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Original Article • Nauplius 33 • 2025 • https://doi.org/10.1590/2358-2936e20240539 linkcopy
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Open-access Holocene Bythocytheridae (Ostracoda: Podocopida) from Southwestern Atlantic deep-sea sediments off Brazil. Part 1: tribes Bythocytherini Sars, 1926 and Jonesini Schornikov, 1981

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    ABSTRACT

    The Bythocytheridae Sars, 1866 is one of the oldest families of cytheroidean Ostracoda, with a fossil record dating back to the Silurian. During the Cenozoic this family became one of the main constituents of the deep-sea ostracod faunas, with some taxa reaching wide geographic distribution. Bythocytherids are, therefore, important for the understanding of evolution and paleobiogeography in marine Ostracoda. The main purpose of this paper is to present a study of the tribes Bythocytherini Sars, 1926 and Jonesini Schornikov, 1981 through the analysis of 47 sediment samples collected from 12 sites in the Rio Grande Rise and adjacent areas. The 11 species herein recorded are attributed to the genera Bythocythere Sars, 1866 (four species), Bythoceratina Hornibrook, 1952 (two species), Retibythere Schornikov, 1981 (two species), Rhombobythere Schornikov, 1982b (two species), and Ruggieriella Colalongo and Pasini, 1980 (one species). The occurrence of the bythocytherid species in the study area seems strongly influenced by the bathymetry, being more abundant in the shallowest site (~1,250 m) and absent beyond 3,000 m water depth.

    Keywords:
    Extreme environments; Marine paleobiodiversity; Micropaleontology; Paleoceanography; Rio Grande Rise

    INTRODUCTION

    The family Bythocytheridae is one of the oldest cytheroidean lineages, with a fossil record ranging from the Middle Silurian to the Holocene (Schornikov, 1988a ; 1990). The carapaces of most bythocytherids are characterized by three morphological features: five adductor scars disposed in a row, an adont/merodont hinge, and a tube-like caudal process. According to Brandão et al. (2023), this family consists of 118 genera, 13 subgenera, and 871 species both living and extinct, some of them reaching wide geographic distributions. The genera PseudocythereSars, 1866 and BythoceratinaHornibrook, 1952 are particularly widespread in the deep-sea Quaternary deposits, although some species also occur in shallow waters (= neritic environment) (e.g., Athersuch et al., 1983; Mostafawi, 2000; Wilson, 2006; Tanaka et al., 2019).

    The carapace morphology of Quaternary bythocytherids varies from smooth (e.g., SclerochilusSars, 1866) to highly sculptured forms, with spines, reticula, and tubercles (e.g., Bythoceratina). Species of some genera (e.g., CytheralisonHornibrook, 1952 and Pseudocythere) hold morphological features shared with taxa of the Paleozoic Thuringian Ecotype (Ordovician-Carboniferous) such as few and well-developed spines in the anterior and posterior margins as well as in the middle of the valves (e.g., Becker et al., 1993; Becker and Blumenstengel, 1995). The diversity of carapace morphologies and a conservative limb morphology in Bythocytheridae have led Schornikov (1988 b ) and Schornikov and Mikhailova (1990) to propose the concept of cyclicity of morphogenesis. According to them, the emergence of a structure can be shifted to earlier or advanced instars according to its adaptiveness. How these morphogenetic processes evolved in bythocytherids is not well understood yet, but the resulting morphological diversity might have contributed to the pandemic and eurybathic patterns observed in several genera.

