MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 ATLAS OF THE DEEP-SEA FAUNA OF THE SOUTHWESTERN ATLANTIC Exploring the Unseen: A Scientific Collection of Unique Deep-Sea Benthic Species Documented During TotalEnergies’ Offshore Studies in the Santos and Campos Basins, Brazil.
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms2 | TotalEnergies The data presented in this atlas were generated through studies conducted under the environmental permitting process regulated by the Brazilian Environment Agency (IBAMA). As such, its regulatory framework serves as a driver for scientific research, contributing to the advancement of knowledge on marine biodiversity. This atlas presents a curated col- lection of deep-sea benthic fauna documented through underwa- ter inspections and environmental characterization studies associa- ted with TotalEnergies’ offshore operations in the Santos and Cam- pos Basins, Southwestern Atlantic. These barely unknown organisms and colonies reveal the extraordinary biodiversity of abyssal ecosystems and highlight the need to advance scientific knowledge in these remote environments. Developed as part of the Biodiversity Action Plan (BAP) of TotalEnergies EP Brasil, this work reflects the company’s approach to transparency, research, and marine conservation. By sharing these findings, the intention is to encourage collaboration, inspire new research, and support efforts that contribute to the protection of deep-sea ecosystems.
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 3 INTRODUCTION The deep ocean remains one of Earth’s most mysterious and least explored environments. In the Southwestern Atlantic, its abyssal zones harbor extraordinary biodiversity, much of it still unknown to science and difficult to observe. Through studies conducted during offshore operations in the Santos and Campos Basins (Lapa, C-M-541, S-M-1711/1815), TotalEnergies has documented benthic taxa, including new records and potential new taxa. Each identification in this atlas was validated by specialized taxonomists, ensuring scientific rigor and reliability. By making the unseen visible, we seek to advance marine research, promote knowledge sharing, foster collaboration between science and industry, and contribute to the long-term preservation of deep-sea ecosystems. THIS ATLAS SEEKS TO: • Share valuable data with the scientific community and the public; • Highlight the ecological importance of deep-sea ecosystems; • Reaffirm TotalEnergies’ commitment to promote biodiversity and transparency, as outlined in its EP Brazilian affiliate’s Biodiversity Action Plan (BAP).
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms4 | TotalEnergies Echinodermata Araeosoma cf. fenestratum Atelecrinidae Benthodytes cf. typica Benthopecten sp. Brisingidae sp. 1 Deima validum validum Echinosigra sp. Freyllidae sp. 1 Hygrosoma cf. petersii Nymphaster arenatus Ophiurida INDEX Porifera Euplectella Hyalonematidae Cnidaria Acanella Anthoptilum Ceriantharia Schizopatidae Umbellula Annelida Admetellinae 5 6 13 14 15 16 17 18 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 19 20 21 12 Crustacea Acutiserolis coineauae Aristeidae Glyphocrangon Heterocarpus Nematocarcinus Galatheoidea Mollusca Graneledone sp. Muusoctopus oregonae Ophiotheutis agassizii Actinopterygii Aldrovandia affinis Allocytus verrucosus Antimora rostrata B. grallator B. phenax Bathysaurus cf. ferox Halosauropsis macrochir Holcomycterus squamosus Ipnops murrayi Macrouridae Synaphobranchidae Peniagone sp. Persicullida Phormosoma placenta Plinthaster dentatus Psychropotes depressa Psychropotes semperiana 7 8 9 10 11
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 5 BIBLIOGRAPHY Lopes. D. A. 2006. Biodiver- sidade de esponjas do mar profundo brasileiro: taxo- nomia da subclasse Hexas- terophora (Porifera, Hexac- tinellidae). Dissertação de Mestrado, Museu Nacional/ Universidade Federal do Rio de Janeiro, 165 p. Saito, T.; Uchida, I. and M. Takeda. 2002. Skeletal grow- th of the deep-sea hexac- tinellid sponge Euplectella oweni, and host selection by the symbiotic shrimp Spon- gicola japonica (Crustacea: Decapoda: Spongicolidae). J. Zool., Lond. 258, 521- 529. HEXACTINELLIDA/ EUPLECTELLIDAE PORIFERA Euplectella Owen, 1841 DIAGNOSIS Columnar or vase-like sponge attached to the sea bottom, thin body wall with several lateral oscula (Lopes, 2006). Appearance of a completely sealed mesh basket skeleton with a flexible phase during development until rigid forms after full growth (Figure 1). DISTRIBUTION Species of the genus are abundant in the North Atlantic and some for the Southwestern Atlantic Ocean, from the middle to the lower slope, between 1,598 and 2,366 m depth along the Brazilian basins. The records in the present investigation: Santos Basin from 1,145 to 1,502 (S-M-1815), from 1,483 to1,847 (S-M- 1711) and (Lapa), from 2,120 to 2,177 m depth and Campos Basin, (C-M-541), from 2,869 to 2,996 m depth (Figure 2). COMMENTS The sophistication of its body wall mistook medieval Europeans who considered it a Chinese glass craftwork. Specimens of Euplectella are usually found in symbiotic association with the shrimp Spongicola spp., and its and spicules have been studied because of their light transmission properties (Saito et al., 2002). Figure 2. Records of the deep-sea sponge Euplectella spp. in the four investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin, and C-M-541, in Campos Basin. Figure 1. An Euplectella spp. specimen observed in Santos Basin at 2,130 m depth. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms6 | TotalEnergies AMPHIDISCOSIDA/ HEXACTINELLIDA PORIFERA Hyalonematidae Gray, 1857 BIBLIOGRAPHY Tabachnick, K.R., Menshe- nina, L.L. (2002). Family Hyalonematidae Gray, 1857. In: Hooper, J.N.A., Van So- est, R.W.M., Willenz, P. (eds) Systema Porifera. Springer, Boston, MA. https://doi. o rg / 1 0.1 0 07/978 -1 - 4 61 5 - 0747-5_127 Tabachnick, K. R., Menshe- nina, L.L, Lopes, D.A., Hajdu, E. 2009. Two new Hyalone- ma species (Hyalonemati- dae: Amphidiscosida) from eastern and south-eastern Brazil, and further Hexacti- nellida (Porifera) collected from seamounts off south- -eastern Brazil by the RV ‘Marion Dufresne’ MD55 ex- pedition. Journal of the Mari- ne Biological Association of the United Kingdom, 89(6), 1243–1250. DIAGNOSIS Generally whip-like, rarely spindle-like in overall shape. Ovoid body, its rhachis is thickest at base. In grown specimens, basalia gathered in a compact twisted tuft (adapted from Tabachnick & Menshenina, 2002) (Figure 1). DISTRIBUTION The family is well-known worldwide in the deep sea, with numerous representatives. Records for Southwestern Atlantic Ocean between 600 and 835 m depth (Tabachnick et al., 2009). Records in the present investigation include: Santos Basin from 1,197 to 1,467 m (S-M-1815), from 1,492 to 1,616 m (S- M-1711), and from 2,113 to 2,179 m (Lapa), and Campos Basin from 2,862 to 2,990 m depth (C-M-541) (Figure 2). COMMENTS Hyalonematidae is usually found isolated on soft bottoms often in association with zoanthids growing along its tuft. Figure 1. A Hyalonematidae sponge observed in Santos Basin at 2,132 m depth. Figure 2. Records of Hyalonematidae in the four investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin, and C-M-541, in Campos Basin. 6 | TotalEnergies Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 7 OCTOCORALLIA/ SCLEROALCYONACEA/ KERATOISIDIDAE CNIDARIA Acanella Gray, 1870 BIBLIOGRAPHY Heestand Saucier, E., Sajjadi, A. & France, S.C. 2017. A ta- xonomic review of the genus Acanella (Cnidaria: Octoco- rallia: Isididae) in the North Atlantic Ocean, with des- criptions of two new species. Zootaxa, 4323 (3), 359–390. Morrissey, D., Gordon, J. D., Saso, E., Bilewitch J. P., Taylor, M. L., Hayes, V., McFa- dden, C. S., Quattrini, A. M., Allcock, L. 2023. Bamboo- zled! Resolving deep evo- lutionary nodes within the phylogeny of bamboo corals (Octocorallia: Scleralcyona- cea: Keratoisididae). Molecu- lar Phylogenetics and Evolu- tion Volume 188, pp. 1-11. Watling, L. Heestand Sau- cier, E. & France, S. C. 2022. Towards a revision of the bamboo corals (Octoco- rallia): Part 4, delineating the family Keratoisididae. Zoota- xa, 5093, pp. 337-375. DIAGNOSIS Bush-like colonies branched dichotomously or in whorls, one to six branches at each node. Colonies moderate in size when anchored in soft substrate by lobate holdfast. Internodes solid, short, and covered with thin coenenchyme. Polyps non- retractile, and perpendicular or angled towards the branch. Polyps armed with needles and/or rods, pharyngeal wall with small thorny stars or short rods (Heestand Saucier et al., 2017) (Figure 1). DISTRIBUTION The genus Acanella Gray, 1870 exhibits a wide-ranging global distribution and can be found on both hard and soft substrata. Bathymetric range, from 300 to 2,875 m depth. Records in the present investigation include: Santos Basin at 1,497 m depth (S- M-1815) and from 1,618 to 1,643 m depth (S-M-1711) (Figure 2). COMMENTS The genus Acanella Gray, 1870 is commonly encountered in deep-sea sedimentary regions, and the identification of these organisms relies primarily on observable traits, such as an articulated axis consisting of an alternating sequence of proteinaceous nodes and calcium carbonate internodes (Morrisey et al., 2023). This feature is a characteristic of the Keratoisididae family. The axis exhibits long white internodes and short dark brown nodes at the base of the colony (Heestand Saucier et al., 2017), which can be easily discerned through images. Furthermore, colonies may possess a calcareous root- like base. The bamboo coral family Keratoisididae displays significant genetic and morphological diversity, as demonstrated by several strongly supported clades in molecular phylogenetic analysis. Moreover, there is compelling evidence indicating the existence of numerous taxa awaiting classification at either the genus or species levels (Watling et al., 2022). Within the North Atlantic region, one confirmed species of Acanella Gray, 1870 has been documented, namely Acanella arbuscula (Johnson, 1862). There are no published records of the genus Acanella for the Brazilian coast, even though its presence is commonly observed in imaging surveys. Figure 1. The deep-sea bamboo coral Acanella spp. observed in Santos Basin at 1,497 m depth. Figure 2. Records of the deep-sea bamboo coral Acanella spp. at 1,497 m depth (S-M-1815) and from 1,618 to 1,643 m depth (S-M-1711), in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms8 | TotalEnergies OCTOCORALLIA/ SCLEROALCYONACEA/ PENNATULOIDEA CNIDARIA Anthoptilum Kölliker, 1880 Figure 2. Records of the deep-sea sea pen Anthoptilum sp. in two of the investigated areas: S-M-1711 and Lapa, in Santos Basin. BIBLIOGRAPHY Arantes, R. C. M, Castro, C. B.; Me- deiros, M. S. & Pires, D. O. 2010. An- thoptilum murrayi Kölliker,1880. In: Lavrado, H. P.; Brasil, M. S. (Orgs). Biodiversidade da região oceânica profunda da Bacia de Campos: Me- gafauna e ictiofauna demersal. Rio de Janeiro, 122-123 p. Schejter, L., Acuña, F.H., Garese, A., Cordeiro-Silva, R.T. & Pérez, C.D. (2018) Sea Pens (Cnidaria: Pennatu- lacea) from Argentine Waters: new distributional records and first re- port of associated anemones. Pan- -American Journal of Aquatic Scien- ces, 13(4), 292–301. Williams, G.C. (1995). Living genera of sea pens (Coelenterata: Octoco- rallia: Pennatulacea): illustrated key and synopses. Zoological Journal of the Linnean Society 113: 93–140. Williams, G.C.; Alderslade, P. (2011). Three new species of pennatulacean octocorals with the ability to attach to rocky substrata (Cnidaria: Antho- zoa: Pennatulacea). Zootaxa 3001: 33–48. Williams, G.; Tracey, D.; Mackay, E. (2014). Pennatulacea (Sea Pens) Descriptions for the New Zealand Region. A field guide of common- ly sampled New Zealand Sea pens including illustrations highlighting technical terms and sea pen mor- phology. New Zealand Aquatic En- vironment and Biodiversity Report No. 132. 22 p DIAGNOSIS Colonies elongate and whip-like, rachis symmetry is bilateral throughout. Axis present throughout length of colony, round to somewhat quadrangular in transverse section. Autozooids numerous and elongate, arranged short in oblique along the rachis. Anthocodiae nonretractile with calyces absent Sclerites absent except for ovals in the interior of the stalk. Color: orange red (Williams, 1995; Williams et al., 2014) (Figure 1). DISTRIBUTION Sea pens are widely distributed across the globe, spanning the Pacific, Indian, and Atlantic Oceans, and extending as far north as the Arctic Circle. Their habitat ranges from depths of 240 to 2500 m (Willians et al., 2014) Records in the present investigation include Santos Basin at 1,850 m depth (S-M-1711) and from 2,116 to 2,188 m depth (Lapa) (Figure 2). COMMENTS Sea pens are found worldwide, primarily in soft-bottom habitats (Williams, 2011). However, some species are capable of dwelling in hard-bottom environments through a suction cup-like expansion at the proximal end of the peduncle (Williams & Alderslade, 2011). The Anthoptillidae family consists of five species, two of which have a global presence: A. grandiflorum and A. murrayi. A. grandiflorum is frequently observed in the deep waters of the Southwestern Atlantic Ocean, particularly in Argentine waters (Schejter et al., 2018). Furthermore, Arantes et al. (2010) reported records of A. murrayi in deep-sea waters off Brazil. Figure 1. The deep-sea whip Anthoptilum sp. observed at 1,850 m depth in Santos Basin. 