Project – The Santos Basin Regional Environmental Characterization (PCR-BS) –
coordinated by CENPES/PETROBRAS, a benthic oceanographic cruise was carried out in 2019. Sediment
samples were collected using a GOMEX-type box corer in 47 stations distributed in eight transects (400–
2,400 m depth range). In total, 12 additional stations (1,300–2,200 m) were defined to cover an area where oil
and gas are exploited. Our results showed that macrofaunal assemblages of the Santos Basin present strong
depth zonation related to changes in organic matter input, as well as to temperature, carbonate, and grain
size. At local scale, the northern sector stood out for having a higher abundance of macrofauna in the upper
slope (400 m) and it probably reflects the oceanographic processes and the organic enrichment caused by the
upwelling events that occur at Cabo Frio region. The zonation pattern and the dominance of some polychaete,
peracarid crustacean, and bivalve families were similar to other SE Brazilian continental margins.
References
Abdul-Jaleel, K. U. 2012. Macrobenthos of the continental
margin (200-1000m) of South Eastern Arabian Sea with
special reference to Polychaetes (phdthesis). School
of Marine Sciences Cochin, University of Science and
Technology, Kochi.
Almada, G. V. de M. B. & Bernardino, A. F. 2017.
Conservation of deep-sea ecosystems within
offshore oil fields on the Brazilian margin, SW
Atlantic. Biological Conservation, 206, 92–101. DOI:
https://doi.org/10.1016/j.biocon.2016.12.026
Almeida, A. G. de & Kowsmann, R. O. 2016. Geomorphology
of the Continental Slope and São Paulo Plateau.
In: Geology and Geomorphology (pp. 33–66).
Amsterdam: Elsevier. DOI: https://doi.org/10.1016/
b978-85-352-8444-7.50010-x
Amaral, A. C. Z., Lana, P., Fernandes, F. & Coimbra, J. 2004.
Parte I - Caracterização do Ambiente e da Macrofauna.
In: Amaral, A. C. Z. & Rossi-Wongtschowski, C. L. D. B.
(eds.), Biodiversidade bentônica das regiões sudeste
e sul do Brasil - plataforma externa e talude superior.
São Paulo: Universidade de São Paulo. DOI:
https://doi.org/10.11606/t.8.2020.tde-15122020-194333
Anderson, M. J., Gorley, R. N. & Clarke, K. R. 2008.
PERMANOVA+ for PRIMER: Guide to Software and
Statistical Methods. Auckland: PRIMER-e.
Arantes, R. C. M., Castro, C. B., Pires, D. O. &
Seoane, J. C. S. 2009. Depth and water mass zonation
and species associations of cold-water octocoral
and stony coral communities in the southwestern
Atlantic. Marine Ecology Progress Series, 397, 71–79.
DOI: https://doi.org/10.3354/meps08230
Belo, W. C. & Silveira, I. C. A. 2013. A variabilidade
vertical do oceano na Bacia de Santos. Boletim
de Geociências - Petrobras, 21(1), 39–62. DOI:
https://doi.org/10.11606/t.21.2011.tde-20042012-152310
Bernardino, A., Berenguer, V. & Ribeiro-Ferreira, V.
Bathymetric and regional changes in benthic
macrofaunal assemblages on the deep Eastern Brazilian
margin, SW Atlantic. Deep Sea Research Part I:
Oceanographic Research Papers, 111, 110–120. DOI:
https://doi.org/10.1016/j.dsr.2016.02.016
Brandini, F., Tura, P. & Santos, P. 2018. Ecosystem
responses to biogeochemical fronts in the South
Brazil Bight. Progress in Oceanography, 164, 52–62.
DOI: https://doi.org/10.1016/j.pocean.2018.04.012
Buhl-Mortensen, L., Buhl-Mortensen, P., Dolan, M.,
Dannheim, J., Bellec, V. & Holte, B. 2012. Habitat
complexity and bottom fauna composition at
different scales on the continental shelf and slope of
northern Norway. Hydrobiologia, 685(1), 191–219. DOI:
https://doi.org/10.1007/s10750-011-0988-6
Buhl-Mortensen, P., Dolan, M., Ross, R., Gonzalez-Mirelis, G.,
Buhl-Mortensen, L., Bjarnadóttir, L. & Albretsen, J.