    The taxonomic study of the Quaternary bythocytherids in Brazil began with Coimbra and Fauth (2002) who proposed two new Bythoceratina species from the Brazilian equatorial shelf. Other Quaternary bythocytherids from the South and Southeastern Brazilian margin include either shallow (neritic) or deep-water occurrences of Pseudocythere, Bythoceratina, and Jonesia Brady, 1866 (Drozinski et al., 2003; Bergue et al., 2006; 2017; 2019; Bergue and Coimbra, 2008; Machado et al., 2020). Southward, along the Argentinian coast, the occurrence of Pseudocythere, Bythoceratina, BythocythereSars, 1866, and NealocythereSchornikov, 1982 a was documented by Whatley et al. (1996; 1997a; 1977b). Recently, Bythoceratina bonaterrae Bergue and Coimbra in Bergue et al., 2021 was described from the continental slope of the Camamu Basin, Northeastern Brazilian margin (Bergue et al., 2021).

    The main objective of this article is to study Holocene bythocytherids of the tribes Bythocytherini Sars, 1926 and Jonesini Schornikov, 1981 in the Southwestern Atlantic Ocean. Moreover, it contributes to the understanding of the bathymetric zonation of their species and the zoogeographic distribution of some genera. The tribe Pseudocytherini Schneider, 1960 will be detailed in another paper. The studied material was obtained in an area including the Vitória-Trindade Chain and the São Paulo Plateau, which records tectonic processes related to the formation of the Rio Grande Rise (RGR) and geologic features linked to magmatism and continental crust fractures (Galvão and Castro, 2017) (Fig. 1). According to Alberoni et al. (2020), these bathymetric highs are characteristic of the Brazilian Meridional Margin, which ranges from the Vitória-Trindade Ridge to the marine limit with Uruguay. The sampled sites are currently under the influence of the North Atlantic Deep Water (NADW) and the Antarctic Bottom Water (AABW) (Silveira et al., 2000). The RGR acts as a huge barrier to the transport of sediments between the oceanic basins of Brazil and Argentina, and affects the northward circulation of the AABW (Galvão and Castro, 2017; Morozov et al., 2020).

    thumbnail of Figure 1.
    Figure 1.
    Study area with location of the sampled sites.

    MATERIAL AND METHODS

    The 47 samples analyzed in this work are composed of biogenic calcareous muds, mostly foraminiferal and nannofossil oozes (Tab. 1). They were obtained from the push cores collected during the Iatá-Piuna/Quelle cruise on board of the R/V “Yokosuka” from the Japan Agency for Marine Earth Science and Technology (JAMSTEC), between 23 April 2013 and 24 May 2013 (Fig. 2). This region was investigated in a broader project named Quelle 2013 (the acronym for Quest for the Limit of Life), which aimed at the study of marine communities from extreme environments as it displays areas of cold seepage. These cores were sampled in different intervals depending on their length, but according to planktonic foraminiferal distribution, all samples are Holocene in age (Bergue et al., 2023).

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    Table 1.
    Location, mass, and bathymetry of the samples analyzed in this study.

    thumbnail of Figure 2.
    Figure 2.
    Occurrence of species in the studied cores. Numbers along the 1338C1 core correspond to the depth in meters.

    The samples were washed through a 0.062 mm mesh with tap water and their resulting residues thoroughly examined for ostracods. The supra-generic taxonomy herein adopted follows Schornikov and Mikhailova (1990) and Brandão et al. (2023). All illustrated material is held at Museu de Paleontologia Irajá Damiani Pinto, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil. Morphological abbreviations: H = height; L = length; LV = left valve; RV = right valve; V = valve. Institutional abbreviations: DSDP: Deep Sea Drilling Project; JAMSTEC: Japan Agency for Marine Earth Science and Technology; MP-O: Museu de Paleontologia Irajá Damiani Pinto, Seção Ostracoda, Porto Alegre, Brazil; ODP: Ocean Drilling Program; OS: Ostracod Collection of the Natural History Museum; USNM: United States National Museum, Washington DC, USA.