8 | TotalEnergies Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 9 HEXACORALLIA CNIDARIA Ceriantharia Perrier, 1983 BIBLIOGRAPHY Forero Mejia, A.C.; Molo- dtsova, T.; Östman, C.; Ba- vestrello, G. & Rouse, G.W. 2020. Molecular phylogeny of Ceriantharia (Cnidaria: Anthozoa) reveals non- -monophyly of traditionally accepted families. Zool. J. Linn. Soc., 190, 397–416. Molodtsova T.M. 2009. Ce- riantharia (Cnidaria) of the Gulf of Mexico. In: Felder DL, Camp DK, eds. Gulf of Mexico, origins, waters and biota. Vol. 1 Biodiversity. Corpus Cristi: Texas A&M University Press, 365–367. Molodtsova, T. N.; Moskalenko, V. N.; Lipukhin, E. V.; Antokhina, T. I.; Anane- va, M. S.; Simakova, U. V. 2023. Cerianthus lloydii (Ce- riantharia: Anthozoa: Cnida- ria): New Status and New Perspectives. Biology, 12(9). Stampar S.N.; Maronna M.M.; Kitahara M.V.; Reimer J.D & Morandini A.C. 2014. Fast-Evolving mitochondrial DNA in Ceriantharia: A re- flection of Hexacorallia pa- raphyly? PLoS ONE. DIAGNOSIS Cylindrical body usually worm-like. The aboral side of the polyp is hollow and dull, opened by a pore. The column, oral disc and tentacles are smooth, and their tissues are simple. The tentacles are divided into marginal tentacles on the edge of the oral disc, and labial tentacles, which are around the mouth (Stampar et al., 2014). DISTRIBUTION The subclass Ceriantharia Perrier, 1893 has a cosmopolitan distribution, occurring from shallow to deep-sea areas. Records in the present investigation include Santos Basin from 1,1,193 to 1,497 m depth (S-M-1815), from 1,612 to 1,850 m depth (S-M-1711) and from 2,127 to 2,135 m depth (Lapa), and Campos Basin from 2,983 to 2,977 m depth (C-M-541) (Figure 2). COMMENTS Ceriantharians are solitary anemone-like anthozoans known from the intertidal zone to the abyssal depths of all oceans, comprising species commonly referred to as tube anemones (Molodtsova, 2009). Tube anemones have a uniform appearance, with an elongated muscular column, a flattened oral disc with two crowns of numerous simple tentacles (Mejia et al., 2019) Adult Ceriantharia are benthic and usually inhabit long, fibrous, mucous tubes, and these tubes are composed of the special cnidae type and encrusted with surrounding particles (Molodtsova et al., 2023). The most important aspects of tube-dwelling anemone taxonomy are related to the shape and arrangement of the mesenteries and cnidome (Stampar et al., 2016). Ceriantharians are common benthic cnidarians inhabiting the deep-sea floor of Brazil, but there are still insufficient taxonomical works for the proper identification of the group. Figure 1. The tube anemone from the subclass Ceriantharia Perrier, 1893 observed in Santos Basin at 1,498 m depth. Figure 2. Records of the tube anemone from the subclass Ceriantharia Perrier, 1893 in the four investigated areas S-M-1815, S-M-1711 and Lapa, in Santos Basin and C-M-541 in Campos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms10 | TotalEnergies CNIDARIA Figure 1. The deep-sea black coral from the family Schizopathidae Brook, 1889 observed in Santos Basin at 1,500 m depth. Figure 2. Record of Schizopatidae in one of the four investigated areas: S-M- 1815, in Santos Basin. HEXACORALLIA/ ANTHIPATHARIA Schizopathidae Brook, 1889 BIBLIOGRAPHY Molodtsova, T. N. 2014. De- ep-sea fauna of European seas: An annotated species check-list of benthic inver- tebrates living deeper than 2000 m in the seas borde- ring Europe. Antipatharia. Invertebrate Zoology, v. 11, p. 3-7. 2014. Molodtsova, T.; Opresko, D. 2023. World List of An- tipatharia. Schizopathi- dae Brook, 1889. Accessed through: World Register of Marine Species at: https:// www.marinespecies.org/ a p h i a . p h p? p = t a x d e t a i l - s&id=266964 Opresko, D. M. 2002. Re- vision of the Antipatharia (Cnidaria: Anthozoa). Part II. Schizopathidae. Zoologis- che Mededelingen, v. 76, p. 411-442. DIAGNOSIS Polyps elongated transversely, with six primary and four secondary mesenteries. Corallum monopodial or branched, stem and branches pinnulate (Figure 1). Pinnules simple or complexly subpinnulate. Spines triangular, laterally compressed, simple or rarely with multiple bifurcations at apex, subequal in size around circumference of axis or larger on one side (Opresko, 2002). DISTRIBUTION The family Schizopathidae Brook, 1889 has a cosmopolitan distribution; it occurs in the Pacific, Indian, Atlantic, and Southern oceans. Its bathymetric range extends from 100 to 8,600 meters in depth. Records in the present investigation include Santos Basin at 1,362 and 1,500 m depth (S-M-1815) (Figure 2). COMMENTS The family Schizopathidae Brook, 1889 comprises corals with elongated polyps in the transverse axis, and 10 mesenteries (6 primary and 4 secondary) (Opresko, 2002). The family includes 13 genera (Molodtsova & Opresko, 2023) with distinct colony morphologies. It encompasses the deepest depth records reached by a black coral: 8,600 and is therefore considered a predominantly deep-sea group (Molodtsova, 2014). 10 | TotalEnergies Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 11 CNIDARIA OCTOCORALLIA/ SCLEROALCYONACEA/ UMBELLULIDAE Umbellula Gray, 1870 Figure 1. Umbellula cf. thomsonii juvenile colony observed in Santos Basin at 1,466 m depth. Figure 2. Records of the sea pen Umbellula in three investigated areas: S-M- 1815, S-M-1711, and Lapa, in Santos Basin. DIAGNOSIS Autozooids arranged in a terminal cluster at the end of a long slender stalk. Symmetry of rachis is bilateral. The conspicuous axis is present throughout the colony, round to x-shape in cross-section. Polyp leaves absent. Autozooids restricted to a terminal cluster containing one to forty polyps. Anthocodiae non-retractile, calyces absent. Siphonozooids present on the rachis at the base of the autozooids and below the terminal cluster on the upper part of the stem. Sclerites may be present in the tentacles, walls of the autozooids, and peduncle (Williams, 1995; López- Gonzales & Drewery, 2022). (Figure 1). DISTRIBUTION The genus Umbellula Gray, 1870 has a cosmopolitan distribution; this species can be found in the Atlantic, Indian, Pacific, Arctic, and Southern Oceans. Its bathymetric range extends from 200 to 6100 meters in depth. Records in the present investigation include Santos Basin at 1,619 m depth (S-M-1711), 1,466 and 1,498 (S-M-1815), and 2,135 m depth (Lapa) (Figure 2). COMMENTS Among octocorals, colonies of Umbellula Gray, 1870 are some of the most instantly recognizable forms (López- Gonzales & Drewery, 2022), owing to their umbrella-shaped colony morphology. Colonies of this sea pen have a proximal peduncle that anchors the colony in soft substrates, followed by an elongated and thin stalk, and a distal crown of polyps (Williams ,1995). Castro & Medeiros (2001) made the first record of the species Umbellula thomsonii (Kölliker, 1874) in Brazilian waters. Species from the genus Umbellula are commonly found inhabiting deep-sea soft bottoms in the Southeast sedimentary basins. In images, it is common to observe juvenile colonies with a short or not-yet-developed stalk (Figure 1). BIBLIOGRAPHY Castro, C. B. & Medeiros, M. S. 2001. Brazilian Pennatula- cea (Cnidaria: Octocorallia). Bulletin of the Biological Society of Washington, Wa- shington, 10:140-159. López-González, P.J. & Drewery, J. (2022) When dis- tant relatives look too alike: a new family, two new genera and new species of deep- -sea Umbellula-like sea pens (Anthozoa, Octocorallia, Pennatulacea). Invertebrate Systematics, 36(3), 199–225. Williams GC (1995) Living genera of sea pens (Coelen- terata: Octocorallia: Penna- tulacea): illustrated key and synopses. Zoological Jour- nal of the Linnean Society 113, 93–140. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms12 | TotalEnergies ERRANTIA/ APHRODITIFORMIA/ POLYNOIDAE ANNELIDA Admetellinae Uschakov 1977 DIAGNOSIS Body arched dorsally, flattened to convex ventrally, longer than wider, wider in the middle segments, and tapering on both ends (oval or fusiform body). More than 50 segments. Elytra present, whitish, more than 20 pairs, located on segments: 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, then on every third segment to the end of the body (Figure 1). Prostomium bilobed, two eyes present. Middle antenna dorsally (with variable position); lateral antennae emerging from the frontal border of two long tapered styles; antennal scales dorsally to lateral antennae. The tentacular segment is achaetous, with two long tentacular, tapered, cirri. The second segment with reduced (rudimentary) parapodia. Nephridial papillae present, shorter than ventral cirri. DISTRIBUTION The subfamily has a wide distribution, occurring in all oceans, except the Arctic Ocean. Species belonging to this group occur in depths ranging from 400 m to 5,760 m, being found mostly in depths from 600-700 m (McIntosh, 1885; Chamberlin, 1919; Pettibone, 1954; Levenstein, 1978; OBIS, 2023). The record in the present investigation is the first for the South Atlantic, in the Santos Basin, at a depth of 1,647m (S-M-1815) (Figure 2). COMMENTS Admetellinae polychaetes are considered rare animals, with few specimens sampled to date. Most of the records and behavior of these worms are known from deep-sea imagery. The morphology of the three anterior segments are key characters for the proper identification of these worms, however, due to the fusiform shape of the body and the combination of several segments and pairs of elytra, it is possible to identify at the subfamily level. A few genera within Polynoidae possess more than 50 segments and more than 20 elytra pairs, all others (excluding members of Admetellinae) have long segments of the same width through the body. There are only two genera within this subfamily, Admetella McIntosh, 1885 and Bathyadmetella Pettibone, 1967, distinguished by the position of middle antennae, tentacular sheath, and notochaetae. Figure 1. A polynoid worm, subfamily Admetellinae, observed in Santos Basin at 1,645 m depth. Figure 2. Record of the Admetellin worm (Polynoidae) in one of the investigated areas: S-M-1815, in Santos Basin. BIBLIOGRAPHY Uschakov, Pavel Vladimiro- vich 1977. Phylogenetic rela- tionships in the family Polynoi- dae (Polychaeta), in D.J. Reish and K. Fauchald eds., Essays on Polychaetous Annelids in Memory of Dr. Olga Hart- man: Los Angeles, University of Southern California, Allan Hancock Press: 29-38. McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. “Challen- ger” during the years 1873-76. Challenger Reports, 12: 1-554. Pettibone, M. H. 1967. Some bathyal polynoids from the central and northeastern Pa- cific (Polychaeta: Polynoidae). Proceedings of the United States National Museum, 121 (3575): 1–15. Chamberlin RV. 1919. The An- nelida Polychaeta. Memoirs of the Museum of Comparative Zoology at Harvard College 48: 1-514. Levenstein RJ. 1978. Annelida (Polychaeta) from the deep- -waters of the Pacific region of Antarctic. Trudy Instituta Okeanologii. Akademiya nauk SSSR 113: 73–87. OBIS, Ocean Biodiversity In- formation System 2023. Ad- metellinae Uschakov, 1977. Available at: https://obis.org/ taxon/237616 (Accessed: 1 Oc- tober 2023). 12 | TotalEnergies Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 13 ISOPODA/ SEROLIDAE CRUSTACEA Acutiserolis coineauae Albuquerque & Costa, 2008 BIBLIOGRAPHY Albuquerque E.L. and Cos- ta, I.A. 2008. A new species of Acutiserolis Brandt, 1988 (Isopoda: Serolidae) from the deep sea of southern Brazil. Nauplius 16: 43–53. Moura, M. and Serejo, S.S. 2010. In: Lavrado, H.P. & Bra- sil, A.C.S. Biodiversidade da Região Oceânica Profunda da Bacia de Campos: Mega- fauna e Ictiofauna Demersal. 164 – 165 p. DIAGNOSIS Broad body, about 1.2 times wider than long, reaching 58 mm in length, dorsoventrally flattened with six posteriorly pointed coxal plates; purplish in color; eyes seleniform small (Figure 1). DISTRIBUTION West Atlantic Ocean (Moura and Serejo, 2010). Brazilian coast, off Bahia to off Rio de Janeiro between 1,190 and 1,555 m depth. Campos Basin, between 1,064 – 1,630 m (Albuquerque and Costa, 2008). The record in the present investigation: Santos Basin from 1,148 to 1,497 m depth (S-M-1815), from 1,487 and 1,847 m depth (S-M- 1711), and at 2,122 m depth (Lapa), and Campos Basin at 2,861 m depth (C-M-541) (Figure 2). COMMENTS Deep-Sea epibenthic species. Figure 1. Acutiserolis coineauae Albuquerque & Costa, 2008 observed in the Santos Basin at 1,641 m depth. Figure 2. Records of Acutiserolis coineauae Albuquerque & Costa, 2008 in three of the investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms14 | TotalEnergies DECAPODA/ ARISTEIDAE CRUSTACEA Aristeidae Wood-Mason in Wood-Mason & Alcock, 1891 BIBLIOGRAPHY Cardoso, I. A., Alves-Junior, F., & Almeida, T. R. G. 2020. Catalogue of Typical Deep- -Sea Decapod Fauna from Brazilian Waters. Deep-Sea Pycnogonids and Crustace- ans of the Americas, 499- 584. Tavares C.R, Serejo C.S. 2007. Taxonomy of Aristei- dae (Dendrobranchiata: Pe- naeoidea) from the central coast of Brazil, collected by the Revizee program, be- tween 19° and 22° S. Zoota- xa 1585:1–44. DIAGNOSIS Redish in color; rostrum sexually dimorphic in several genera, being elongate in females and juvenile males, short in adult males; usually only 3 dorsal rostral/postrostral spines; lacking ventral spines. Carapace lacking postorbital and pterygostomian spines; antennal and branchiostegal spines always present; postantennal spine rarely present and hepatic spines often lacking; cervical and postcervical sulci sometimes present, most often reaching dorsal midline, or almost absent and visible only laterally. Eye with optic calathus bearing mesial tubercle; ocular scale and styliform projection lacking. Antennule with prosartema reduced to setose boss; flagella unequal, dorsal flagellum short, flattened for most of its length, inserted proximally on third segment; ventral flagellum sexually dimorphic in some genera (Tavares and Serejo, 2007) (Figure 1). DISTRIBUTION Aristeids are typically pelagic, in depths ranging from 