Classification and Mapping of Benthic Biotopes
in Arctic and Sub-Arctic Norwegian Waters. Frontiers in
Marine Science, 7, 1–15. DOI: https://doi.org/10.3389/
fmars.2020.00271
Capítoli, R. & Bonilha, L. 1991. Projeto Talude. FIPEC
Relatório final. In: Vooren, C. M. (ed.) (pp. 79–92).
Rio Grande: Fundação Universitária do Rio Grande.
Carreira, R., Lazzari, L., Ceccopieri, M., Rozo, L.,
Martins, D., Fonseca, G., Vieira, D. & Massone, C. 2023.
Sedimentary provinces of organic matter accumulation
in the Santos Basin, SW Atlantic: insights from
multiple bulk proxies and machine learning analysis.
Cartes, J., Fanelli, E., López-Pérez, C. & Lebrato, M.
Deep-sea macroplankton distribution (at 400
to 2300m) in the northwestern Mediterranean in
relation to environmental factors. Journal of Marine
Systems, 113–114, 75–87. DOI: https://doi.org/10.1016/
j.jmarsys.2012.12.012
Carvalho, R., Wei, C. L., Rowe, G., Schulze, A. 2013.
Complex depth-related patterns in taxonomic and
functional diversity of polychaetes in the Gulf of Mexico.
Deep-Sea Research Part II, 66–77. DOI: 10.1016/
j.dsr.2013.07.002
Clarke, K., Somerfield, P. J. & Warwick, R. 2001.
The distribution of Antarctic marine benthic
communities (3rd ed.). Plymouth: PRIMER-e. DOI:
https://doi.org/10.1029/ar070p0219
Coelho-Souza, S., López, M., Guimarães, J., Coutinho, R. &
Candella, R. 2012. Biophysical interactions in the Cabo
Frio upwelling system, southeastern Brazil. Brazilian
Journal of Oceanography, 60(3), 353–365. DOI:
https://doi.org/10.1590/s1679-87592012000300008
Cosson, N., Sibuet, M. & Galeron, J. 1997. Community
structure and spatial heterogeneity of the deepsea macrofauna at three contrasting stations in the
tropical northeast Atlantic. Deep Sea Research Part I:
Oceanographic Research Papers, 44(2), 247–269. DOI:
https://doi.org/10.1016/s0967-0637(96)00110-0
Danovaro, R., Fabiano, M., & Della Croce, N. 1993. Labile
organic matter and microbial biomasses in deep-sea
sediments (Eastern Mediterranean Sea). Deep-Sea
Research Part II, 40(5), 953–965. DOI: https://
doi.org/10.1016/0967-0637(93)90083-F
Bathyal macrofauna from Santos Basin
Ocean and Coastal Research 2023, v71(suppl 3):e23032 16
Moura et al.
Davies, A., Duineveld, G., Lavaleye, M., Bergman, M.,
Van Haren, H. & Roberts, J. 2009. Downwelling
and deep-water bottom currents as food supply
mechanisms to the cold-water coral Lophelia
pertusa (Scleractinia) at the Mingulay Reef Complex.
Limnology and Oceanography, 54(2), 620–629. DOI:
https://doi.org/10.4319/lo.2009.54.2.0620
Davison, J., Van Haren, H., Hosegood, P., Piechaud, N. &
Howell, K. 2019. The distribution of deep-sea sponge
aggregations (Porifera) in relation to oceanographic
processes in the Faroe-Shetland Channel. Deep Sea
Research Part I: Oceanographic Research
Papers, 146, 55–61. DOI: https://doi.org/10.1016/
j.dsr.2019.03.005
De Smet, B., Pape, E., Riehl, T., Bonifácio, P., Colson, L. &
Vanreusel, A. 2017. The Community Structure of
Deep-Sea Macrofauna Associated with Polymetallic
Nodules in the Eastern Part of the Clarion-Clipperton
Fracture Zone. Frontiers in Marine Science, 4, 1–14.
DOI: https://doi.org/10.3389/fmars.2017.00103
Falcão, A., Curbelo-Fernandez, M., Borges, A.,
Filgueiras, V., Kowsmann, R. & Martins, R. 2017.