    SYSTEMATICS

    Class Ostracoda Latreille, 1802

    Subclass Podocopa Müller, 1894

    Order Podocopida Sars, 1866

    Suborder Cytherocopina Baird, 1850

    Superfamily Cytheroidea Baird, 1850

    Family Bythocytheridae Sars, 1926

    Subfamily Bythocytherinae Sars, 1926

    Tribe Bythocytherini Sars, 1926

    Diagnosis. Carapace with hinge margin straight, caudal process usually tube-like, positioned in continuation of dorsal margin, or strongly reduced in some cases. Macrosculpture usually represented by alar process, spines and crossed ridges, nodes and crossed sulci. Mesosculpture reticulated. External marginal ridge present. Inner sculpture might present one or two dike-like alae. Concrescence zone wide with simple marginal pore-canals. Vestibulum usually pronounced in anterior half. Besides wide normal pore-canals with bristles, conical pore-canals without bristles might occur. Fulcral point conspicuous (adapted from Schornikov, 1981).

    Genus Bythocythere Sars, 1866

    Type-species. Bythocythere turgidaSars, 1866 by original designation.

    Remarks. Bythocythere is an eurybathic genus with wide geographic distribution, including Indian, Pacific, and Austral oceans. In some of these occurrences it has been misidentified as, for instance, ?Rhombobythere sp. 2 in Didié and Bauch (2001) and ?Rhombobythere sp. 1 in Corrège (1993), which correspond to Bythocythere instead.

    Bythocythere bathyatos Whatley and Coles, 1987

    (Fig. 3 A )

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    Figure 3.
    SEM images of the recorded species. Bythocythere bathyatosWhatley and Coles, 1987: A, (MP-O-3170) RV external view. Bythocythere eugeneschornikoviYasuhara et al., 2009 a : B, RV external view (MP-O-3101); C, LV external view (MP-O-3171). Bythocythere sp. 1: D, female RV external view (MP-O-3108); E, male RV external view (MP-O-3172). Bythocythere sp. 2: F, RV external view (MP-O-3173); G, LV external view (MP-O-3174); H, LV internal view (MP-O-3175); I, (A-2?) RV external view (MP-O-3176); J, (A-1?) LV external view (MP-O-3177). Bythoceratina sp. 1 (MP-O-3178): K, LV external view; L, internal view of the same specimen. Bythoceratina sp. 2: M, (MP-O-3179) RV external view. Retibythere sp. 1: N, RV external view (MP-O-3107); O, internal view of the same specimen. Scale bars = 0.1 mm.

    Bythocythere bathyatosWhatley and Coles, 1987: 59, pl. 1, figs. 26-31. - Didié and Bauch, 2001: 107, pl. 3, figs. 27, 28. - Alvarez-Zarikian, 2009: 3, pl. 2, fig. 3. - Jöst et al., 2022: 325, pl. 3, fig. C.

    Holotype. Female RV OS 12501.

    Type-locality and age. DSDP Site 607, core 18 (41°00.07’N; 32°57.44’W). Early Pliocene.

    Material examined. MP-O-3170 (sample 1338C5 0.02 m) RV, L = 0.48 mm, H = 0.28 mm.

    Occurrence and material. Samples 1343C8 0.0 m (1v), 1336C1 0.06 m (1v), and 1338C5 0.02 m (1v).

    Stratigraphic and geographic distributions. Early Pliocene: North Atlantic, DSDP Site 607 (Whatley and Coles, 1987). Quaternary: North Atlantic (Didié and Bauch, 2001; Alvarez- Zarikian, 2009). Holocene: North Atlantic (Jöst et al., 2022). Southwestern Atlantic (this study).

    Bythocythere eugeneschornikovi Yasuhara et al., 2009a

    (Fig. 3 B , C)

    Bythocythere eugeneschornikoviYasuhara et al., 2009 a : 890, pl. 2, figs. 5-10. - Bergue et al., 2021: 4, fig. 3S. - Bergue et al., 2023: 11, fig. 3O.

    Holotype. Male LV, USNM 537145.

    Type-locality and age. ODP Site 1055B, sample 94-96 cm, North Atlantic (32°47.041’N 76°17.179’W). Quaternary.