60 to 5420 meters. In Brazil found between 61 and 5,419m depth (Cardoso et. al. 2020). The records in the present investigation: Santos Basin at 1,487 m depth (S-M-1711) and from 2,120 to 2,155 m depth (Lapa), and Campos Basin (C-M-541) between 2865,72 to 2996,44 m depth. COMMENTS Aristeids shrimp normally living in waters outside the continental shelf to abyssal depths, with one species having been found at a depth of more than 5,400 m. In Brazilian waters, the family Aristeidae Wood-Mason in Wood-Mason & Alcock, 1891 is represented by seven genus and eight species: Aristaeomorpha foliacea (Risso, 1827) (61m – 1,300m), Aristaeopsis edwardsiana (Johnson, 1867) (200m – 1,850 m), Aristeus antennatus (Risso, 1816) (150m – 1,799m), Aristeus antillensis A. Milne-Edwards and Bouvier, 1909 (200 to 2,057m), Hemipenaeus carpenteri Wood-Mason, 1891 (900m – 3,900m), Hepomadus tener Smith, 1884 (765 – 5,400 m), Cerataspis monstrosus Gray, 1828 (752 – 5,413 m) and Plesiopenaeus coruscans (Wood-Mason in Wood-Mason and Alcock, 1891) (900m – 2,367m) (Cardoso et. al. 2020). Figure 1. Aristeid shrimp observed in the Santos Basin at 1,487 m depth. Figure 2. Records of Aristeid shrimps in three of the investigated areas: S-M-1711 and Lapa, at Santos Basin, and C-M-541, at Campos Basin. Atlas of Benthic Fauna 14 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 15 CRUSTACEA BIBLIOGRAPHY Cardoso, I. A., Alves-Junior, F., & Almeida, T. R. G. 2020. Catalogue of Typical Deep- -Sea Decapod Fauna from Brazilian Waters. Deep-Sea Pycnogonids and Crustace- ans of the Americas, 499- 584. Serejo, C.S.; YOUNG, P. S.; Cardoso, I. A.; Tavares, C.; Rodrigues, C.; Almeida, T. C. 2007. Abundância, diver- sidade e zonação dos Crus- táceos no talude da costa central do Brasil (11º - 22º S) coletados pelo Progra- ma REVIZEE/Score Cen- tral.: prospecção pesqueira. In: Costa, P.A.S.; Olavo, G.; Martins, A.S.. (Eds.). Biodi- versidade da fauna marinha profunda na costa central brasileira. Rio de Janeiro: Museu Nacional. p. 133-162 (Série Livros n24). DIAGNOSIS Cream white with reddish nuances in color; integument rigid; dorsally and ventrally flattened or concave, armed laterally with pairs of teeth; carapace strongly sculptured with ridges, grooves, spines and tubercles (Figure 1). DISTRIBUTION Indian Ocean, western Pacific Ocean, eastern Atlantic Ocean and western Atlantic Ocean. Glyphocrangonids inhabit deep water regions, with a bathymetric distribution between 150 and 6,370 m depth. In Brazil found from Amapá do Rio de Janeiro (Serejo et. al. 2007), between 256 and 3,219 m depth (Cardoso et. al. 2020). The records in the present investigation: Santos Basin (S-M-1711), between 1,488 and 1,852 m depth, (S-M-1815) between 1,363 and 1,501 m depth. COMMENTS The genus Glyphocrangon A. Milne-Edwards, 1881 is the sole accepted genus within the family Glyphocrangonidae Smith, 1884. In Brazilian waters, the genus is represented by eight species: Glyphocrangon aculeata A. Milne- Edwards, 1881 (443m to 1,760m depth), Glyphocrangon alispina Chace, 1939 (548m to 2,094 m depth), Glyphocrangon aurantiaca Holthuis, 1971 (410m to 733m depth), Glyphocrangon longirostris (Smith, 1882) (1,280m to 2,500 m depth), Glyphocrangon neglecta Faxon, 1896 (365m to 1,050 m depth), Glyphocrangon nobilis A. Milne-Edwards, 1881 (410m to 2,150 m depth), Glyphocrangon sculpta (Smith, 1882) (1,645m to 3,219 m) and Glyphocrangon spinicauda A. Milne-Edwards, 1881 (256 m to 692 m depth) (Serejo et. al. 2007 and Cardoso et. al. 2020). Figure 1. Glyphocrangon spp. A. Milne-Edwards, 1881 observed in the Santos Basin at 1,487 m depth. Figure 2. Records of Glyphocrangon spp. A. Milne-Edwards, 1881 in the two Investigated areas: S-M-1711 and S-M-1815, at Santos Basin. DECAPODA/ GLYPHOCRANGONIDAE Glyphocrangon A. Milne-Edwards, 1881 Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms16 | TotalEnergies CRUSTACEA BIBLIOGRAPHY Cardoso, I. A., Alves-Junior, F., & Al- meida, T. R. G. 2020. Catalogue of Typical Deep-Sea Decapod Fau- na from Brazilian Waters. Deep-Sea Pycnogonids and Crustaceans of the Americas, 499-584. Chan, B., Lin, I., Shih, T., & Chan, T. (2008). Bioluminescent emissions of the deep-water pandalid shrimp, Heterocarpus sibogae De Man, 1917 (Decapoda, Caridea, Pandalidae) under laboratory conditions. Crus- taceana, 81(3), 341-350. https://doi. org/10.1163/156854008783564064 Crosnier, A. and Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale, 19:1-409 Rego, A. B. B., Cardoso, I. A. 2010. Ta- xonomy and distribution of shrimps genus Heterocarpus A. Milne-E- dwards, 1881 collected on the Brazilian slope by the Program REVIZEE – cen- tral score. Nauplius 18(2):117–128 Tavares, M. S. 1999. New species and new records of deep-water caridean shrimps from the South Atlantic Oce- an (Crustacea, Decapoda). Zoosyste- ma 21:671–677 DIAGNOSIS Cream white and reddish in color; Carapace with one or more well-marked longitudinal carinae Rostrum elongated, laterally compressed; antennular flagella simple, two distal segments not arising side by side from penultimate segment (Figure 1). DISTRIBUTION Shrimps of the genus Heterocarpus A. Milne- Edwards, 1881 can be found in all tropical and some temperate deep seas, in depths varying from 73 to 2.834 m (Chace, 1985). In Brazil found between 278 and 1,718 m depth (Tavares, 1999; Rego & Cardoso, 2010 and Cardoso et. al. 2020). The records in the present investigation: Santos Basin (S-M-1711), at 1,485 m depth. COMMENTS Adults are benthic, while juveniles and larvae are often caught in the water column (Crosnier and Forest, 1973). Many Heterocarpus shrimp are known to use bioluminescence as a defense mechanism, spitting it at predators. (Chan et. al. 2008). Despite having benthic habits, they migrate through the water column at night (Rego & Cardoso, 2010). In Brazilian waters five species have been recorded: Heterocarpus dorsalis Bate, 1888 (185 m – 1,400 m); Heterocarpus ensifer A. Milne-Edwards, 1881 (88 m - 885 m); Heterocarpus inopinatus Tavares, 1999 (150 m – 1,718 m); Heterocarpus laevigatus Bate, 1888 (366 m - 966 m) and Heterocarpus oryx A. Milne-Edwards, 1881 (118 m - 1,774 m) (Cardoso et. al. 2020). Figure 1. Heterocarpus spp. A. Milne-Edwards, 1881 observed in the Santos Basin at 1,485 m depth. Figure 2. Record of Heterocarpus spp. A. Milne-Edwards, 1881 in one of the investigated areas: S-M-1711, in Santos Basin. DECAPODA/ PANDALIDAE Heterocarpus A. Milne-Edwards, 1881 Atlas of Benthic Fauna 16 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 17 CRUSTACEA BIBLIOGRAPHY Cardoso, I.A., Burukovsky, R.N. 2014. Nematocarcinus Milne Edwards, 1881 (Crus- tacea, Decapoda) from Sou- thwestern Atlantic, including the Southern Mid-Atlantic Ridge area. Zootaxa, 3887: 437–458. Cardoso, I. A., Alves-Junior, F., & Almeida, T. R. G. 2020. Catalogue of Typical Deep- -Sea Decapod Fauna from Brazilian Waters. Deep-Sea Pycnogonids and Crustace- ans of the Americas, 499- 584. Chace, F.A., Jr 1986. The Ca- ridean shrimps (Crustacea: Decapoda) of the Alba- tross Philippine Expedition, 1907–1910. Part 4: Families Oplophoridae and Nema- tocarcinidae. Smithsonian Contributions to Zoology No. 432: i-iv, 1-82. DIAGNOSIS Redish in color; rostrum rather elongated, laterally compressed; carapace relatively smooth, not strongly carved; simple antennular flagella, without accessory branches; two pairs of anterior pereiopods with chelae, three posterior pairs exceptionally long (Figure 1). DISTRIBUTION Nematocarcinids are typically demersal, in depths ranging from 51 to 3931 meters (Chace 1986). In Brazil found between 190 and 3,075 m depth (Cardoso et. al. 2020). The records in the present investigation: Santos Basin from 1,148 to 1,499 m depth (S-M-1815) and from 1,483 to 1,849 m depth (S-M-1711), and Campos Basin at 3,002 m depth (C-M-541) (Figure 2). COMMENTS In Brazilian waters, the genus Nematocarcinus A. Milne- -Edwards, 1881 is represented by four species: Nemato- carcinus gracilipes Filhol, 1884 (190 m - 1860 m), Nema- tocarcinus parvus Burukovsky, 2000 (600 m – 1,050 m), Nematocarcinus rotundus Crosnier Forest, 1973 (421 m – 1,629 m) and Nematocarcinus tenuipes Spence-Bate, 1888 (630 m – 3,075 m). Figure 1. Blue antimoral, spp. A. Milne-Edwards, 1881 observed in the Campos Basin at 3,001 m depth. Figure 2. Records of Nematocarcinus A. spp. Milne-Edwards, 1881 in three of the investigated areas: S-M-1815 and S-M-1711, in Santos Basin, and C-M-541 in Campos Basin. DECAPODA/ NEMATOCARCINIDAE Nematocarcinus A. Milne-Edwards, 1881 Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms18 | TotalEnergies ANOMURA/ GALATHEOIDEA CRUSTACEA BIBLIOGRAPHY Rodrigues, C. 2010. In: Lavra- do, H.P. & Brasil, A.C.S. Bio- diversidade da Região Oce- ânica Profunda da Bacia de Campos: Megafauna e Ictio- fauna Demersal. 210 – 215 p. Rodrigues, C. and Serejo, C.S. 2010. In: Lavrado, H.P. & Brasil, A.C.S. Biodiversidade da Região Oceânica Pro- funda da Bacia de Campos: Megafauna e Ictiofauna De- mersal. 216 – 217 p. DIAGNOSIS Body depressed, symmetrical, more or less ovate; rostrum usually well developed and prominent, variable in form; eyes well developed. Abdomen bent upon itself but not folded against thorax. (Figure 1). DISTRIBUTION Species of Galatheoidea are found from intertidal waters to the abyssal plains of the deep sea. Circumglobal distribution. The records in the present investigation: Santos Basin (S-M-1711 and S-M-1815), between 1,483 and 1,850 m depth (Figure 2). COMMENTS Galatheids are mostly relatively small; some exhibiting spectacular colors, especially those that live as commensals on brightly colored invertebrates. They are a significant component of the benthic fauna of coral reefs. Some species are found in deeper waters of the continental slope, exceeding 2,000 meters deep. Figure 1. Galatheoidea observed in the Santos Basin at 1,483 m depth. Figure 2. Records of Galatheoidea in three of the investigated areas: S-M- 1815, S-M-1711, and Lapa, in Santos Basin. Galatheoidea Samouelle, 1819 Atlas of Benthic Fauna 18 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 19 MOLLUSCA BIBLIOGRAPHY Haimovici, M; Costa, P.A.S.; Santos, R.A.; Martins, A.S.; Olavo, G. 2007. Composição de espécies, distribuição e abundância de cefalópodes do talude da região central do Brasil. In: Costa, P.A.S.; Olavo, G.; Martins, A.S. (Eds.) Biodiversidade da fauna ma- rinha profunda na costa cen- tral brasileira. Rio de Janeiro: Museu Nacional. p.109-132 (Série Livros n.24). MolluscaBase eds. (2023). MolluscaBase. Graneledo- ne Joubin, 1918. Accessed through: World Register of Marine Species at: https:// www.marinespecies.org/ a p h i a . p h p? p = t a x d e t a i l - s&id=157018 on 2023-11-30 Voight, J.R. 2000. A deep- -sea octopus (Graneledone cf. boreopacifica) as a shell- -crushing hydrothermal vent predator. Journal of Zoology, 252(3): 335-341. DIAGNOSIS Spherical to slightly ovoid mantle with a rough appearance. Purplish in color. Styles present, not mineralized. Arms moderate to long, length 2 to 4 times the length of the mantle. Webs approximately equal in depth. Suction cups in single file (Figure 1). DISTRIBUTION The genus is circumglobal in deep sea habitats. On the Brazilian coast, there is only one record in the Campos Basin, at a depth of 1653m (Haimovici et. al. 2007). The record in the present investigation: Santos Basin (S-M-1815), at 1,462 m (Figure 2). COMMENTS The genus Graneledone contains 10 valid species (MolluscaBase, 2023). There is only one record of an unidentified species belonging to the genus Graneledone along the Brazilian coast. (Haimovici et. al. 2007). Figure 1. Graneledone sp. observed in the Santos Basin at 1,462 m depth. Figure 2. Records of Graneledone sp. in one of the investigated areas: S-M- 1815, in Santos Basin. OCTOPODA/ ENTEROCTOPODIDAE Graneledone Joubin, 1948 Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms20 | TotalEnergies MOLLUSCA BIBLIOGRAPHY Gleadall, I.G. 2004. Some old and new genera of octopus. Interdisciplinary Information Sciences, 10:99-112. Haimovici, M; Costa, P.A.S.; Santos, R.A.; Martins, A.S.; Olavo, G. 2007. Composição de espécies, distribuição e abundância de cefalópodes do talude da região central do Brasil. In: Costa, P.A.S.; Olavo, G.; Martins, A.S. (Eds.) Biodiversidade da fauna ma- rinha profunda na costa cen- tral brasileira. Rio de Janeiro: Museu Nacional. p.109-132 (Série Livros n.24). MolluscaBase eds. (2023). MolluscaBase. Muusoctopus Gleadall, 2004. Accessed through: World Register of Marine Species at: https:// www.marinespecies.org/ a p h i a . p h p? p = t a x d e t a i l - s&id=527126 on 2023-11-30 DIAGNOSIS Globose mantle, broad head, relatively large eyes. Long, slender, cylindrical arms; arms 1 and 2 longer than 3 and 4. Suckers arranged in two moderately spaced rows. Large funnel, free for half its length. Skin lacks well-defined patch-and-groove system, displaying white and reddish hues. (Figure 1). DISTRIBUTION Muusoctopus oregonae (Toll, 1981) occurs in the West Atlantic Ocean, Caribbean and Brazil: Campos Basin. Found between 488m and 1,066m depth (Haimovici et. al. 2007). The record in the present investigation: Santos Basin (S- M-1711), at 1,850 m (Figure 2). COMMENTS The genus Muusoctopus contains 27 valid species (MolluscaBase, 2023). Muusoctopus oregonae (Toll, 1981) is the only species recorded for Brazil (Haimovici et. al. 2007). Muusoctopus species have no ink sac. Figure 1. M.oregonae (Toll, 1981) observed in the Santos Basin at 1,850 m depth. Figure 2. Records of M.oregonae (Toll, 1981) in one of the investigated areas: S-M-1711, in Santos Basin. OCTOPODA/ ENTEROCTOPODIDAE Muusoctopus oregonae (Toll, 1981) Atlas of Benthic Fauna 20 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 21 MOLLUSCA BIBLIOGRAPHY