Importância ecológica e econômica da Bacia
de Campos: ambiente transicional na margem
continental do Oceano Atlântico Sudoeste. In:
Curbelo-Fernandez, M. P. & Braga, A. C. (eds.),
Ambiente Bentônico: caracterização ambiental regional
da Bacia de Campos. Rio de janeiro: Elsevier. DOI:
https://doi.org/10.1016/b978-85-352-7263-5.50001-1
Figueiredo Jr, A., Carneiro, J. & Santos-Filho, J.
Santos Basin continental shelf morphology,
sedimentology, and slope sediment distribution
Ocean and Coastal Research. Ocean and Coastal
Research, 71(3), 1–15.
Figueiredo Jr, A. & Madureira, L. 2004. Topografia,
composição, refletividade do substrato marinho e
identificação de províncias sedimentares na Região
Sudeste-Sul do Brasil. Instituto Oceanográfico USP.
Fonseca, G. & Vieira, D. 2023. Overcoming the challenges
of data integration in ecosystem studies with machine
learning pipelines: an example from the PCRBS.
Ocean and Coastal Research.
Gage, J. 2004. Diversity in deep-sea benthic macrofauna:
the importance of local ecology, the larger scale, history
and the Antarctic. Deep Sea Research Part II: Topical
Studies in Oceanography, 51(14–16), 1689–1708.
DOI: https://doi.org/10.1016/j.dsr2.2004.07.013
Gage, J., Angel, M. & Tyler, P. 1991. Deep-Sea Biology:
A Natural History of Organisms at the Deep-Sea
Floor. The Journal of Animal Ecology. Cambridge:
Cambridge University Press, 233 pp. DOI: https://doi.
org/10.2307/5527
Galéron, J., Menot, L., Renaud, N., Crassous, P.,
Khripounoff, A., Treignier, C. & Sibuet, M. 2009.
Spatial and temporal patterns of benthic macrofaunal
communities on the deep continental margin in the
Gulf of Guinea. Deep Sea Research Part II: Topical
Studies in Oceanography, 56(23), 2299–2312. DOI:
https://doi.org/10.1016/j.dsr2.2009.04.011
Galéron, J., Sibuet, M., Mahaut, M. & Dinet, A. 2000. Variation
in structure and biomass of the benthic communities
at three contrasting sites in the tropical Northeast
Atlantic. Marine Ecology Progress Series, 197, 121–137.
DOI: https://doi.org/10.3354/meps197121
Galéron, J., Sibuet, M., Vanreusel, A., Mackenzie, K.,
Gooday, A., Dinet, A. & Wolff, G. 2001. Temporal
patterns among meiofauna and macrofauna taxa
related to changes in sediment geochemistry at an
abyssal NE Atlantic site. Progress in Oceanography,
(1–4), 303–324. DOI: https://doi.org/10.1016/s0079-
(01)00059-3
Galucci, F., Fonseca, G., Vieira, D., Yaginuma, L.,
Gheller, P., Brito, S. & Corbisier, T. 2023. Predicting
large-scale spatial patterns of marine meiofauna:
implications for environmental monitoring. Ocean and
Coastal Research.
Glover, A. & Smith, C. 2003. The deep-sea floor ecosystem:
current status and prospects of anthropogenic change
by the year 2025. Environmental Conservation,
(3), 219–241. DOI: https://doi.org/10.1017/
s0376892903000225
Glover, A., Smith, C., Paterson, G., Wilson, G.,
Hawkins, L. & Sheader, M. 2002. Polychaete species
diversity in the central Pacific abyss: local and
regional patterns, and relationships with productivity.
Marine Ecology Progress Series, 240, 157–170. DOI:
https://doi.org/10.3354/meps240157
Grassle, F. J. & Maciolek, N. 1992. Deep-Sea Species
Richness: Regional and Local Diversity Estimates from
Quantitative Bottom Samples. The American Naturalist,
(2), 313–341. DOI: https://doi.org/10.1086/285329
Gray, J. & Elliott, M. 2009. Ecology of Marine Sediments
(2nd ed.). New York: Oxford University Press. DOI:
https://doi.org/10.1093/oso/9780198569015.001.0001
Guimarães, L. M., França, E. J. D., Arruda, G. N. de &
Albergaria-Barbosa, A. C. R. de. 2020. Historical inputs
of polycyclic aromatic hydrocarbons in the preserved
tropical estuary of the Itapicuru River, Bahia, Brazil.