    Material examined . MP-O-3101 (sample 1343C8 0.0 m) RV, L = 0.80 mm, H = 0.40 mm (Fig. 3 B ). MP-O-3171 (sample 1346C7 0.0 m) LV, L = 0.83 mm, H = 0.40 mm (Fig. 3 C ).

    Occurrence and material. Samples 1343C8 0.0 m (2v), 1344C5 0.0 m (2v), and 1347C6 0.0 m (3v).

    Stratigraphic and geographic distribution. Pleistocene-Holocene: Camamu Basin, Brazil (Bergue et al., 2021). Quaternary: ODP Site 1055 (Yasuhara et al., 2009 a ). Holocene: São Paulo Plateau, Southwestern Atlantic (Bergue et al., 2023; this study).

    Bythocythere sp. 1

    (Fig. 3 D , E)

    Bythocythere mylaensisSciuto, 2009. - Bergue et al., 2023: 12, figs. 4e, f.

    Material examined. MP-O-3108 (sample 1345C6 0.0 m) female RV, L = 0.62 mm, H = 0.40 mm (Fig. 3 D ); MP-O-3172 (Sample 1338C1 0.02 m), male RV, L = 0.52 mm, H = 0.33 mm (Fig. 3 E ).

    Occurrence and material. Samples 1345C6 0.0 m (1v) and 1338C1 0.02 m (2v).

    Stratigraphic and geographic distributions. Holocene: São Paulo Plateau, Southwestern Atlantic (Bergue et al., 2023; this study).

    Remarks. Bergue et al. (2023) identified one of the specimens herein illustrated (MP-O-3108, figure 3D) as Bythocythere mylaensisSciuto, 2009. In the present study, two RV with more inflated posteroventral region were recorded. We consider this morphological difference as sexual dimorphism, which in some Bythocytherini is expressed in the height/length ratio (Schornikov, 1982 a ). However, considering that sexual dimorphism was not mentioned in the description of B. mylaensis, we prefer to keep the specimens herein studied in open nomenclature. Bythocythere sp. 1 is very similar to ?Rhombobythere sp. 2 in Didié and Bauch, 2001 and ?Rhombobythere sp. 1 in Corrège, 1993 recorded, respectively, in the North Atlantic and in the Coral Sea (Australasia), but differs in the size of the puncta and in the outline of the caudal process.

    Bythocythere sp. 2

    (Fig. 3 F -J)

    Material examined. MP-O-3173, RV, L = 0.55 mm, H = 0.32 mm (Fig. 3 F ). MP-O-3174 (sample 1338C5 0.05 m) LV, L = 0.60 mm, H = 0.32 mm (Fig. 3 G ). MP-O-3175 (1338C5 0.05 m) LV, L = 0.57 mm, H = 0.30 mm (Fig. 3 H ). MP-O-3176 (sample 1338C5 0.09 m) (A-2?) RV, L = 0.49 mm, H = 0.25 mm (Fig. 3 I ). MP-O-3177 (sample 1338C5 0.03 m) (A-2?) LV, L = 0.40 mm, H = 0.22 mm (Fig. 3 J ).

    Occurrence and material. Samples 1338C1 0.08 m (1v), 1338C5 0.03 m (1v), 1338C5 0.05 m (2v), and 1338C5 0.09 m (1v).

    Remarks. Bythocythere sp. 2 is similar to Bythocythere sp. 1 in terms of surface sculpture but is lower and differs in the outline of the posterior margin. The weakly developed reticulation and the oblique ventral outline observed in the juveniles (Fig. 3I, J) make this species similar to ?Rhombobythere sp. 3 in Didié and Bauch (2001). However, the material herein figured has anterior margin less rounded and caudal process more obtuse.

    Genus Bythoceratina Hornibrook, 1952

    Type-species. Bythoceratina mestayeraeHornibrook, 1952, by original designation.