Haimovici, M; Costa, P.A.S.; Santos, R.A.; Martins, A.S.; Olavo, G. 2007. Composição de espécies, distribuição e abundância de cefalópodes do talude da região central do Brasil. In: Costa, P.A.S.; Olavo, G.; Martins, A.S. (Eds.) Biodiversidade da fauna ma- rinha profunda na costa cen- tral brasileira. Rio de Janeiro: Museu Nacional. p.109-132 (Série Livros n.24). Villanueva R.; Collins, M.A.; Sanchez, P.; Voss, N.A. 2002. Systematics, distribution and biology of the cirra- te octopods of the genus Opisthoteuthis (Mollusca, Cephalopoda) in the Atlan- tic Ocean, with description of two new species. Bulletin of Marine Science, v. 71, n. 1, p. 933-986. DIAGNOSIS Semi-gelatinous body in a bell-shaped form. Saccular, gelatinous mantle, about one third of the total length, nearly twice as long as wide, with a rounded posterior end. Moderately long fins, broadly rounded, positioned laterally between the eyes and the apex of the mantle. Head broader than the mantle, without a visible neck region. Pallial aperture narrow, enveloping the base of the funnel. Elongated funnel. Arms semi-gelatinous, subequal, moderately long (Figure 1). DISTRIBUTION Opisthoteuthis agassizii A. E. Verrill, 1883, is recognized as being limited to the West Atlantic Ocean and the Ca- ribbean (Villanueva et. al. 2002), in the Campos Basin at depths between 1001 m to 2075 m (Haimovici et. al., 2007). The record in the present investigation: Santos Basin (S-M-1815), between 1452m and 1497m depth (Figure 2). COMMENTS The dumbo octopus exhibits four distinct movement modes: crawling, taking off, swimming by flapping their ear-like fins, and a passive mode characterized by umbrella-style drifting. They employ these methods either simultaneously or individually. They exhibit deep pelagic habits and scan the ocean floor for food. Common prey items include crustaceans, mollusks, and worms. OCTOPODA/ OPISTHOTEUTHIDAE Opisthoteuthis cf. agassizii A. E. Verrill, 1883 Figure 1. Opisthoteuthis cf. agassizii A. E. Verrill, 1883 observed in the Santos Basin at 1,151 m depth. Figure 2. Records of Opisthoteuthis cf. agassizii A. E. Verrill, 1883 in one of the investigated areas: S-M-1815, in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms22 | TotalEnergies ECHINOIDEA/ ECHINOTHURIOIDA/ ECHINOTHURIIDAE ECHINODERMATA Araeosoma cf. fenestratum (Thomson, 1872) BIBLIOGRAPHY Pawson, D.L. 1982. Papers from the Echinoderm Con- ference. 8. Deep-sea echi- noderms in the Tongue of the Ocean, Bahama Islands: a survey, using the research submersible Alvin. Australian Museum Memoir 16: 129–145. Schultz, H. 2011. Sea Urchins III - Worldwide Regular De- ep-Water Species. Heinke & Peter Schultz Partner. Thomson, C.W. 1872. Noti- ce of a new family of the Echinodermata. Proceedin- gs of the Royal Society of Edinburg 7 (1869-1872): 615- 617. DIAGNOSIS Test fragile, very flexible, low, circular, and hemispherical. Oral side almost flat, although slightly round towards the ambitus. Ventral primary spines ending in a hoof-like tip. Aboral spines large, numerous. Color ivory white (based on Thomson, 1872) (Figure 1). DISTRIBUTION Northeast Atlantic, from the Rockall Trough, to Portugal and in the West Atlantic from the West lndies to Brazil. Bathymetric range, from 148 to 1,180 m depth. Records in the present investigation include: Santos Basin at 1,850 m depth (S-M-1711) and from 2,116 to 2,188 m depth (Lapa) (Figure 2). COMMENTS Distinctive oral spines bearing hoof-like tips are one of the most characteristic features of echinothuriid sea urchins (Thomson, 1872). Their test usually collapses and most spines fall or break after sampling (Pawson, 1982). The identification was based on comparison with other in situ photographs, considering that no specimens were collected from the region for confirmation, and distinctive characteristics in the test and pedicellariae types, could not be observed. This species clearly differs from the upper bathyal East Atlantic congener Asterosoma violaceum Mortensen, 1903 by its color. Figure 1. The sea urchin Araeosoma cf. fenestratrum (Thomson, 1982) observed in Santos Basin at 2,145 m depth. Figure 2. Records of the sea urchin Araeosoma cf. fenestratrum (Thomson, 1982) in two of the investigated areas: S-M-1711 and Lapa, in Santos Basin. 22 | TotalEnergies Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 23 CRINOIDEA/ COMATULIDA/ ATELECRINIDAE ECHINODERMATA BIBLIOGRAPHY Campos, L.S., Moura, R.B., Veríssimo, I., Curbelo-Fernandez, M.P., Caval- canti, G.H., Brasil, A.C.S. 2010. ROV imaging of deep-sea echinoderms from the Brazilian continental mar- gin, Southwest Atlantic. In: Harris, L., Boetger, A., Walker, C.H. Echinoder- ms: Durham. Proceedings of the 12th International Echinoderm Conferen- ce. 147-152 pp. Gondim, A.I., Moura, R.B., Christof- fersen, M.L. Catalogue of living cri- noids (Echinodermata: Crinoidea) from Brazil. Zootaxa 4995 (1): 027– 055. Hess, H. & Messing, C.G. 2011. Arti- culata. Treatise on Invertebrate Pa- leontology, Part T, Echinodermata 2 Revised, Crinoidea, vol. 3. Seldon, P. (ed.), Ausich, W.I. (coord.). Universi- ty of Kansas Paleontological Institu- te, xxx + 261 p. Messing, C.G. 2013. A revision of the genus Atelecrinus PH Carpenter (Echinodermata: Crinoidea). Zoota- xa 3681 (1): 1–043. Messing, C.G. & Dearborn, J.H. 1990 Marine Flora and Fauna of the Nor- theastern United States. Echinoder- mata: Crinoidea. National Oceanic and Atmospheric Administration (NOAA) Technical Report, National Marine Fisheries Service 91, 30 pp. DIAGNOSIS Arms 10, long and delicate, extended almost parallel to the sediment. Arm tips slightly curved upwards, with longer brachials, but lacking pinnules. Arms whitish, with black pinnules (Figure 1) (based on Hess & Messing, 2011). DISTRIBUTION The family Atelecrinidae includes upper to middle slope species of feather stars (~200– 1500 m) with rare deeper records (down to >3,500 m) from the the Atlantic, Indian and tropical Pacific Oceans (Messing 2013). In the present investigation, it was recorded in Santos Basin from 1,150 to 1,501 m depth (S-M-1815) and from 1,481 to 1,492 m depth (S-M-1711) (Figure 2). COMMENTS In Brazil, the family Atelecrinidae is currently represented by Atelecrinus helgae A.H. Clark, 1913 (Gondim et al. 2021). Campos et al. (2010) recognized atelecrinid crinoids based on images from Campos Basin, but it was not possible to assert their identification at species level. They also occurred on undisturbed unconsolidated substrates. According to Messing (1990), dredged specimens of atelecrinids usually occur singly or in small numbers. Figure 1. An atelecrinid crinoid observed on the sediment in the Santos Basin at 1,614 m depth. Figure 2. Records of ophiuroids in two of the investigated areas: S-M-1815 and S-M-1711 in Santos Basin. Atelecrinidae Bather, 1899 Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 24 HOLOTHUROIDEA/ ELASIPODIDA/ PSYCHROPOTIDAE ECHINODERMATA Benthodytes cf. typica Théel, 1882 BIBLIOGRAPHY Billett, D.S.M., Hansen, B., Huggett, Q.J. 1985. Pelagic Holothurioidea (Echinodermata) of the northeast Atlantic. In: Keegan, B.F. & O’Connor, B.D.S. (Eds.) Echinodermata: Pro- ceedings of the Fifth International Echinoderms Conference, Galway, Ireland A.A. Balkema, Rotterdam, 399–411. Borrero-Pérez, G.H., Dueñas L.F., León, J., Puentes, V. 2020 Deep-sea holothurians (Echinodermata, Holo- thuroidea) from the Colombian Sou- thern Caribbean Sea. Check List 16 (3): 535–551. Hansen, B. 1975. Systematics and biology of the deep-sea holothu- rians. Galathea Report 13:1–262. Moura, R.B. 2016. Taxonomia e distri- buição de holoturias (Echinoderma- ta, Holothuroidea) do mar profundo brasileiro. Tese de Doutorado. Pro- grama de Pós-Graduação em Biolo- gia Animal, Universidade Federal de Pernambuco. Pawson, D.L. 1982. Papers from the Echinoderm Conference. 8. Deep- -sea echinoderms in the Tongue of the Ocean, Bahama Islands: a sur- vey, using the research submersible Alvin. Australian Museum Memoir 16: 129–145. Pawson, D.L., Nizinski, M.S., Ames, C.L., Pawson, D.L. 2015. Deep-sea echinoids and holothurians (Echino- dermata) near cold seeps and coral communities in the northern Gulf of Mexico. Bulletin of Marine Science, 91(2): 167–204. DIAGNOSIS Body flattened, ellipsoid, approximately 1.5x to 2x long as wide, surrounded by a very broad brim, wider laterally. Body wall rather thick, soft, and transparent. Mouth ventral, anus dorsal. Dorsal papillae absent. Color light violet to bluish, with lateral bands, radiating from the middle towards the circumference of the body (Figure 1) (after Théel, 1882). DISTRIBUTION Cosmopolitan species with depth range from 1,873 to 4,700 m. Indian Ocean, Pacific Ocean, and numerous records in the Atlantic Ocean, especially off the west coast of the United States (Hansen, 1975; Pawson et al., 2015; Moura, 2016; Borrero-Pérez et al., 2020). Records in the present investigation include: Santos Basin at 2,124 m depth (Lapa), and Campos Basin, from 2,860 to 2,998 m depth (C-M-541) (Figure 2). COMMENTS Benthodytes sanguinolenta Théel, 1882 and Benthodytes typica are the most common species of the genus, occurring from the lower slope to abyssal depths in the Atlantic. Although the latter is considered a well-known species in the Gulf of Mexico, Pawson et al. (2015) were unable to clearly distinguish it based on ROV images. Benthodytes sanguinolenta is usually larger than B. typica, but the main diagnostic characteristics used to identify these species are the number of tentacles, arrangement of oral papillae, and shape and size of their reduced ossicles, which cannot be observed macroscopically. Individuals observed in Bahamas were light brownish to cream, with little evidence of shades of red, while the anterior end was dark brown (Pawson, 1982). Images provided by Borrero-Pérez et al. (2020) off Colombia also showed color variation and documented the swimming behavior of an adult individual, which has been previously reported only for juveniles (Billett et al., 1985). Figure 1. The holothuroid Benthodytes cf. typica observed in Campos Basin at 2,997 m depth. Figure 2. Records of Benthodytes cf. typica in two of the investigated areas: Lapa, in Santos Basin, and C-M-541, in Campos Basin. Atlas of Benthic Fauna 24 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 25 ASTEROIDEA/ NOTOMYOTIDA/ BENTHOPECTINIDAE ECHINODERMATA Benthopecten Verrill, 1884 BIBLIOGRAPHY Clark, A.M. & Downey, M.E. 1992.Starfishes of the Atlan- tic. Chapman and Hall, Lon- don, 794 pp. Mah, C.L. 2023. World As- teroidea Database. Bentho- pecten Verrill, 1884. Acces- sed through: World Register of Marine Species at: ht- tps://www.marinespecies. org/aphia.php?p=taxdetail- s&id=123239 on 2023-10-15. DIAGNOSIS Body stellate. Disk small. Arms five, long, tapering distally. Interradial arcs rounded. Marginal plates large, covered each with spinules and at least a single spine. Odd interradial marginal plate bearing a conspicuously long vertical spine. Color red to orange on the disk fading to cream on middle section of each arm (based on Clark & Downey, 1992) (Figure 1). DISTRIBUTION Benthopecten species are found in deep-water, mostly from the middle to the lower slope, and four of them occur in the Atlantic: B. folini (Perrier, 1894), B. simplex (Perrier, 1881), B. spinosissimus (Sladen, 1889), and B. spinosus Verrill, 1884 (Mah, 2023). The record in the present investigation: Santos Basin at 1,849 m depth (S- M-1711) (Figure 2). COMMENTS Many of the details used for species identification, such as the form and occurrence of central disk spines, number of furrows, subambulacral and suboral spines, could not be observed in the provided image. Because of their fragility, even when sampled, many specimens of this group of sea stars have broken arms, making it difficult to observe basic characteristics useful for identification, such as the number of marginal plates (Clark & Downey, 1992). Figure 1. The sea star Benthopecten sp. observed in Santos Basin at 1,849 m depth. Figure 2. Record of the sea star Benthopecten sp. in one of the investigated areas: S-M-1711, in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021ECHINODERMATA ASTEROIDEA/ BRISINGIDA/ BRISINGIDAE Brisingidae G.O. Sars, 1875 Figure 1. A brisingid seastar observed in Santos Basin at 2,127 m depth. Note the smaller arm in regeneration. Figure 2. Records of brisingid seastars in one of the investigated areas: Lapa, in Santos Basin. DIAGNOSIS Disk small circular and well-defined. Madreporite large to moderate. Arms long, slender, usually 7-20, with distinct gonadal regions. Abactinal plates forming costae across the arms. Interradial arcs acute. Marginal spines also long and slender, with numerous pedicellariae. Color reddish to orange (Figure 1). DISTRIBUTION Brisingid sea stars occur worldwide, with some species observed in high abundance in deep-sea environments, from 100 and 4,000 m depth (Clark & Downey, 1992). The records in the present investigation include: Santos Basin, from 2,109 to 2,161 m depth (Lapa) (Figure 2). COMMENTS Brisingid sea stars are mostly found on hard substrates, such as sponges and corals, with arms extended in feeding posture (Clark & Downey, 1992; Campos et al., 2010). For this reason, they can be mistaken with comatulid crinoids. In Brazil, Novodinia antillensis and Brisinga sp. are the only known records of brisingids from the middle slope of Campos Basin (Campos et al., 2010). It was not possible to ascertain the identification of those specimens observed here to a lower level due to the lack of details in the images. BIBLIOGRAPHY Campos, L.S., Moura, R.B., Alcântara, P., Vasconcellos, R.F., Curbelo-Fernandez, M.P., Veríssimo, I., Cavalcan- ti, G.H. 2010. On two new records of the Family Brisin- gidae (Echinodermata: As- teroidea) from the Brazilian continental