Marine Pollution Bulletin, 156, 111218. DOI: https://doi.
org/10.1016/j.marpolbul.2020.111218
Hessler, R. & Jumars, P. 1974. Abyssal community analysis
from replicate cores in the central North Pacific.
Deep Sea Research and Oceanographic Abstracts,
(3), 185–209. DOI: https://doi.org/10.1016/0011-
(74)90058-8
Hughes, D. & Gage, J. 2004. Benthic metazoan
biomass, community structure and bioturbation
at three contrasting deep-water sites on the
northwest European continental margin. Progress in
Oceanography, 63(1–2), 29–55. DOI: https://doi.org/
1016/j.pocean.2004.09.002
Ingole, B., Sautya, S., Sivadas, S., Singh, R. & Nanajkar, M.
Macrofaunal community structure in the western
Indian continental margin including the oxygen
minimum zone. Marine Ecology, 31(1), 148–166. DOI:
https://doi.org/10.1111/j.1439-0485.2009.00356.x
Jumars, P., Dorgan, K. & Lindsay, S. 2015. Diet of Worms
Emended: An Update of Polychaete Feeding Guilds.
Annual Review of Marine Science, 7(1), 497–520. DOI:
https://doi.org/10.1146/annurev-marine-010814-020007
Kohonen, T. 2001. Self-organizing maps. Berlin: SpringerVerlag.
Bathyal macrofauna from Santos Basin
Ocean and Coastal Research 2023, v71(suppl 3):e23032 17
Moura et al.
Kokesh, B. S., Kidwell, S. M., Tomašových, A., & Walther, S. M.
Detecting strong spatial and temporal variation
in macrobenthic composition on an urban shelf using
taxonomic surrogates. Marine Ecology Progress Series,
, 13–30. DOI: https://doi.org/10.3354/meps13932
Lavrado, H., Omena, E. & Bernardino, A. 2010. Macrofauna
bentônica do talude continental e cânions da Bacia de
Campos. In: Lavrado, H. P. & Brasil, A. C. S. (eds.),
Ambiente Bentônico: caracterização ambiental regional
da Bacia de Campos, Atlântico Sudoeste. Rio de
Janeiro: Elsevier. DOI: https://doi.org/10.1016/b978-85-
-7263-5.50009-6
Lavrado, H., Omena, E. & Bernardino, A. 2017. Ambiente
Bentônico: caracterização ambiental regional da Bacia
de Campos, Atlântico Sudoeste. In: Falcão, H. P., A. P. C.,
Lavrado (ed.). Rio de Janeiro: Elsevier.
Levin, L. & Gooday, A. 2003. Ecosystems of the Deep
Oceans. In: Tyler, P. A. (ed.) (Vol. 28, pp. 111–178).
Amsterdam: Elsevier.
Levin, L. & Sibuet, M. 2012. Understanding Continental
Margin Biodiversity: A New Imperative. Annual Review
of Marine Science, 4(1), 79–112. DOI: https://doi.org/
1146/annurev-marine-120709-142714
Mahiques, M., Silveira, I. C. A. da, Mello e Sousa, S. H. de &
Rodrigues, M. 2002. Post-LGM sedimentation on
the outer shelf-upper slope of the northernmost
part of the São Paulo Bight, southeastern Brazil.
Marine Geology, 181(4), 387–400. DOI: https://doi.