    Bythoceratina sp. 1

    (Fig. 3 K , L)

    Material examined. MP-O-3178 (sample 1335C1 0.01 m) LV, L = 0.80 mm, H = 0.43 mm.

    Occurrence and material. 1335C1 0.01 m (1v).

    Remarks. This very reticulated specimen differs from the other species of Bythoceratina in having an obliquely rounded anterior margin, and a slight convexity in the middle of the dorsal margin. The solum of each fossa is pierced by two or three pits. Bythoceratina sp. 1 is probably a new species, but the single specimen obtained does not allow a taxonomic description.

    Bythoceratina sp. 2

    (Fig. 3 M )

    Material examined. MP-O-3179 (sample 1344C6 0.0 m) RV, L = 0.9 mm, H = 0.5 mm (approximately).

    Occurrence and material. Sample 1344C6 0.0 m (1v).

    Remarks. It is unclear whether the specimen here illustrated is an adult. However, it holds all characteristics of the genus.

    Genus Retibythere Schornikov, 1981

    Type-species. Retibythere bialataSchornikov, 1981, by original designation.

    Remarks. Schornikov (1981) proposed the genus Retibythere based on living specimens from the far-east marine regions off Soviet Union (Okhotsk Sea and Japan Sea). Subsequently, Schornikov (1987) described a new subgenus Retibythere (Bathybythere) and reassigned to it the species Cytherura scaberrimaBrady, 1886. Other published combinations of this species are Bythoceratina scaberrima (Brady, 1886) and Cythere scaberrima (Brady, 1886). In terms of carapace morphology, Retibythere differs from Bythoceratina by the presence of non-crenulated median hinge element. Moreover, the presence of a ventral tooth-socket distinguishes Retibythere from the remaining Bythocytherini. According to Brandão et al. (2023), the following taxa are presently ascribed to Retibythere: Retibythere (Bathybythere) Schornikov, 1987, Retibythere (Bathybythere) scaberrima (Brady, 1886), Retibythere (Bathybythere) vandenboldi (Ruggieri, 1960), Retibythere (Retibythere) Schornikov, 1981, Retibythere (Retibythere) acutialataSchornikov, 1981, and Retibythere (Retibythere) bialataSchornikov, 1981. The two species of Retibythere recorded in the present study have socket-like structures at the ventral margin of the RV but differ in the hinge median element. Although external similarity and the presence of the tooth-socket on the ventral margin are strong morphological evidence to place them in the same genus, the hinge morphology of Retibythere remains an important issue to be investigated to reinforce this hypothesis.

    Retibythere sp. 1

    (Fig. 3 N , O; 4A, B)

    Bythoceratina sp. Bergue et al., 2023: 12, fig. 4c, d.

    Material examined. MP-O-3107 adult RV (sample 1344C5 0.0 m) L = 0.78 mm, H = 0.4 mm (Figs. 3N-O, 4A). MP-O-3180, A-1 LV (sample 1335C1 0.0 m) L = 0.62 mm, H = 0.31 mm (Fig. 4 B ).

    Occurrence and material. Samples 1335C1 0.0 m (1v) and 1344C5 0.0 m (1v).

    Remarks. This species differs from Retibythere (Retibythere) bialataSchornikov, 1981 in having only one anterodorsal tubercle.

    Retibythere sp. 2

    (Fig. 4 C -E)

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    Figure 4.
    SEM images of the recorded species. Retibythere sp. 1: A, detail of the ventral margin (MP-O-3107); B, juvenile LV external view (MP-O-3180). Retibythere sp. 2: C, RV external view (MP-O-3181); D, LV external view (MP-O-3182); E, RV (MP-O-3181) internal view. Rhombobythere sp. 1: F, LV external view (MP-O-3183); G, same specimen internal view. Rhombobythere sp. 2: H, LV external view (MP-O-3185); I, same specimen internal view; J, RV external view (MP-O-3184); K, same specimen internal view. Ruggieriella mcmanusiYasuhara et al., 2009 a : L, female LV external view (MP-O-3186); M, male LV external view (MP-O-3187); N, LV external view (MP-O-3188); O, LV external view (MP-O-3189). Scale bars = 0.1 mm.