margin. In: Har- ris, L., Echinoderms: Durham. pp. 139-146. Clark, A.M. & Downey, M.E. 1992. Starfishes of the Atlan- tic. Chapman and Hall, Lon- don, 794 pp. Downey, M.E. 1986. Revision of the Atlantic Brisingida (Echinodermata: Asteroi- dea), with description of a new genus and family. Smi- thsonian Contributions to Zoology 435:43. Atlas of Benthic Fauna 26 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 27 HOLOTHUROIDEA/ SYNALLACTIDAE/ DEIMATIDAE ECHINODERMATA Deima validum validum Théel, 1879 DIAGNOSIS Body ovoid and symmetrical, flattened ventrally. Color in vivo varies from white to pinkish orange. Mouth and anus ventral. Dorsal and non-retractile papillae arranged in two rows along both radii but only visible nearest to the mediodorsal area. Tube feet modified into papillae arranged ventrally in two simple rows along the radii (Figure 1). DISTRIBUTION Cosmopolitan subspecies, occurring from 724 to 5,424 m depth, not recorded in the Arctic and Antarctica. Records in the Indian Ocean, from the Arabian Sea and Benguela Bay; West Pacific; East Atlantic, from Porcupine Seabight to Angola; West Atlantic, from Florida, in the United States down to Rio de Janeiro, in Brazil (Hansen, 1975; Bohn, 2006; Moura & Campos, 2010; Rogacheva et al., 2013; Borrero-Pérez et al., 2020). Records in the present investigation include: Santos Basin from 1,199 to 1,501 (S- M-1815), from 1,485 to 1,850 (S-M-1711) and from 2,124 to 2,178 (Lapa), and Campos Basin at 2,986 m depth (C-M- 541) (Figure 2). COMMENTS Deima validum validum is one of the most common holothuroids found on the middle slope of the Campos Basin (> 1,000 m), but occurs less abundantly on the lower slope (> 3,000 m). It can be easily recognized by the shape of the body and the presence of long dorsal papillae, which can be curved towards the posterior region. Figure 1. The holothuroid Deima validum validum Théel, 1879 observed in Santos Basin at 1,614 m depth. Figure 2. Records of Deima validum validum Théel, 1879 in the four investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin, and C-M-541, in Campos Basin. BIBLIOGRAPHY Hansen, B. 1975. Systematics and biology of the deep-sea holothurians. Galathea Re- port 13:1–262. Bohn, J.M. 2006. Crinoidea and Holothuroidea (Echi- nodermata) of the abyssal Angola Basin–Results of the DIVA-1 expedition of FS “Me- teor” (Cruise M48/1). Zoota- xa 1276: 1–31. Borrero-Pérez, G.H., Dueñas L.F., León, J., Puentes, V. 2020 Deep-sea holothurians (Echinodermata, Holothu- roidea) from the Colombian Southern Caribbean Sea. Check List 16 (3): 535–551. Moura, R.B. & Campos, L.S. 2010. Deima validum vali- dum. In: Lavrado, H.P. & Bra- sil, A.C.S. Biodiversidade da Região Oceânica Profunda da Bacia de Campos: Mega- fauna e Ictiofauna Demersal. Rogacheva A., Gebruk A., Alt C. 2013. Holothuroidea of the Charlie Gibbs Fracture Zone area, northern Mid-Atlantic Ridge. Marine Biology Rese- arch 9 (5–6): 587–623. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms28 | TotalEnergies ASTEROIDEA/ BRISINGIDA/ FREYELLIDAE ECHINODERMATA Freyellidae Downey, 1986 BIBLIOGRAPHY Clark, A.M. & Downey, M.E. 1992.Starfishes of the Atlan- tic. Chapman and Hall, Lon- don, 794 pp. Downey, M.E. 1986. Revision of the Atlantic Brisingida (Echinodermata: Asteroi- dea), with description of a new genus and family. Smi- thsonian Contributions to Zoology 435:43. DIAGNOSIS Disk small, circular, and merging with the arms. Madreporite small. Arms long, slender, usually 6-17, more attached to the disc. Abactinal plates forming a mosaic on proximal part of the arms. Interradial arcs gently curved, interradial plates bare. Marginal spines also long and slender, with numerous pedicellariae. Color whitish (Figure 1). DISTRIBUTION Freyellidae are also deep-sea dwellers, but found at a broader range than brisingids, at depths from 950 to 5,600 m (Downey, 1986). The records in the present investigation include: Santos Basin, at 1,848 m depth (S- M-1711) and from 2,127 to 2,178 m depth (Lapa) (Figure 2). COMMENTS Differently from brisingids, freyellid sea stars can be considered detritivores and are found on soft substrate (Clark & Downey, 1992). Because of the arrangement of their proximal arm plates, they are unable rise their arms vertically over the disk (Downey et al., 1986). Some species, however, are able to raise the distal portions of their arms (Clark & Downey, 1992). Due to the lack of detailed images, the identification of the specimens observed here is tentative. Figure 1. A possible freyellid seastar observed in Santos Basin at 2,132 m depth. Figure 2. Records of freyellid seastars in two of the investigated areas: S-M-1711 and Lapa, in Santos Basin. Atlas of Benthic Fauna 28 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 29 HOLASTEROIDA/ CERATOPHYSIDAE ECHINOIDEA Echinosigra phiale Thomson, 1873 BIBLIOGRAPHY Minin, K.V., Mironov, A.N., Pe- trov, N.B., Vladychenskaya, I.P. 2024. Evolutionary and biogeographic patterns in the deep-sea echinoid fa- milies Pourtalesiidae Agas- siz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoo- logical Journal of the Linne- an Society, 202 (4): 1–30. Mironov, A.N. 2008. Pourta- lesiid sea urchins (Echino- dermata: Echinoidea) of the northern Mid-Atlantic Ridge. Marine Biology Research, 4(1-2): 3–24. Mironov, A.N. 2014. Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates li- ving deeper than 2000 m in the seas bordering Europe. Echinoidea. Invertebrate Zo- ology 11: 120–129. Mortensen T. 1907. Echinoi- dea. Danish Ingolf-Expedi- tion, 4(2):1-200. Schultz, H. 2009. Sea Ur- chins II - worldwide irregular deep water species. Heinke & Peter Schultz Partner Scientific Publications: Hem- dingen. 501-849 pp. DIAGNOSIS Test thin, bottle-shaped (TW/TL<0.30), with an anterior funnel-like deep peristomial groove, a slender neck (AW/ AL 0.21–0.33), and a dilated body with ventral and dorsal keels, that ends in a rostrate tail-like structure (based on Mortensen, 1907 and Mironov, 2008) (Figure 1). DISTRIBUTION Echinosigra phiale occurs from the northern and north-eastern Atlantic, from 1,535 to 3,800 m (Mironov, 2014). The record in the present investigation: Santos Basin at 1,642 m depth (S-M-1711) (Figure 2). COMMENTS Based on phylogenetic reconstructions, Minin et al. (2024) showed that the former subgenus Echinosigra (Echinosigra) and Echinosigra (Echinogutta) belonged to distant lineages and raised the latter to a genus rank. Underwater photographs of these thin-shelled deep- sea echinoids are uncommon in literature, as they are considered most-specialized burrowers, submerging through the silty sediment (Minin et al. 2024). Detailed morphological information on the apical system and pedicellariae could not be obtained, as the observed specimen was not collected. Figure 1. The bottle-shaped sea urchin Echinosigra phiale (Thomson, 1873) observed in Santos Basin at 1,642 m depth. Figure 2. Record of the sea urchin Echinosigra phiale (Thomson, 1873) in one of the investigated areas: S-M-1711, in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms30 | TotalEnergies ECHINODERMATA BIBLIOGRAPHY Grassle, J.F., Sanders, H.L., Hessler, R. R., Rowe G.T., McLellan, T. 1975. Pattern and zonation: a study of the bathyal megafauna using the research submersible Al- vin. Deep-Sea Research 22: 457-481. Mironov, A.N. 2006. Echi- noids from seamounts of the north-eastern Atlantic; onshore/offshore gradients in species distribution. In: Mironov, A.N., Gebruk, A.V., Southward, A.J. Biogeogra- phy of the North Atlantic se- amounts. 96–133 pp. Pawson, D.L. 1982. Papers from the Echinoderm Con- ference. 8. Deep-sea echi- noderms in the Tongue of the Ocean, Bahama Islands: a survey, using the research submersible Alvin. Australian Museum Memoir 16: 129–145. Yepes-Gaurisas, D. & Guer- rero-Correa, G. 2022. First record and range extension of the deep-sea urchin Hy- grosoma petersii (A. Agas- siz, 1880) (Echinodermata, Echinoidea) for the southern Caribbean. Bulletin of Marine and Coastal Research 51 (1): 207–214. DIAGNOSIS Test large, very flexible, low, pentagonal, and hemispherical. Oral side almost flat, although slightly round towards the ambitus. Aboral spines long, thin, and numerous. Ventral primary spines curved, bearing a white hoof-like structure at the tip. Color varying from dark purple to blackish (Figure 1) (after Yepes-Gaurisas & Guerrero-Correa, 2022). DISTRIBUTION Hygrosoma petersii is the only of the three closely-related species in the genus that occurs in the Atlantic Ocean, with most records in the North Atlantic, from 200 to 3,200 m (Mironov, 2006; Yepes- Gaurisas & Guerrero-Correa, 2022). In the South Atlantic is also found off South Africa. Records in the present investigation include: Santos Basin from 1,849 to 1,853 m depth (S-M-1711) and from 2,110 to 2,189 m depth (Lapa) (Figure 2). COMMENTS The identification proposed here is provisional, considering that no specimens were collected from the region for confirmation. Some individuals displayed the same behavior observed for Hygrosoma petersii in the Bahamas islands, walking quickly on ventral spines (Grassle et al., 1975, Pawson, 1982). Despite the presence of foraminifera and pteropod shells in the intestinal contents of some specimens, the diet of this species in the Bahamas consisted almost exclusively of Sargassum and Thalassia fragments (Pawson, 1982). Figure 1. The sea urchin Hygrosoma cf. petersii (A. Agassiz, 1880) observed in Santos Basin at 2,129 m depth. Figure 2. Records of the sea urchin Hygrosoma cf. petersii (A. Agassiz, 1880) in two of the investigated areas: S-M-1711 and Lapa, in Santos Basin. ECHINOIDEA/ ECHINOTHURIOIDA/ ECHINOTHURIIDAE Hygrosoma cf. petersii (A. Agassiz, 1880) Atlas of Benthic Fauna 30 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 31 ECHINODERMATA BIBLIOGRAPHY Campos, L.S., Vasconcelos, R.F., Mou- ra, R.B., Frensel, R. 2010a. Nymphaster arenatus (Perrier, 1881). In: Lavrado, H.P. & Brasil, A.C.S. Biodiversidade da Região Oceânica Profunda da Bacia de Campos: Megafauna e Ictiofauna Demersal. 268-169 pp. Campos, L.S., Moura, R.B., Veríssimo, I., Curbelo-Fernandez, M.P., Cavalcan- ti, G.H., Brasil, A.C.S. 2010b. ROV ima- ging of deep-sea echinoderms from the Brazilian continental margin, Sou- thwest Atlantic. In: Harris, L., Boetger, A., Walker, C.H. Echinoderms: Durham. Proceedings of the 12th International Echinoderm Conference. 147-152 pp. Costa, P.A.S., Mincarone, M.A., Costa Braga, A., Martins, A.S., Lavrado, H.P., Haimovici, M. & Falcao, A.P. 2015. Me- gafaunal communities along a depth gradient on the tropical Brazilian con- tinental margin. Marine Biology Rese- arch, 11 (10): 1053–1064. Mah, C. 2020. New species, occurren- ce records and observations of preda- tion by deep-sea Asteroidea (Echino- dermata) from the North Atlantic by NOAA ship Okeanos Explorer. Zoota- xa 4766 (2): 201–260. Wagstaff, M.C., Howell, K.L., Bett, B.J., Billett, D.S.M., Brault, S., Stuart, C.T. & Rex, M.A. 2014. diversity of deep-sea holothurians and asteroids along a bathymetric gradient (NE Atlantic). Marine Ecology Progress Series 508, 177–185. DIAGNOSIS Body flat. Disk large. Arms elongated, wide basally, narrowing to the tip. Interradial margins arcuate. Abactinal plates completely covered with granules. Superomarginals squarish, in contact across the midline of the arms (Figure 1). DISTRIBUTION In the Northwest Atlantic, from the Georgia, Gulf of Mexico to southeastern Brazil, and in the Northeast Atlantic, from Ireland to Morocco, Canary and Cape Verde Islands, from 225-3,000 m depth (Clark & Downey, 1992; Campos et al. 2010 a, b). Records in the present investigation include: Santos Basin from 1,152 to 1,498 m depth (S-M-1815) and from 1,489 to 1,852 m depth (S- M-1711) (Figure 2). COMMENTS According to Costa et al. (2015) and Wagstaff et al. (2014), Nymphaster arenatus is considered a scavenger and a predator, as illustrated in-situ observation by Mah (2020). Figure 1. The seastar Nymphaster arenatus (Perrier, 1881) observed in the Santos Basin at 1,847 m depth. Figure 2. Records of the seastar Nymphaster arenatus (Perrier, 1881) in two of the investigated areas: S-M-1815 and S-M-1711 in Santos Basin. ASTEROIDEA/ VALVATIDA/ GONIASTERIDAE Nymphaster arenatus (Perrier, 1881) Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms32 | TotalEnergies ECHINODERMATA BIBLIOGRAPHY Madsen, F.J. 1973. The Ophiodermatidae. Galathea Report 11: 133–143. Pawson, D.L. 1982. Papers from the Echinoderm Con- ference. 