org/10.1016/S0025-3227(01)00225-0
Menot, L., Sibuet, M., Carney, R., Levin, L., Rowe, G.,
Billett, D., Poore, G., Kitazato, H., Vanreusel, A.,
Galéron, J., Lavrado, H., Sellanes, J., Ingole, B. &
Krylova, E. 2010. New Perceptions of Continental
Margin Biodiversity. In: Mcintyre, A. D. (ed.), Life in the
World’s Oceans: Diversity, Distribution, and Abundance
(pp. 79–102). Hoboken: Wiley-Blackwell. DOI: https://
doi.org/10.1002/9781444325508.ch5
Mohriak, W. 2003. Geologia, Tectônica e Recursos Minerais
do Brasil. In: Bizzi, L., Schobbenhaus, C., Vidotti, R., &
Gonçalves, J. H. (eds.), Geologia, Tectônica e Recursos
Minerais do Brasil (pp. 375–413). Brasília, DF:
CPRM. DOI: https://doi.org/10.5724/gcs.05.25.0375
Moreira, D., Dalto, A., Figueiredo Jr, A.,
Valerio, A., Detoni, A., Bonecker, A., Signori, C.,
Namiki, C., Sasaki, D., Pupo, D., Silva, D., Kutner, D.,
Duque-Castaño, D., Marcon, E., Gallotta, F., Paula, F.,
Galucci, F., Roque, G., Campos, G., Fonseca, G.,
Mattos, G., Lavrado, H., Silveira, I., Costa, J., SantosFilho, J., Carneiro, J., Moreira, J., Rozo, L., Araujo, L.,
Lazzari, L., Silva, L., Michelazzo, L., Fernandes, L.,
Dottori, M., Araújo Jr, M., Chuqui, M., Ceccopieri, M.,
Borges-Silva, M., Kampel, M., Bergo, N., Silva, P.,
Tura, P., Moura, R., Romano, R., Martins, R., Carreira, R.,
Toledo, R., Bonecker, S., Disaró, S., Rodrigues, S.,
Corbisier, T., Vicente, T., Paiva, V., Pellizari, V., Belo, W.,
Brandini, F. & Souza, S. 2023. Multidisciplinary scientific
cruises for environmental characterization in the
Santos Basin – methods and sampling design.
Oliveira, D., Cordeiro, L. & Carreira, R. 2013.
Characterization of organic matter in cross-margin
sediment transects of an upwelling region in the
Campos Basin (SW Atlantic, Brazil) using lipid
biomarkers. Biogeochemistry, 112(1–3), 311–327.
DOI: https://doi.org/10.1007/s10533-012-9726-z
Pabis, K., Sobczyk, R., Siciński, J., Ensrud, T. &
Serigstadt, B. 2019. Natural and anthropogenic
factors influencing abundance of the benthic
macrofauna along the shelf and slope of the Gulf of
Guinea, a large marine ecosystem off West Africa.
Oceanologia, 62(1), 83–100. DOI: https://doi.org/
1016/j.oceano.2019.08.003
Palma, M., Quiroga, E., Gallardo, V., Arntz, W., Gerdes, D.,
Schneider, W. & Hebbeln, D. 2005. Macrobenthic
animal assemblages of the continental margin off
Chile (22° to 42°S). Journal of the Marine Biological
Association of the United Kingdom, 85(2), 233–245.
DOI: https://doi.org/10.1017/s0025315405011112h
Puerta, P., Johnson, C., Carreiro-Silva, M., Henry, L.,
Kenchington, E., Morato, T., Kazanidis, G., Rueda, J.,
Urra, J., Ross, S., Wei, C., González-Irusta, J., ArnaudHaond, S. & Orejas, C. 2020. Influence of Water
Masses on the Biodiversity and Biogeography
of Deep-Sea Benthic Ecosystems in the North
Atlantic. Frontiers in Marine Science, 7, 1–25. DOI:
https://doi.org/10.3389/fmars.2020.00239
R Core Team. 2021. R: A language and environment for
statistical computing. R Foundation for Statistical
Computing. Vienna, Austria.
Ramirez-Llodra, E., Brandt, A., Danovaro, R., De Mol, B.,
Escobar, E., German, C., Levin, L., Martinez Arbizu, P.,
Menot, L., Buhl-Mortensen, P., Narayanaswamy, B.,
Smith, C., Tittensor, D., Tyler, P., Vanreusel, A. &
Vecchione, M. 2010. Deep, diverse and definitely
different: unique attributes of the world’s largest
ecosystem. Biogeosciences, 7(9), 2851–2899. DOI:
https://doi.org/10.5194/bg-7-2851-2010
Rex, M. A. 1981. Community Structure in the Deep-Sea
Benthos. Annual Review of Ecology and Systematics,
(1), 331–353. DOI: https://doi.org/10.1146/annurev.