    Material examined. MP-O-3181 RV (sample 1335C6 0.05 m), L = 0.83 mm, H = 0.47 mm (Figs. 4C and 4E). MP-O-3182 LV (sample 1335C1 0.0 m), L = 0.85 mm, H = 0.5 mm (Fig. 4 D ).

    Occurrence and material. Samples 1335C1 0.0 m (1v) and 1335C6 0.05 m (1v).

    Remarks. Several studies have illustrated specimens of Bythoceratina scaberrima (Brady, 1866) that are similar to Retibythere sp. 2 (e.g., Yasuhara et al., 2014; Gemery et al., 2015; Jöst et al., 2022). According to Benson (1973), no type-series was designated when B. scaberrima was described. However, Brady (1886: 198) states that: “when we examine the outline of the two or three detached valves… that have been used to describe the species” (translated from French), thus designating a syntype series of at least three valves. A re-analysis of the collection should lead to the designation of a lectotype and two paralectotypes or, in the case of loss or destruction of the material, of a neotype. Rare reports of B. scaberrima or its subspecies have illustrations of the carapace in inner view (Benson, 1973; Colalongo and Pasini, 1980), while in others it is not illustrated at all (Guernet, 2005; Sciuto, 2005). Considering the hinge morphology (see genus remarks), differences in the outline, and the absence of a type series for the comparison, the material herein studied is kept in open nomenclature.

    Genus Rhombobythere Schornikov, 1982 b

    Type-species. Rhombobythere intertextaSchornikov, 1982 b , by original designation.

    Remarks. Rhombobythere is an eurybathic genus whose known geographic distribution is currently restricted to the North Atlantic and Australasia. Schornikov (1982 b ) described eight species of this genus in Australia from samples between 8 and 186 m water depth. We consider that Bythocythere sp. identified by Bentley (1988) in Brisbane Water (Australia) is in fact Rhombobythere foveolataSchornikhov, 1982b. Moreover, ?Pseudocythere sp. 2 reported by Didié and Bauch (2001) and Alvarez Zarikian (2009) from the Rockall Plateau and the IODP Site U1314, respectively, is possibly another species of Rhombobythere. In the present study, the two Rhombobythere species occurred only in the shallowest site (1338) located in the Rio Grande Rise (Tab. 1; Fig. 1).

    Rhombobythere sp. 1

    (Fig. 4 F , G)

    Material examined. MP-O-3183 (sample 1338C5 0.01 m) LV, L = 0.80 mm, H = 0.48 mm.

    Occurrence and material. Samples 1338C1 0.02 m (7v), 1338C1 0.08 m (6v), 1338C1 0.10 m (7v), 1338C1 0.16 m (3v), 1338C5 0.01 m (3v), 1338C5 0.03 m (7v), 1338C5 0.09 m (3v), 1338C5 0.04 m (2v), 1338C5 0.05 m (4v), 1338C5 0.10 m (6v), 1338C5 0.11 m (3v), 1338C5 0.12 m (2v), and 1338C5 0.13 m (2v).

    Remarks. Although relatively abundant, most of the specimens of Rhombobythere sp. 1 in the studied material are juveniles. Only one adult LV was found, however it is broken. We chose to keep this species in open nomenclature until more and better-preserved specimens of both valves are found.

    Rhombobythere sp. 2

    (Fig. 4 H -K)

    Material examined. MP-O-3184 (1338C5 0.11 m) RV, L = 0.74 mm, H = 0.35 mm (figs. 4J, K); MP-O-3185 (1338C5 0.11 m) LV, L = 0.71 mm, H = 0.35 mm (figs. 4H, I).