8. Deep-sea echi- noderms in the Tongue of the Ocean, Bahama Islands: a survey, using the research submersible Alvin. Australian Museum Memoir 16: 129–145. DIAGNOSIS Disk flat, with radial elevations. Scales or plates not visible, probably thin and very small. Radial shields ellipsoid, located near the margin of the disk, separated from each other. Arms unbranched, stout, tapering, with tips slightly raised from the sediment. Arm length around four times the disk diameter. Lateral arm spines very short. Color in vivo orange with arms distally fading to white (Figure 1). DISTRIBUTION The records in the present investigation: Santos Basin from 1,488 to 1,850 m depth (S-M-1711) and from 2,116 to 2,188 m depth (Lapa) (Figure 2). COMMENTS Although many deep-sea ophiuroids are part of the macrofauna, larger specimens also occur buried in the sediment or associated with other benthic organisms, such as corals and sponges. Among few species identified from images in the Atlantic, Bathypectinura heros (Lyman, 1879) is a highly active uniformly orange ophiuroid commonly found in the main Atlantic, Indian Ocean and Western Pacific from 240 to 2,960 m depth. (Madsen, 1973; Pawson, 1982). However, most morphological details of ophiuroids are not usually seen in in situ photographs, which results in a higher identification level. Figure 1. An ophiuroid observed in Santos Basin at 2,143 m depth. Figure 2. Records of ophiuroids in two of the investigated areas: S-M-1711 and Lapa, in Santos Basin. OPHIUROIDEA/ OPHIURIDA Ophiurida Müller & Troschel, 1840 sensu O’Hara et al., 2017 Atlas of Benthic Fauna 32 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 33 ECHINODERMATA BIBLIOGRAPHY Borrero-Pérez, G.H., Dueñas L.F., León, J., Puentes, V. 2020 Deep-sea holothurians (Echinodermata, Holothu- roidea) from the Colombian Southern Caribbean Sea. Check List 16 (3): 535–551. Gebruk, A.V., Smirnov, A.V., Rogacheva, A.V. 2014 Deep- -sea fauna of European seas: An annotated species che- ck-list of benthic invertebra- tes living deeper than 2000 m in the seas bordering Eu- rope. Holothuroidea. Inverte- brate Zoology 11(1): 156–180. Rogacheva, A., Gebruk, A., Alt, C.H.S. 2012. Swimming deep-sea holothurians (Echi- nodermata: Holothuroidea) on the northern Mid-Atlan- tic Ridge. Zoosymposia 7: 213–224. Rogacheva, A., Gebruk, A., Alt, C.H.S. 2013. Holothu- roidea of the Charlie Gibbs Fracture Zone area, northern Mid-Atlantic Ridge. Marine Biology Research 9 (5–6): 587–623. DIAGNOSIS Body ovoid. Dorsal papillae anteriorly placed, forming a short velum. Mouth in a ventral lobe, surrounded by ten tentacles. Ventral tube feet restricted to posterior lobes. Guts visible through the body wall. Colour translucent light pink, with reddish tentacle disks (Figure 1) (based on Rogacheva et al. 2012; Borrero-Pérez et al., 2020). DISTRIBUTION The genus Peniagone includes around 30 lower-slope to abyssal- depth species widely distributed in the world oceans, except for the Arctic (Gebruk et al., 2014). In the present investigation this species occurred in Santos Basin, from 1,488 to 1,849 (S-M-1711) and from 2,109 to 2,177 m depth (Lapa) and in Campos Basin at 3,000 m depth (C-M-541) (Figure 2). COMMENTS Species of Peniagone, Elpidiidae, Amperima, and Ellipinion as considered frequent swimmers, but also spend some time feeding on the seafloor (Rogacheva et al. 2012). After defecating, these sea cucumbers use their tentacles and body undulations as an effective method for swimming. Peniagone includes most species in the family, also showing the highest diversity of external morphological features (Rogacheva et al., 2013). HOLOTHUROIDEA/ ELASIPODIDA Peniagone Théel, 1882 Figure 1. A swimming individual of Peniagone sp. observed in C-M-541, in Campos Basin, at at 3,000 m depth. Figure 2. Records of Peniagone sp. in three of the investigated areas: S-M-1711 and Lapa, in Santos Basin, and C-M-541 in Campos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms34 | TotalEnergies ECHINODERMATA BIBLIOGRAPHY O’Loughlin, P.M., Ahearn, C. 2005. A review of pygal-fur- rowed Synallactidae (Echi- nodermata: Holothuroidea), with new species from the Antarctic, Atlantic and Paci- fic oceans. Mem Mus Victo- ria, 62(2):147–179. Miller, A.K., Kerr, A.M., Paulay, G., Reich, M., Wilson, N.G., Carvajal, J.I. & Rouse, G.W. 2017. Molecular phylogeny of extant Holothuroidea (Echi- nodermata). Molecular Phy- logenetics and Evolution. 111: 110-131. Pawson, D.L., Nizinski, M.S., Ames, C.L., Pawson, D.L. 2015. Deep-sea echinoids and holothurians (Echino- dermata) near cold seeps and coral communities in the northern Gulf of Mexico. Bul- letin of Marine Science, 91(2): 167–204. DIAGNOSIS Body more or less cylindrical to fusiform, mostly or totally covered with dead foraminifera and/or pteropod shells. Color dirty cream to gray (after Pawson et al., 2015) (Figure 1). DISTRIBUTION The order Persiculida consists of deep-sea holothuroid genera spread worldwide, some still requiring taxonomic revision. Records in the present investigation include: Santos Basin, from 2,109 to 2,187 m depth (Lapa) (Figure 2). COMMENTS Some of the genera currently included in the order Persiculida – Oloughlinius, Pseudostichopus (Pseudostichopodidae) and Molpadiodemas (Molpadiodemidae) were previously classified into the family Synallactidae (O’Loughlin & Ahearn, 2005. Miller et al., 2017). These are very difficult to determine from images alone. Even when collected specimens are available, body wall ossicles, which are usually very informative for holothuroid taxonomy, are absent in many species, encrusting their body walls with sediment attachments. For instance, specimens of Molpadiodemas involutus are usually covered with dead foraminifera Globigerina spp., whereas specimens of Oloughlinius, like many specimens observed here, are found entirely covered with pteropod mollusks of the genus Clio spp. (O’Loughlin & Ahearn, 2002; Pawson et al., 2015). Figure 1. A persiculid sea cucumber covered by pteropod shells observed in Santos Basin at 2,131 m depth. HOLOTHUROIDEA/ PERSICULIDA Persiculida Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 Figure 2. Records of the persiculid sea cucumbers in one of the investigated areas: Lapa, in Santos Basin. Atlas of Benthic Fauna 34 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 35 ECHINOIDEA/ ECHINOTHURIOIDA/ ECHINOTHURIIDAE ECHINODERMATA Phormosoma placenta Thomson, 1872 BIBLIOGRAPHY Emson, R.H. & Young, C.M. (1998) Form and function of the primary spines of two bathyal echinothuriid sea urchins. Acta Zoologica, Stockholm, 79 (2): 101 - 111. Gondim, A.I., Moura, R.B., Christoffer- sen, M.L. & Dias, T.L.P. 2018. Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from nor- theastern Brazil. Zootaxa 4529 (1): 1-72. Mironov, A.N. 2014. Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates li- ving deeper than 2000 m in the seas bordering Europe. Echinoidea. Inverte- brate Zoology, 11 (1): 120 - 129. Pawson, D.L. 1982. Papers from the Echinoderm Conference. 8. Deep-sea echinoderms in the Tongue of the Oce- an, Bahama Islands: a survey, using the research submersible Alvin. Australian Museum Memoir 16: 129–145. Pawson, D.L. Pawson, D.L., Nizinski, M.S., Ames, C.L., Pawson, D.L. 2015. Deep-sea echinoids and holothurians (Echinodermata) near cold seeps and coral communities in the northern Gulf of Mexico. Bulletin of Marine Science, 91(2): 167–204. Serafy, D.K. & Fell, F.J. 1985. Marine flo- ra and fauna of the Northeastern Uni- ted States. Echinodermata: Echinoidea. NOAA Technical Report National Mari- ne Fisheries Service, 33, 1 - 27. DIAGNOSIS Test low, fragile, hemispherical, with sharp round margin, limited by pointed ventral spines without the hoof-like structures found in other echinothurioids. Aboral spines long, many enclosed by fleshy sac. Color varying from red to brownish in live specimens (based on Pawson et al., 2015) (Figure 1). DISTRIBUTION Phormosoma placenta is the only of four species in the genus that occurs in the Atlantic Ocean. In the West Atlantic, from the North Carolina, United States, to the Southeastern Brazil, from 38 down to 4100 m (Mironov, 2014; Gondim et al. 2018), but it is very rare at depths shallower than 500 m (Serafy & Fell, 1985). Records in the present investigation: Santos Basin from 1,646 to 1,851 m depth (S-M-1711) (Figure 2). COMMENTS One of the most conspicuous characters that can be observed in live specimens of Phormosoma is their fleshy spines, which may partially or totally obscure the dorsal surface of the test. The main function of these spines is most probably defense (Emson & Young 1998). Figure 1. The sea urchin Phormosoma placenta Thomson, 1872 observed in Santos Basin at 1,849 m depth. Figure 2. Records of the sea urchin Phormosoma placenta Thomson, 1872 in one of the investigated areas in Santos Basin (S-M-1711). Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms36 | TotalEnergies ASTEROIDEA/ VALVATIDA/ GONIASTERIDAE ECHINODERMATA Plinthaster dentatus (Perrier, 1884) BIBLIOGRAPHY Clark, A.M. & Downey, M.E. 1992. Starfishes of the Atlan- tic. Chapman and Hall, Lon- don, 794 pp. Cunha, R., Martins, L.R., Me- negola, C., Souto, C. 2021. Taxonomy of the sea stars (Echinodermata: Asteroi- dea) from Bahia State, inclu- ding ontogenetic variation and an illustrated key to the Brazilian species. Zootaxa 4955 (1): 001–078. Halpern, J.A. 1970. A mo- nographic revision of the goniasterid sea stars of the North Atlantic. Unpublished PhD Dissertation, University of Miami, Florida, 253 pp. Mah, C. 2020. New species, occurrence records and ob- servations of predation by deep-sea Asteroidea (Echi- nodermata) from the North Atlantic by NOAA ship Oke- anos Explorer. Zootaxa 4766 (2): 201–260. DIAGNOSIS Body pentagonal to stellate. Disk wide with a large madreporite. Arms short. Abactinal plates flat, block-like. Terminal arm plate large, tapering distally. Color cream. (based on Clark & Downey, 1992) (Figure 1). DISTRIBUTION Plinthaster dentatus occurs widely in the Atlantic (Clark & Downey, 1992; Ventura & Nobre, 2007; Mah, 2020; Cunha et al., 2021). In the Western Atlantic, from the North Carolina, United States to the Southern Brazil. In the Northeast Atlantic, in the Rockall Trough, Newfoundland to Gulf of Guinea, Azores, Canary and Cape Verde Islands, and Faroe Islands. According to Mah (2020), records from the South Pacific should be confirmed. From 229 to 2,910 m depth. Records in the present investigation include: Santos Basin, 1,484 and 1,850 m depth (S-M-1711) (Figure 2). COMMENTS According to Mah (2020), little data are available on the biology of Plinthaster dentatus in contrast to the number of specimens available in scientific collections. Based on ROV images from the North Atlantic the author identified this species probably feeding on encrusting organisms on Madrepora rubble and geodiid sponges. Foraminifera, crinoid arms, corals fragments were also found among the feeding items for this species (Halpern, 1970). Figure 1. The seastar Plinthaster dentatus (Perrier, 1884) observed in the Santos Basin at 1,850 m depth. Figure 2. Records of the seastar Plinthaster dentatus (Perrier, 1884) in one of the investigated areas: S-M-1711, in Santos Basin. Atlas of Benthic Fauna 36 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 37 HOLOTHUROIDEA/ ELASIPODIDA/ PSYCHROPOTIDAE ECHINODERMATA Psychropotes depressa (Théel, 1882) BIBLIOGRAPHY Borrero-Pérez, G.H., Dueñas L.F., León, J., Puentes, V. 2020 Deep-sea holothurians (Echinodermata, Holothu- roidea) from the Colombian Southern Caribbean Sea. Check List 16 (3): 535–551. Moura, R.B. 2016. Taxonomia e distribuição de holoturias (Echinodermata, Holothu- roidea) do mar profundo brasileiro. Tese de Doutora- do. Programa de Pós-Gra- duação em Biologia Animal, Universidade Federal de Pernambuco. Pawson, D.L., Nizinski, M.S., Ames, C.L., Pawson, D.L. 2015. Deep-sea echinoids and holothurians (Echino- dermata) near cold seeps and coral communities in the northern Gulf of Mexico. Bul- letin of Marine Science, 91(2): 167–204. Rogacheva, A., Gebruk, A., Alt, C.H.S. 2012. Swimming deep-sea holothurians (Echi- nodermata: Holothuroidea) on the northern Mid-Atlan- tic Ridge. Zoosymposia 7: 213–224. Rogacheva, A., Gebruk, A., Alt, C.H.S. 2013. Holothu- roidea of the Charlie Gibbs Fracture Zone area, northern Mid-Atlantic Ridge. Marine Biology Research 9 (5–6): 587–623. DIAGNOSIS Body arched dorsally, flattened ventrally, approximately twice as long as wide, surrounded by a ventrolateral brim. Mouth and anus ventral. Body wall translucent, light violet, darker ventrally. Dorsal longitudinal muscles and intestine visible through the body wall. Prominent dorsal projection approximately one-sixth of the body length located about one-third of the body length from the posterior end. Three or more pairs of short and blunt projections equally spaced along the inner edges of the dorsal radii, the posterior-most pair largest, placed at mid-body (based on Pawson et al., 2015 and Borrero-Pérez et al. 2020) (Figure 1). DISTRIBUTION Cosmopolitan species with depth range from 957 to 4,200 m. In the Western Atlantic, this species has been recorded in the Gulf of Mexico and off Cuba, Colombia, and Brazil (Rogacheva et al., 2013; Pawson et al. 2015; Moura, 2016; Borrero-Perez et al., 2020). Records in the present investigation include: Santos Basin from 1,463 to 1,498 m (S-M-1815), from 1,484 to 1,652 m (S-M-1711) and from 2,113 to 2,187 m (Lapa), and Campos Basin from 2,861 to 2,999 m depth (C-M-541) (Figure 2). COMMENTS Smaller specimens with shorter papillae were observed here, shallower in the middle slope. Although commonly observed on the seafloor, P. depressa can occasionally swim, usually in response to disturbance or when changing its location (Rogacheva et al., 2012). Figure 1. The holothuroid Psychropotes depressa (Théel, 1882) observed in Campos Basin at 2,863 m depth. Figure 2. Records of Psychropotes depressa (Théel, 1882) in the four investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin, and C-M-541, in Campos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms38 | TotalEnergies HOLOTHUROIDEA/ ELASIPODIDA/ PSYCHROPOTIDAE ECHINODERMATA Psychropotes semperiana Théel, 1882 BIBLIOGRAPHY Bohn, J.M. 2006. Crinoidea and Holothuroidea (Echi- nodermata) of the abyssal Angola Basin - Results of the DIVA-1 expedition of FS ME- TEOR (Cruise M48/1). Zoo- taxa 1276: 1-31. Gebruk, A.V., Smirnov, A.V., Rogacheva, A.V. 2014 Deep- -sea fauna of European seas: An annotated species che- ck-list of benthic invertebra- tes living deeper than 2000 m in the seas bordering Eu- rope. Holothuroidea. Inverte- brate Zoology 11(1): 156–180. Hansen, B. 1975. Systematics and biology of the deep-Sea holothurians. Part. 1. Elasipo- da. Scientific Results of the Danish Deep-Sea Expedition Round the World 1950–52. Galathea Report. 