es.12.110181.001555
Rex, M., Etter, R., Morris, J., Crouse, J., Mcclain, C.,
Johnson, N., Stuart, C., Deming, J., Thies, R. &
Avery, R. 2006. Global bathymetric patterns of standing
stock and body size in the deep-sea benthos. Marine
Ecology Progress Series, 317, 1–8. DOI: https://doi.org/
3354/meps317001
Saaedi, H., Bernardino, A., Shimabukuro, M., Falchetto, G. &
Sumida, P. 2019. Macrofaunal community structure and
biodiversity patterns based on a wood-fall experiment
in the deep South-west Atlantic. Deep Sea Research
Part I: Oceanographic Research Papers, 145, 73–82.
DOI: https://doi.org/10.1016/j.dsr.2019.01.008
Sallorenzo, I. A. 2013. Caracterização da macroinfauna
bentônica da plataforma continental da Bacia de
Campos: questões metodológicas (candthesis).
Universidade Federal Fluminense, Niterói.
Seeliger, U., Odebrecht, C. & Castelo, J. 1998.
Os ecossistemas costeiro e marinho do extremo sul do
Brasil. Editora Ecoscientia. Rio Grande: Ecoscientia.
Shields, M. & Blanco-Perez, R. 2013. Polychaete abundance,
biomass and diversity patterns at the Mid-Atlantic
Ridge, North Atlantic Ocean. Deep Sea Research
Part II: Topical Studies in Oceanography, 98, 315–325.
DOI: https://doi.org/10.1016/j.dsr2.2013.04.010
Bathyal macrofauna from Santos Basin
Ocean and Coastal Research 2023, v71(suppl 3):e23032 18
Moura et al.
Shields, M. & Hughes, D. 2009. Large-scale variation in
macrofaunal communities along the eastern Nordic
Seas continental margin: A comparison of four
stations with contrasting food supply. Progress in
Oceanography, 82(2), 125–136. DOI: https://doi.org/
1016/j.pocean.2009.05.001
Shimabukuro, M., Carrerette, O., Alfaro-Lucas, J.,
Rizzo, A., Halanych, K. & Sumida, P. 2019. Diversity,
Distribution and Phylogeny of Hesionidae (Annelida)
Colonizing Whale Falls: New Species of Sirsoe and
Connections Between Ocean Basins. Frontiers in
Marine Science, 6(478), 1–26. DOI: https://doi.org/
3389/fmars.2019.00478
Shimabukuro, M., Rizzo, A. E., Alfaro-Lucas, J. M.,
Fujiwara, Y. & Sumida, P. Y. G. 2017. Sphaerodoropsis
kitazatoi, a new species and the first record of
Sphaerodoridae (Annelida: Phyllodocida) in SW
Atlantic abyssal sediments around a whale carcass.
Deep-Sea Research Part II, 146, 18–26. DOI: https://
doi.org/10.1016/j.dsr2.2017.04.003
Silveira, I. C. A., Bernardo, P. S., Lazaneo, C. Z.,
Amorim, J. P. M., Borges-Silva, M., Martins, R. C.,
Santos, D. M. C., Dottori, M., Belo, W. C., Martins, R. P.
Guerra, L. A. A. & Moreira, D. L. 2023. Oceanographic
conditions of the continental slope and deep waters
in Santos Basin: the SANSED cruise (winter 2019).
Ocean and Coastal Research, 71(3), 1–12. DOI: http://
doi.org/10.1590/2675-2824071.2206icas
Stuart, C. & Rex, M. 2009. Bathymetric patterns of deepsea gastropod species diversity in 10 basins of the
Atlantic Ocean and Norwegian Sea. Marine Ecology,
(2), 164–180. DOI: https://doi.org/10.1111/j.1439-
2008.00269.x
Sumida, P., Alfaro-Lucas, J., Shimabukuro, M., Kitazato, H.,
Perez, J., Soares-Gomes, A., Toyofuku, T., Lima, A.,
Ara, K. & Fujiwara, Y. 2016. Deep-sea whale fall fauna
from the Atlantic resembles that of the Pacific Ocean.
Scientific Reports, 6(1), 22139. DOI: https://doi.org/
1038/srep22139
Sumida, P. & Pires-Vanin, A. 1997. Benthic associations
of the shelf-break and upper slope off Ubatuba-SP,
South-eastern Brazil. Estuarine Coastal and Shelf
Science, 44, 779–784.