    Occurrence and material. Samples 1338C1 0.02 m (1v), 1338C1 0.10 m (1v), and 1338C5 0.01 m (2v).

    Remarks. Rhombobythere sp. 2 differs from Rhombobythere sulcataSchornikov, 1982 b in the outline of the dorsal margin and the absence of a posteromedian tubercle. It is possibly a new species, but the paucity of specimens precludes its description.

    Tribe Jonesini Schornikov, 1981

    Diagnosis. Carapace with caudal process of different shapes or absent. Dorsal margin straight or with fringe immediately after middle. Surface smooth with meso- or microsculpture of longitudinal pattern. Outer marginal ridge absent. Hinge lophodont, not crenulated, with narrow, long, weak, projected terminal teeth. Anterior or median inner rib frequently well-develop. Concrescence zone wide, with narrow simple or branched pore-canals. Vestibulum wide, not developed only at oral concavity. Inner ridge more developed on left than on right valve. Normal pore canals very thin, simple or with rim. Fulcral point sometimes not pronounced, two frontal scars sometimes fused (adapted from Schornikov, 1981).

    Genus Ruggieriella Colalongo and Pasini, 1980

    Type-species. Ruggieriella decemcostataColalongo and Pasini, 1980, by original designation.

    Remarks. Ruggieriella seems to be restricted to the deep-sea of the Atlantic and Mediterranean (Colalongo and Pasini, 1980; Cronin, 1983; Bergue and Coimbra, 2008; Yasuhara et al., 2009 a ; Yasuhara et al., 2021; Jöst et al., 2022). The present record seems to be the southernmost occurrence of the genus.

    Ruggieriella mcmanusi Yasuhara et al., 2009 a

    (Fig. 4 L -O)

    Rugieriella mcmanusiYasuhara et al., 2009 a : 892, pl. 4, figs. 1-5. - Yasuhara et al., 2021: 9, fig. 5.1.

    Ruggieriella sp. aff. R. mcmanusiYasuhara et al.; 2009 a - Bergue et al., 2023: 11, fig. 3N.

    Holotype. Adult RV, USNM 537137.

    Type-locality and age. ODP Site 1055B, sample 98-100 cm, North Atlantic (32°47.041’N 76°17.179’W). Quaternary.

    Material examined. MP-O-3186 (1335C6 0.02 m) female LV, L = 0.45 mm, H = 0.28 mm (Fig. 4 L ); MP-O-3187 (1345C7 0.0 m) male LV, L = 0.5 mm, H = 0.25 mm (Fig. 4 M ). MP-O-3188 (1344C5 0.0 m) male LV, L = 0.44 mm, H = 0.23 mm (Fig. 4 N ); MP-O-3189, (sample 1344C5 0.0 m) male LV, L = 0.42 mm, H = 0.21 mm (fig. 4O).

    Occurrence and material. Samples 1344C5 0.0 m (3v), 1335C6 0.02 m (1v), and 1345C7 0.0 m (1v).

    Stratigraphic and geographic distribution. Quaternary: ODP Sites 1055 and 925 (Yasuhara et al., 2009 a ; 2021). Holocene: São Paulo Plateau, Southwestern Atlantic (Bergue et al., 2023 and this study).

    Remarks.Bergue et al. (2023) recorded some specimens with a well-marked anterior marginal rib and a curved penultimate lateral rib, which were identified as Ruggieriella sp. aff. R. mcmanusi (fig. 3N in Bergue et al. 2023). The examination of more specimens in this work, however, demonstrated the possibility that the degree of development of these ribs could be more parsimoniously interpreted as a continuous variation and, consequently, an intraspecific variation.