13: 1–262. Moura, R.B. 2016. Taxonomia e distribuição de holoturias (Echinodermata, Holothu- roidea) do mar profundo brasileiro. Tese de Doutora- do. Programa de Pós-Gra- duação em Biologia Animal, Universidade Federal de Pernambuco. Théel, H. 1882. Report on the Holothurioidea dredged by HMS Challenger during the years 1873-1876. Part I. Scientific results of H.M.S. Challenger 1873-1876. Zoolo- gy IV 13: 1-176. DIAGNOSIS Body flattened anteriorly, but more elevated posteriorly, where an unpaired dorsal appendage, measuring around 1/3 of the body length, is placed. Body wall dark purple. Ventral surface flattened limited by a narrow brim of fused tube feet around the body. Mouth and anus ventral. (Figure 1). DISTRIBUTION North and West Indian Ocean; Most records along the Atlantic Ocean, especially in the North Atlantic, from 1,433 to 5,610 m (Théel, 1882; Hansen, 1975; Bohn, 2006; Gebruk et al. 2014; Moura, 2016). The record in the present investigation: Campos Basin at 2,984 m depth (C-M-541) (Figure 2). COMMENTS The dorsal appendage is not frequently preserved in sampled specimens (Hansen, 1975). Figure 1. The holothuroid Psychropotes semperiana Théel, 1882 observed in Campos Basin at 2,984 m depth. Figure 2. Record of the holothuroid Psychropotes semperiana Théel, 1882 in one of the investigated areas in Campos Basin (C-M-541). Atlas of Benthic Fauna 38 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 39 ACTINOPTERYGII/ NOTACANTHIFORMES/ HALOSAURIDAE ACTINOPTERYGII Aldrovandia affinis (Günther, 1877) BIBLIOGRAPHY Cohen, D.M., A.W. Ebeling, T. Iwamoto, S.B. McDowell, N.B. Marshall, D.E. Rosen, P. Sonoda, W.H. Weed III and L.P. Woods, 1973. Fishes of the western North Atlantic. Part six. New Haven, Sears Found. Mar. Res., Yale Univ. Costa, P.A.S. 2010. In: Lavra- do, H.P. & Brasil, A.C.S. Bio- diversidade da Região Oce- ânica Profunda da Bacia de Campos: Megafauna e Ictio- fauna Demersal. 306 – 307 p. Sulak, K.J., 1986. Halosauri- dae. p. 196-197. In M.M. Smi- th and P.C. Heemstra (eds.) Smiths’ sea fishes. Springer- -Verlag, Berlin. DIAGNOSIS Body rather elongate, variable coloration: white to grey- brown, underside darker. Insertion of the pelvic fin only slightly anterior to the origin of the dorsal fin. Lacks scale on the opercle and head (Costa, 2010). Adult reaches 60 cm in length (Figure 1). DISTRIBUTION The species is circumglobal (except in polar and subpolar regions as well as the Mediterranean). Western Indian Ocean; Eastern Indian Ocean; Western Central Pacific Ocean; Eastern Central Pacific Ocean; Southwest Pacific Ocean; Southeast Pacific Ocean; Northwestern Atlantic Ocean; Eastern Central Atlantic Ocean; Western Central Atlantic Ocean; Southeast Atlantic Ocean; Southwest Atlantic Ocean; In Brazil found between 1,067 and 1,632 m depth (Costa, 2010). The records in the present investigation include: Santos Basin, from 1,148 to 1,500 m depth (S-M-1815), from 1,492 to 1,848 m depth (S-M-1711) and from 2,117 to 2,160 m depth (Lapa) (Figure 2). COMMENTS Feeds on benthic invertebrates: polychaetes, bivalves, amphipods and ophiuroids; large amounts of sediment can be found in your digestive tract. Figure 1. Gilbert’s halosaurid fish, Aldrovandia affiis (Günther, 1877) observed in Santos Basin at 2,160 m depth. Figure 2. Records of Aldrovandia affiis (Günther, 1877) in three of the investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms40 | TotalEnergies ZEIFORMES/ OREOSOMATIDAE ACTINOPTERYGII Allocyttus verrucosus (Gilchrist, 1906) BIBLIOGRAPHY Costa, P.A.S. 2010. In: Lavra- do, H.P. & Brasil, A.C.S. Bio- diversidade da Região Oce- ânica Profunda da Bacia de Campos: Megafauna e Ictio- fauna Demersal. 362 – 363 p. Kailola, P.J., M.J. Williams, P.C. Stewart, R.E. Reichelt, A. McNee and C. Grieve, 1993. Australian fisheries re- sources. Bureau of Resource Sciences, Canberra, Austra- lia. 422 p. Karrer, C., 1986. Oreosomati- dae. p. 438-440. In M.M. Smi- th and P.C. Heemstra (eds.) Smiths’ sea fishes. Springer- -Verlag, Berlin. Lyle, J.M. and D.C. Smith, 1997. Abundance and biolo- gy of warty oreo (Allocyttus verrucosus) and spiky oreo (Neocyttus rhomboidalis) (Oreosomatidae) off sou- th-eastern Australia. Mar. Freshwat. Res. 48:91-102. DIAGNOSIS Body diamond-shaped with two rows of large, warty scales flanking both sides between the pelvic and anal fins. Coloration variable: adults exhibit a blackish hue, while prejuveniles appear silvery with a greenish back and prominent grey blotches that vanish at approximately 12 cm (Karrer, 1986). Possesses notably large eyes. Adult length about 30 cm (Figure 1). DISTRIBUTION Eastern Atlantic: off Mauritania, Gulf of Guinea, Namibia and South Africa. Western Atlantic: off Suriname, southern Brazil and Uruguay. Western Indian Ocean: Mozambique Channel, Walters Bank. Eastern Indian Ocean: East Indian and West Australian Ridge, Australia. South Pacific: New Zealand and off Chile. In Brazil found between 1,059 and 1,324 m depth (Costa, 2010). The records in the present investigation include: Santos Basin, at 1,150 and 1,497 m depth (S-M-1815) and at 1,847 depth (S-M-1711) (Figure 2). COMMENTS Displays seasonally synchronized reproduction. After spawning, the eggs float close to the surface (Kailola et.al., 1993). Juveniles are pelagic and live at depths below 1000 m (Lyle & Smith, 1997). Feeds on invertebrates, and fishes. Figure 1. Warty oreodory, Allocyttus verrucosus (Gilchrist, 1906) observed in Santos Basin at 1,847 m depth. Figure 2. Records of Allocyttus verrucosus (Gilchrist, 1906) in two of the investigated areas: S-M-1815 and S-M-1711, in Santos Basin. Atlas of Benthic Fauna 40 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 41 GADIFORMES/ MORIDAE ACTINOPTERYGII Antimora rostrata (Günther, 1878) DIAGNOSIS Body rather elongated, compressed; variable coloration: greyish-blue, olive green, deep purple, or black. Pointed snout with bony grooves widening beneath the eyes. Extremely long dorsal fin ray, divided dorsal and anal fins. Adult length: 40-75 cm (Figure 1). DISTRIBUTION The species is circumglobal in temperate and subantarctic latitudes (except in the North Pacific), archibenthic to abyssobenthic in habit. Western Indian Ocean; Eastern Indian Ocean; Antarctic Indian Ocean; Eastern Central Pacific Ocean; Southwest Pacific Ocean; Southeast Pacific Ocean; Eastern Central Atlantic Ocean; Western Central Atlantic Ocean; Northwestern Atlantic Ocean; Southwestern Atlantic Ocean; Antarctic Atlantic Ocean. In Brazil found between 1,059 and 1,324 m depth (Costa, 2010). The records in the present investigation include: Santos Basin from 1,148 to 1,459 m depth (S-M-1815), at 1,613 m depth (S-M- 1711) and from 2,133 to 2135 m depth (Lapa) (Figure 2). COMMENTS Feeds on benthic invertebrates. Figure 1. Blue antimoral, Antimora rostrata (Günther, 1878) observed in Santos Basin at 1,458 m depth. Figure 2. Records of Antimora rostrata (Günther, 1878) in three of the investigated areas: S-M-1815, S-M_1711 and Lapa, in Santos Basin. BIBLIOGRAPHY Cohen, D.M., T. Inada, T. Iwa- moto and N. Scialabba, 1990. FAO species catalogue. Vol. 10. Gadiform fishes of the world (Order Gadiformes). An annotated and illustrated catalogue of cods, hakes, grenadiers and other gadi- form fishes known to date. FAO Fish. Synop. 125(10). Rome: FAO. 442 p. Costa, P.A.S. 2010. In: Lavra- do, H.P. & Brasil, A.C.S. Bio- diversidade da Região Oce- ânica Profunda da Bacia de Campos: Megafauna e Ictio- fauna Demersal. 342 – 343 p. LOPES, P. R. D. et al. Ob- servações sobre Antimo- ra rostrata (Günther, 1878) (Actinopterygii: Moridae) no Estado do Rio de Janei- ro, Brasil (Oceano Atlântico Ocidental). Arquivos de Ci- ências do Mar. Fortaleza, v. 44, n. 1, p. 107-110, 2011. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms42 | TotalEnergies ACTINOPTERYGII Bathypterois grallator (Goode & Bean, 1886) BIBLIOGRAPHY Franco, M.A.L. 2010. In: La- vrado, H.P. & Brasil, A.C.S. Biodiversidade da Região Oceânica Profunda da Bacia de Campos: Megafauna e Ic- tiofauna Demersal. Sulak, K. 1977. The syste- matics and biology of Ba- thypterois (Pisces, Chlo- rophthalmidae) with a revised classification of ben- thic myctophiform fishes. Rosenstiel School of Marine and Atmospheric Science. University of Miami. DIAGNOSIS Pectoral fin not divided into two parts and without greatly produced rays, none extending beyond origin of anal fin. Outer ventral and lower caudal rays longer than standard length. Adipose fin absent (Figure 1). DISTRIBUTION The genus is circumglobal beneath temperate and tropical seas, archibenthic to abyssobenthic in habit. Western Indian Ocean; Western Pacific Ocean; Northeast Australia; Eastern Atlantic Ocean: Mediterranean, Equatorial Africa; Northwestern Atlantic Ocean: United States coast from 40 N to Caribbean Sea; Gulf of Mexico; Southwestern Atlantic Ocean: Campos Basin, Rio Grande Rise. Benthic (878-4,720 m) (Sula, 1977). In Brazil found between 1,302 and 1,644 m depth (Franco, 2007). Records in the present investigation include: Santos Basin from 2,131 to 2,188 m depth (Lapa), and Campos Basin from 2,899 to 2,986 m depth (C-M-541) (Figure 2). COMMENTS Adults are benthic, while juveniles were captured in midwater trawls. The elaborate pectoral fins of Bathypterois are undoubtedly employed as sensory devices to detect the presence of its planktonic prey (copepods and other crustaceans). The stiff elongate pelvic and caudal fin rays seem to serve to elevate the fish above the substrate. Figure 1. Tripodfish Bathypterois grallator (Goode & Bean, 1886) observed in Campos Basin at 2,985 m depth. Figure 2. Records of Bathypterois grallator (Goode & Bean, 1886) in two of the investigated areas: Lapa, in Santos Basin, and C-M-541, in Campos Basin. AULOPIFORMES/ IPNOPIDAE Atlas of Benthic Fauna 42 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 43 AULOPIFORMES/ IPNOPIDAE ACTINOPTERYGII Bathypterois phenax Parr, 1928 BIBLIOGRAPHY Franco, M.A.L., Braga, A.C., Nunan, G.W.A. & Costa, P.A.S. 2009. Fishes of the family Ipnopidae (Teleos- tei: Aulopiformes) collected on the Brazilian continental slope between 11° and 23°S. Journal of Fish Biology 75: 797–815. Merrett, N.R., 1990. Chloro- phthalmidae. p. 351-360. In J.C. Quero, J.C. Hureau, C. Karrer, A. Post and L. Salda- nha (eds.) Check-list of the fishes of the eastern tropical Atlantic (CLOFETA). JNICT, Lisbon; SEI, Paris; and UNES- CO, Paris. Vol. 1. Sulak, K.J., 1977. The syste- matics and biology of Ba- thypterois (Pisces, Chloro- phthalmidae). Galathea Rep, 14, pp.49-108. DIAGNOSIS Body elongated, covered by large well marked scales, dark grey to black in color; head dark, with minute eyes, and black fins; pectoral fins thrown forward of the head (Figure 1). DISTRIBUTION The species is only known from the Atlantic Ocean, archibenthic to abyssobenthic in habit. Eastern Atlantic Ocean: Off Sierra Leone and Liberia (Merrett, 1990) and off South Africa; Western Atlantic Ocean: off the North American coast from latitude 39° to 9° N, including the Gulf of Mexico and Caribbean Sea (Sulak, 1977) and off Brazil from latitude 13° to 23° S, between 900 and 2,100 m depth (Franco et al., 2009). The records in the present investigation include: Santos Basin from 2,130 to 2,153 m depth (Lapa), and Campos Basin from 2,983 to 2,997 m depth (C-M-541) (Figure 2). COMMENTS This species is frequently observed inhabiting the middle and lower regions of the continental slope. It exhibits a benthic resting behavior, employing its pelvic fins and the lower rays of the caudal fin for support. While adult individuals exhibit benthic behavior, juvenile specimens exhibit pelagic habits. Its primary dietary source consists of benthopelagic plankton. Figure 1. Blackfin spiderfish Bathypterois phenax Parr, 1928 observed in Campos Basin at 2,998 m depth. Figure 2. Records of Bathypterois phenax Parr, 1928 in the two of the investigated areas: Lapa, in Santos Basin, and C-M-541, in Campos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms44 | TotalEnergies ACTINOPTERYGII/ AULOPIFORMES/ BATHYSAURIDAE FEROX Bathysaurus cf. ferox Günther, 1878 DIAGNOSIS Body elongated, cylindrical, grey to brown in color; head greatly depressed, bony, with large greenish-black eyes and an enormous mouth bearing multiple series of long, sharp, needle- like teeth; dorsal fin long-based; caudal fin paddle-like; fins often dark, especially ventral, pelvic, and pectoral (Figure 1). DISTRIBUTION Bathysaurus ferox Günther, 1878 exhibits a broad, nearly cosmopolitan distribution in deep waters, occurring in the Atlantic, Pacific, and Indian Oceans. In the Atlantic, it has been recorded from the Davis Strait to the Rio Grande Rise, Brazil, including the Northeastern Atlantic (from the British Isles to the Gulf of Guinea) and the South Atlantic, with additional records off South Africa. In the Pacific, it occurs in the southwestern (Australia and New Zealand) and southeastern regions (Nazca and Sala y Gómez ridges). The species is typically bathydemersal, occurring at depths ranging from approximately 600 to 3,500 m, most commonly between 1,000 and 2,500 m, associated with the continental slope and abyssal plain. Occurrence data compiled from global databases such as GBIF support this wide distribution, with records dispersed across different ocean basins, reflecting a circumglobal pattern in deep-sea environments. Records in the present investigation include: Santos Basin from 1,614 to 1,626 m depth (S-M-11) and at 2,122 m depth (Lapa), and from Campos Basin from 2,976 to 2,986 m depth (C-M-541) (Figure 2). COMMENTS Two species of Bathysaurus are known from the Brazilian coast: Bathysaurus ferox Günther, 1878, and Bathysaurus mollis Günther, 1878. These are the only recognized species of the genus and function as apex predators in deep-sea abyssal environments. Although both share a lizard-like appearance and hermaphroditic nature, they differ in morphology, depth preference, and fin structure. Bathysaurus mollis is typically pale and unpigmented, with flabby, translucent flesh, whereas Bathysaurus ferox is darker, with grey-brown pigmentation. Adults of both species are predominantly benthic. Figure 1. Bathysaurus cf. ferox Günther, 1878 observed in Santos Basin at 1,626 m depth. Figure 2. Records of Bathysaurus cf. ferox Günther, 1878, in three of the investigated areas: S-M-1711 and Lapa, in Santos Basin and C-M-541, in Campos Basin. BIBLIOGRAPHY Froese, R.; Pauly, D. (eds.). FishBase. Bathysaurus ferox. Available at: https://www. fishbase.org. GBIF. Bathysaurus ferox Günther, 1878. Available at: https://www.gbif.org/spe- cies/2401415. Bray, D.J. Bathysaurus ferox. Fishes of Australia. Available at: https://fishesofaustralia. net.au. Perez, J. A. A.; Gavazzoni, L.; Souza, L. H. P.; Sumida, P. Y. G.; Kitazato, H. 2020. Deep- -sea habitats and megafau- na on the slopes of the São Paulo Ridge, SW Atlantic. Frontiers in Marine Science, v. 7, 572166. DOI: 10.3389/ fmars.2020.572166 Menezes, N.A., P.A. Buckup, J.L. Figueiredo and R.L. Mou- ra, 2003. Catálogo das espé- cies de peixes marinhos do Brasil. São Paulo. Museu de Zoologia da Universidade de São Paulo, 160 pp. Atlas of Benthic Fauna 44 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 45 ACTINOPTERYGII BIBLIOGRAPHY Bergstad, O.A., L Clark, H.