Sumida, P., Yoshinaga, M., Ciotti, A. & Gaeta, S. 2005.
Benthic response to upwelling events off the SE
Brazilian coast. Marine Ecology Progress Series, 291,
–42. DOI: https://doi.org/10.3354/meps291035
Sumida, P., Yoshinaga, M., Madureira, L. A. S.-P. &
Hovland, M. 2004. Seabed pockmarks associated with
deepwater corals off SE Brazilian continental slope,
Santos Basin. Marine Geology, 207(1–4), 159–167.
DOI: https://doi.org/10.1016/j.margeo.2004.03.006
Taghon, G., Nowell, A. & Jumars, P. 1980. Induction of
Suspension Feeding in Spionid Polychaetes by High
Particulate Fluxes. Science, 210(4469), 562–564. DOI:
https://doi.org/10.1126/science.210.4469.562
Thistle, D. 2003. Ecosystems of the deep oceans. In:
Tyler, P. A. (ed.) Ecosystems of the World (Vol. 28).
Amsterdam: Elsevier BV.
Tselepides, A., Papadopoulou, N.-, Podaras, D.,
Plaiti, W. & Koutsoubas, D. 2000. Macrobenthic
community structure over the continental margin
of Crete (South Aegean Sea, NE Mediterranean).
Progress in Oceanography, 46(2–4), 401–428. DOI:
https://doi.org/10.1016/s0079-6611(00)00027-6
Tyler, P. 2003. Ecosystems of the Deep Ocean. Planets,
Stars and Nebulae Studied with Photopolarimetry
(Vol. 28). Amsterdam: Elsevier, 434–443 pp. DOI:
https://doi.org/10.2307/j.ctt2050vsn.29
Valentin, J. 2001. The Cabo Frio Upwelling System,
Brazil. In: Seeliger, U. & K. B. (ed.) Coastal Marine
Ecosystems of Latin America (Vol. 144, pp. 97–104).
Berlin: Springer-Verlag.
Van Der Grient, J. & Rogers, A. 2021. Environmental
influence on the distribution of polychaete families
and feeding guilds in benthic communities of the
Grand Banks and Flemish Cap (NW Atlantic).
Deep Sea Research Part I: Oceanographic Research
Papers, 171, 103498. DOI: https://doi.org/10.1016/
j.dsr.2021.103498
Viana, A., Faugeres, J., Kowsmann, R., Lima, J.,
Caddah, L. & Rizzo, J. 1998. Hydrology, morphology
and sedimentology of the Campos continental
margin, offshore Brazil. Sedimentary Geology,
(1–4), 133–157. DOI: https://doi.org/10.1016/
s0037-0738(97)00090-0
Vieira, D. & Fonseca, G. 2022. iMESc: An Interactive Machine
Learning App for Environmental Science. Accessed:
https://zenodo.org/record/6484391#.ZF1R63bMLDc
Washburn, T., Menot, L., Bonifácio, P., Pape, E.,
Błażewicz, M., Bribiesca-Contreras, G., Dahlgren, T.,
Fukushima, T., Glover, A., Ju, S., Kaiser, S., Yu, O. &
Smith, C. 2021. Patterns of Macrofaunal Biodiversity
Across the Clarion-Clipperton Zone: An Area Targeted
for Seabed Mining. Frontiers in Marine Science, 8,
–22. DOI: https://doi.org/10.3389/fmars.2021.626571
Wickham, H. 2016. Ggplot2: Elegant Graphics for Data
Analysis. Journal of the Royal Statistical Society
Series A: Statistics in Society. New York: Springer
Verlag, 245–246 pp.
Würzberg, L. W., Peters, J., Schüller, M. & Brandt, A. 2011.
Diet insights of deep-sea polychaetes derived from fatty
acid analyses. Deep Sea Research Part II, 58(1/2), 153–
DOI: https://doi.org/10.1016/j.dsr2.2010.10.014
Yasuhara, M. & Danovaro, R. 2014. Temperature impacts
on deep-sea biodiversity. Biological Reviews, 91(2),
–287. DOI: https://doi.org/10.1111/brv.12169