    CONCLUDING REMARKS

    The late Quaternary changes in the oceanic circulation and, consequently, in global climate, have modulated the distribution, abundance, and diversity of deep-sea ostracods during the Late Cenozoic (Yasuhara et al., 2009 b ; Yasuhara and Danovaro, 2016). As these organisms usually display low populational densities (Jöst et al., 2017), their record is more susceptible to the effect of oceanographic changes on calcite preservation (Majoran et al., 1997; Yasuhara et al., 2008, 2016). Therefore, the fossil record of bythocytherids in the study area possibly responds to the influence of centennial to millennial-scale events associated with shifts in the Inter Tropical Convergence Zone (ITCZ) in the southwestern Atlantic Ocean during the Holocene (Gyllenkreutz et al., 2010; Pivel et al., 2010; Chiesi et al., 2014; Lessa et al., 2016).

    Besides the influence of water mass properties on ostracod carapace preservation, the physiography of the study area, which includes plateaus, ridges, and seamounts (Alberoni et al. 2020; Gaurisas and Bernardino, 2023) possibly exerted some influence on the pattern of occurrence of the bythocytherids. However, despite the existence of cold seeps in this region (Freire et al., 2017; Fujikura et al., 2017) their influence on the diversity of the ostracod faunas could not be ascertained (Bergue et al., 2023). Therefore, the available data do not allow concluding whether the taphonomy or autecology exerted more influence on the composition of assemblages (Fig. 2). Notwithstanding, the existence of bathymetric zonation is clearly demonstrated by the absence of bythocytherids in the sites deeper than 3,000 m water depth (i.e., 1336, 1337, 1340, and 1341), despite the occurrence of other ostracod taxa therein (Bergue et al., 2023). A possible explanation for such zonation is the transition between the North Atlantic Deep Water (NADW) and the Antarctic Bottom Water (AABW) near this bathymetric interval (Silveira et al., 2020). The NADW temperature is around 3-4°C while in the AABW it is lower than 2°C (Silveira et al. 2000; Morozov et al., 2020), and previous studies proposed that these differences between the NADW and the AABW influence Ostracoda distribution (Dingle and Lord, 1990; Yasuhara et al., 2009 b ).

    Finally, it is relevant to mention that most of the species herein studied have not been recorded in the RGR before, probably because previous studies developed therein focused mainly on Pliocene and older (Benson, 1977; Benson and Peypouquet, 1983; Bergue et al., 2019). The only possible exception is Retibythere sp. 2, which could be conspecific to B. scaberrima sensu Bergue et al. (2019). Even considering that the general patterns of geographic and bathymetric distributions presented in this paper might be biased by the paucity of data from some regions, they supply additional ecological information on five bythocytherid genera.

    ACKNOWLEDGEMENTS

    The authors express their gratitude to the Brazilian Geological Survey for the samples used in this study and the two anonymous reviewers who contributed to the improvement of this work.

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    ADDITIONAL INFORMATION AND DECLARATIONS

    • Consent for publication
      All authors declare that they have reviewed the content of the manuscript and gave their consent to submit the document.
    • Study association
      This study is a follow up on Bergue et al. (2023).
    • Data availability
      All data generated and analyzed during this study are presented in this article.
    • Funding and grant disclosures
      This study was funded by the National Council for Scientific and Technological Development (CNPq) grant n° 402860/2021-7.
    • Study permits
      Not applicable.

    Edited by

    • Associate Editor:
      Lourdes Elmoor Loureiro
    • Editor-in-chief
      Christopher Tudge

    Data availability

    All data generated and analyzed during this study are presented in this article.

    Data citations

    Brandão, SN; Antonietto, LS; Nery, DG; Pereira, JS; Praxedes, RA; Santos, SG and Karanovic, I. 2023. World Ostracoda Database. Accessed at Accessed at https://www.marinespecies.org/ostracoda on 2023-05-04.

    Publication Dates

    • Publication in this collection
      10 Jan 2025
    • Date of issue
      2025

    History

    • Received
      08 Jan 2024
    • Accepted
      24 June 2024
    • Corrected
      28 Jan 2025
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