Ø. Hansen and N. Cousins, 2012. Distribution, population bio- logy, and trophic ecology of the deepwater demersal fish Halosauropsis macrochir (Pis- ces: Halosauridae) on the Mi- d-Atlantic Ridge. PLoS ONE 7(2): e31493. doi:10.1371/jour- nal.pone.0031493 Cohen, D.M., A.W. Ebeling, T. Iwamoto, S.B. McDowell, N.B. Marshall, D.E. Rosen, P. So- noda, W.H. Weed III and L.P. Woods, 1973. Fishes of the western North Atlantic. Part six. New Haven, Sears Found. Mar. Res., Yale Univ. Mincarone, M. M., Martins, A. S., Costa, P. A. S., Braga, A. C. & Haimovici, M., 2017. Peixes ma- rinhos da Bacia de Campos: uma revisão da diversidade. In Comunidades Demersais e Bioconstrutores: caracteriza- ção ambiental regional da Ba- cia de Campos, Atlântico Su- doeste (Curbelo-Fernandez, M. P. & Braga, A. C., eds), pp. 187–216, vol. 4. Rio de Janeiro: Elsevier. Sulak, K.J., 1986. Halosauridae. p. 593-598. In P.J.P. Whitehe- ad, M.-L. Bauchot, J.-C. Hure- au, J. Nielsen and E. Tortonese (eds.) Fishes of the north-eas- tern Atlantic and the Mediter- ranean. UNESCO, Paris. vol. 2. DIAGNOSIS Body rather elongate; greyish-black to bluish-black in color. Absence of scales on the top of the head and presence of scales on the opercular bone. Adult length reaches 60 cm (Figure 1). DISTRIBUTION Western Indian Ocean; Eastern Indian Ocean; Antarctic Indian Ocean; Northwest Pacific Ocean; Southwest Pacific Ocean; Eastern Central Atlantic Ocean; Western Central Atlantic Ocean; Northwestern Atlantic Ocean; Northeastern Atlantic Ocean; Southwestern Atlantic Ocean; Southeastern Atlantic Ocean; Antarctic Atlantic Ocean. The records in the present investigation include: Santos Basin, from 1,455 to 1,498 m depth (S-M-1815), from 1,482 to 1,647 m depth (S-M-1711) and from 2,131 to 2,216 m depth (Lapa) (Figure 2). COMMENTS The species is bathydemersal. Feeds on polychaetes, crustaceans, echinoderms and echiurids. Hovers above or rests on bottom. Long slender pectorals are commonly held erect pointing upward and forward, these may serve a sensory function (Sulak, 1986). Figure 1. Abyssal halosaur, Halosauropsis macrochir (Günther, 1878) observed in Santos Basin at 1,455 m depth. Figure 2. Records of Halosauropsis macrochir (Günther, 1878) in three of the investigated areas: S-M-1815, S-M-1711 and Lapa, in Santos Basin. NOTACANTHIFORMES/ HALOSAURIDAE Halosauropsis macrochir (Günther, 1878) Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms46 | TotalEnergies ACTINOPTERYGII BIBLIOGRAPHY MINCARONE, M.M. 2010. In: Lavrado, H.P. & Brasil, A.C.S. Biodiversidade da Região Oceânica Profunda da Ba- cia de Campos: Megafauna e Ictiofauna Demersal. 350 – 351 p. MINCARONE, M.M.; NIEL- SEN, J.G.; COSTA, P.A.S. 2008. Deep-sea ophidiiform fishes collected on the Bra- zilian continental slope, be- tween 11° and 23°S. Zootaxa, v. 1770, p. 41-64. Nielsen, J.G., 1986. Ophi- diidae. p. 1158-1166. In P.J.P. Whitehead, M.-L. Bauchot, J.-C. Hureau, J. Nielsen and E. Tortonese (eds.) Fishes of the North-eastern Atlan- tic and the Mediterranean. UNESCO, Paris. Vol 3. Nielsen, J.G., D.M. Cohen, D.F. Markle and C.R. Robins, 1999. Ophidiiform fishes of the world (Order Ophidii- formes). An annotated and illustrated catalogue of pe- arlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Fish. Synop. 125(18):178p. Rome: FAO. DIAGNOSIS Robust, elongated body; rounded snout; small eyes; pectoral fin elongated, about head length; greyish-white body with dark fins. Adult length: 28-31 cm (Figure 1). DISTRIBUTION Eastern Atlantic: off the Azores and south of the Canary Islands (Ref. 6555). Western Atlantic: USA; Bahamas and Brazil. In Brazil, reported only off Rio de Janeiro; found between 1,122 and 1,147 m depth (Mincarone, 2010). The record in the present investigation: Santos Basin (Lapa), at 2,188 m depth. COMMENTS Rare species; unknown biology and ecology (Mincarone, 2010). Figure 1. Holcomycteronus squamosus (Roule, 1916) observed in the Santos Basin at 2,188 m depth. Figure 2. Record of Holcomycteronus squamosus (Roule, 1916) in one of the investigated areas: Lapa, in Santos Basin. OPHIDIIFORMES/ OPHIDIIDAE Holcomycteronus squamosus (Roule, 1916) Atlas of Benthic Fauna 46 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 47 ACTINOPTERYGII BIBLIOGRAPHY Franco, M.A.L., Braga, A.C., Nunan, G.W.A. & Costa, P.A.S. 2009. Fishes of the family Ipnopidae (Teleos- tei: Aulopiformes) collected on the Brazilian continental slope between 11° and 23°S. Journal of Fish Biology 75: 797–815. Haedrich, R.L. and N.R. Mer- rett, 1988. Summary atlas of deep-living demersal fishes in the North Atlantic Basin. J. Nat. Hist. 22:1325-1362. DIAGNOSIS Body slender, moderately compressed dorsoventrally, covered with distinct dark gray scales; dark head with bioluminescent eye spots on top; translucent caudal fin (Figure 1). DISTRIBUTION This species is considered restricted to the Atlantic Ocean (Franco et al. 2009). Eastern Atlantic Ocean: North Africa; Northwestern Atlantic Ocean: Gulf of Mexico and Caribbean Sea to the South Atlantic Ocean: west of Tristan da Cunha and along the Brazilian coast, from latitude 10º to 21º S (Franco et. al. 2009). Records in the present investigation include: Santos Basin at 1,497 m depth (S-M-1815) and 2,133 m depth (Lapa) (Figure 2). COMMENTS The pelagic larvae have normal eyes. During development and metamorphosis into the adult form, the eyes become flattened and enlarged, losing the optic nerve and lens in the process. Figure 1. Ipnops murrayi Günther, 1878 observed in the Santos Basin at 2,133 m depth. Figure 2. Records of Ipnops murrayi Günther, 1878 in two of the investigated areas, S-M-1815 and Lapa in Santos Basin. AULOPIFORMES/ IPNOPIDAE Ipnops murrayi Günther, 1878 Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms48 | TotalEnergies ACTINOPTERYGII BIBLIOGRAPHY Costa, P.A.S., Braga, A.C., Melo, M.R.S., Nunan, G.W., Martins, A.S., Olavo, G. 2007. Assembléias de teleósteos demersais no talude da cos- ta central brasileira. In: Cos- ta, P.A.S., Olavo, G., Nunan, G.W.A., et. al. (Eds.), Biodi- versidade da Fauna Marinha Profunda na Costa Central Brasileira - Série Livros 24. Museu Nacional, Rio de ja- neiro, pp. 87–107. Froese, R. and Pauly, D. Edi- tors. 2023. FishBase. Ma- crouridae Bonaparte, 1831. Accessed through: World Register of Marine Species at: https://marinespecies. org/aphia.php?p=taxdetail- s&id=125471 on 2023-11-20. Nóbrega, M.F., Garcia Júnior, J., Oliveira, M. A., Oliveira, J.E. L. 2023. Continental slope fishes of the Potiguar Basin off Northeast Brazil in the Equatorial Atlantic Oce- an. Deep-Sea Research Part I-Oceanographic Research Papers, v. 192, p. 103937. DIAGNOSIS Elongated body, large head, and short trunk ending in a long, pointed tail. No caudal fin. One or two dorsal fins possible. True fin spines lacking. Leading dorsal fin ray may be spinous. Anal and dorsal fins merge, anal fin base larger than second dorsal fin. (Figure 1). DISTRIBUTION This family occur in all oceans (except the high Arctic), they are primarily benthopelagic deep-sea fishes; a few species are bathypelagic. Most species are found between 200 m and 2,000 m, one species to more than 7,012 m. In Brazil found between 100 and 2,220 m depth (Costa, 2010 and Nóbrega et. al. 2023). The records in the present investigation: Santos Basin at between 1,150 and 2,135 m depth (S-M-1815) at 2,135 m depth (Lapa), and Campos Basin (C-M-541) at 2,870 and 3,000 m depth (Figure 2). COMMENTS The family has 38 genera distributed in four subfamilies (Froese & Pauly, 2023). In the Campos Basin, there are 22 known species, classified into 14 genera. Figure 1. Macrouridae Bonaparte, 1831 observed in the Campos Basin at 3,001 m depth. Figure 2. Records of Macrouridae Bonaparte, 1831 in three of the investigated areas: S-M-1815 and Lapa in Santos Basin, and C-M-541, in Campos Basin. GADIFORMES/ MACROURIDAE Macrouridae Bonaparte, 1831 Atlas of Benthic Fauna 48 | TotalEnergies
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 49 BIBLIOGRAPHY Carpenter, K.E. (Ed.) 2002. The living marine resour- ces of the Western Central Atlantic. Volume 2: Bony fishes part 1 (Acipenseridae to Grammatidae). FAO Spe- cies Identification Guide for Fishery Purposes and Ame- rican Society of Ichthyo- logists and Herpetologists Special Publication No. 5. Rome, FAO. 2002. pp. 601- 1374. Melo, M.R.S. 2010. In: Lavra- do, H.P. & Brasil, A.C.S. Bio- diversidade da Região Oce- ânica Profunda da Bacia de Campos: Megafauna e Ictio- fauna Demersal. 314 – 321 p. DIAGNOSIS Small to medium-sized eels, the largest species reaching 1.8 m, but most less than 1 m. Body stout to elongate, reaching 1.8 m, but most less than 1 m; plain brown or grey in color, sometimes countershaded. Head variable. Eye well developed to reduced. Snout may be short and blunt or moderately elongate.Mouth usually large; jaws nearly equal, sometimes snout projects slightly beyond lower jaw and sometimes vice versa. Anterior nostril tubular, near tip of snout; posterior nostril on side of snout, at or below mideye level. Gill openings low on body, belowpectoral fins (when present). Dorsal and anal fins well developed, confluent with caudal fin. Pectoral fins present or absent. Scales present or absent. Lateral line variable, often complete, sometimes reduced to a few pores at anterior end and sometimes no pores at all (Carpenter, 2002) (Figure 1). DISTRIBUTION Worldwide (Atlantic, Indian and Pacific); tropical and temperate. Habitat: benthic, deep water or deep-sea (larvae pelagic); depth range 0 - 2,820 m; common 450 - 600m. In Capos Basin 750 – 200m (Melo, 2010). Records in the present investigation: Santos Basin from 1,148 to 1,497 m depth (S-M-1815), from 1,487 to 1,847 m depth (S-M-1711), and at 2,122 m depth (Lapa), and Campos Basin at 2, 861 m depth (Figure 2). COMMENTS The Synaphobranchidae is divided into three subfamilies. The Simenchelyinae comprises a single genus and species, Simenchelys parasitica. The Synaphobranchinae includes species most representative of the family, distributed across four genera. The Ilyophinae is the most species-rich and morphologically diverse subfamily within Synaphobranchidae, encompassing seven genera. Ilyophines exhibit considerable variation in body shape, dentition, presence or absence of pectoral fins, eye size, and snout ornamentation. Most lack scales, and in some species, the anus is positioned far forward, nearly under the pectoral fins. ACTINOPTERYGII ANGUILLIFORMES/ SYNAPHOBRANCHIDAE Synaphobranchidae Johnsons, 1862 Figure 1. Synaphobranchidae Johnson, 1862 observed in the Santos Basin at 1,497 m depth. Figure 2. Records of Synaphobranchidae Johnson, 1862 in the four investigated areas: S-M-1815, S-M-1711, and Lapa, in Santos Basin, and C-M-541, in Campos Basin. Atlas of Benthic Fauna
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021 Atlas of Benthic Organisms50 | TotalEnergies CONCLUSION In the vast and little-known depths of the Southwestern Atlantic, life thrives in forms that remain largely invisible to science. Documenting this hidden world is essential, not only to appreciate its complexity but to ensure its preservation. This atlas was created with that ambition. It gathers several benthic taxa documented during offshore studies and transforms observations into shared knowledge. More than a compilation of records, this atlas reflects TotalEnergies’ commitment to promoting biodiversity and stands as a concrete example of collaboration with the scientific community. LOOKING AHEAD, OUR GOAL IS CLEAR: to deepen understanding of the invaluable biodiversity of South Atlantic deep-sea ecosystems by sharing data and inspiring similar initiatives to strengthen knowledge about this unique natural heritage for generations to come.
MAGENTA YELLOWCYAN 30-34 Rue du Chemin Vert 75011 Paris +33 (0)1 85 56 97 00 www.carrenoir.com Ce fichier est un document d’exécution créé sur Illustrator version CS6. TECHNIQUETONS RECOMMANDÉS TOTAL TOT_21_00008_TotalEnergies_Logo_CMYK JFB/MG Date : 13/07/2021TotalEnergies | 51 TECHNICAL TEAM Organizers- TotalEnergies EP Brazil Aline Wyllie Lacerda Rodrigues, MSc. Maxence Chéreau, MSc. Organizers- ATLANTIS SMA Maria Patricia Curbelo Fernandez, DSc. Alexandre Luiz Neves Borges, MSc. Organizer- Environpact Vinicius Couto Alves Taxonomists Paulo Marcio Santos Costa, DSc., FIPERJ Brazil Rafael Bendayan de Moura, DSc., IB-UFRJ Brazil Renata Carolina Mikosz Arantes, DSc., UFSC Brazil Vinicius da Rocha Miranda, DSc. IB- UFRRJ Brazil Layout Brandesign
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