Rev. bras. paleontol. 18(3):xxx-xxx, Setembro/Dezembro 2015 © 2015 by the Sociedade Brasileira de Paleontologia doi: 10.4072/rbp.2015.2.01 BIOSTRATIGRAPHIC AND PALEOZOOGEOGRAPHIC REVIEW OF THE UPPER APTIAN-ALBIAN OSTRACODS OF RIACHUELO FORMATION, SERGIPE- ALAGOAS BASIN, NORTHEASTERN BRAZIL LUCAS SILVEIRA ANTONIETTO, DERMEVAL APARECIDO DO CARMO Laboratory of Micropaleontology, Institute of Geosciences, University of Brasilia, Darcy Ribeiro Campus, 70919-970, Brasilia, DF, Brazil. antoniettols@gmail.com MARTA CLAUDIA VIVIERS PETROBRAS-CENPES/DIVEX, Cidade Universitária, Quadra 7, Ilha do Fundão, 21949-900, Rio de Janeiro, RJ, Brazil. mcviviers@petrobras.com.br RODRIGO RODRIGUES ADÔRNO Serviço Geológico do Brasil, DEGEO/DIPALE-CPRM/REPO, Avenida Lauro Sodré, 2561, CEP. 76801-58, Porto Velho, Rondônia, Brazil. rodrigo.adorno@cprm.gov.br ABSTRACT – A biostratigraphic and paleozoogeographic revision of upper Aptian-Albian ostracods of the Riachuelo Formation, in the Sergipe-Alagoas Basin, Brazil, was conducted. The studied material comprised 336 samples from several outcrops, along with well cores drilled by Petróleo Brasileiro S.A. (PETROBRAS). After taxonomic analysis, including the description of Reticulocosta edrianae n. sp., a new biozonation scheme was obtained for the Albian interval of that formation, and the paleozoogeographic affi nities of the Brazil-Central West Africa province were reviewed through comparison with results published in the current literature. Several changes were made to the previously established ostracod zones. A new code for the Aracajuia benderi Zone (OSE-1), MSA-0, was created, with impact in all of its subzones. The names of the Pattersoncypris ex. gr. angulata (OSE-1.1), ‘Patellacythere’ sp. GA E 27 (OSE- 1.3) and Veenia guianensis (OSE-1.4) subzones were changed to Harbinia sinuata? (MSA-0.1), Praebythoceratina amsittenensis (MSA-0.3) and Reticulocosta edrianae (MSA-0.4), respectively. The Sergipella viviersae Subzone (OSE- 1.2) had its code changed for MSA-0.2; also, a stratotype was stablished for it, and its lower limit repositioned in time (from early Albian to upper Aptian). The Metacytheropteon aff. Metacytheropteron sp. GA C 24 Subzone (OSE-1.5) was renamed MSA-0.5. The uppermost Albian Aracajuia antiqua Latest Occurrence Interval Zone (MSA-1) is herein proposed, following the reclassifi cation of specimens of the Subzone Aracajuia benderi (OSE-1.6) from this species to Aracajuia antiqua. Paleozoogeographic analyses showed some faunal interchange with Tethyan provinces, beginning in the early Albian, at both specifi c and generic levels. Limited northward marine infl uxes across the Walvis Ridge were also observed during this age, while larger scale southward migrations occurred in the early Cenomanian. The presence of Brachycytherinae species in the Brazil-Central West Africa province as early as the Aptian indicates a new zoogeographic origin for this subfamily, while explaining the occurrence of the group in both the Austral realm and part of the Tethyan realm during the Albian. Key words: Early Cretaceous, South Proto-Atlantic Ocean, marine ostracods, MSA-1 Zone, province interchange. RESUMO – Uma revisão bioestratigráfi ca e paleozoogeográfi ca dos ostracodes do Aptiano superior-Albiano da Formação Riachuelo, bacia de Sergipe-Alagoas, Brasil, foi proposto no presente trabalho. O material estudado compreendeu 336 amostras de vários afl oramentos, juntamente com testemunhos de poços perfurados pela Petróleo Brasileiro S.A. (PETROBRAS). Após a análise taxonômica, incluindo a descrição de Reticulocosta edrianae sp. nov., um novo zoneamento bioestratigráfi co foi obtido para a formação, e as afi nidades paleozoogeográfi cas da província Brasil-África Central Ocidental (incluindo a bacia de Sergipe-Alagoas) foram revistas por meio da comparação com resultados publicados na literatura existente. Várias mudanças foram feitas nas biozonas previamente estabelecidas com base em ostracodes. O código da Zona Aracajuia benderi (OSE-1) foi alterado para MSA-0. As subzonas Pattersoncypris ex. gr. angulata, (OSE-1.1) ‘Patellacythere’ sp. GA E 27 (OSE-1.3) e Veenia guianensis (OSE-1.4) tiveram seus nomes e espécies guia alterados respectivamente para Harbinia sinuata (MSA-0.1), Praebythoceratina amsittenensis (MSA-0.3) e Reticulocosta edrianae (MSA-0.4), respectivamente. A subzona Sergipella viviersae (OSE-1.2) teve seu código alterado para MAS-0.2, e um novo estratótipo foi proposto para a mesma. A Subzona Metacytheropteron aff. Metacytheropteron sp. GA C 24 (OSE-1.5) foi renomeada como MSA-0.5. A Zona Diferencial Superior Aracajuia antiqua (MSA-1), do Albiano terminal, é aqui proposta, uma vez que espécimes tidos como fósseis guia para a Subzona Aracajuia benderi (OSE-1.6) são aqui identifi cados como pertencendo à espécie Aracajuia antiqua. Análises paleozoogeográfi cas evidenciaram intercâmbio de 1 2 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 faunas da Província Brasil-África Central Ocidental com províncias Tetianas a partir do Albiano, tanto em nível de espécie quanto de gênero. Infl uxos marinhos limitados para o norte através da Cordilheira de Walvis também foram observados durante esta idade, enquanto migração de maior intensidade em direção ao sul ocorreu no Eocenomaniano. A presença de espécies de Brachycytherinae na Província Brasil-Central África Ocidental a partir do Aptiano indica uma nova origem zoogeográfi ca para esta subfamília, ao mesmo tempo em que explica a ocorrência do grupo tanto em regiões austrais quanto tetianas durante o Albiano. Palavras-chave: Eocretáceo, Proto-Oceano Atlântico Sul, ostracodes marinhos, Zona MSA-1, intercâmbio faunístico. INTRODUCTION The Sergipe-Alagoas Basin is a marginal basin located in northeastern Brazil (9-11°30ʼS/37-35°30ʼW). Its onshore The present work is a biostratigraphic and paleozoo- portion is between 16 and 50 km long and 170 km wide, geographic revision of the upper Aptian-Albian ostracods covering a total area of approximately 11,000 km2 (Feijó, of the Riachuelo Formation, in the Sergipe-Alagoas Basin, 1994), bounded by the Pernambuco-Paraíba to the northeast northeastern Brazil. Its main objectives are to upgrade the (Maragogi high) and the Jacuípe basin to the southwest (Itapuã biostratigraphic framework based on ostracods for the Albian fault) (Figure 1). It has been the subject of several studies, of the basin and to re-evaluate the correlations between the mainly due to the amount of data acquired from efforts such as ostracod biozones and those based on planktic and benthic well and outcrop sampling, investigations of seismic lines and foraminifera and cephalopods. At the same time, we also seek gravimetric and magnetic surveys. Most of these studies have to determine the relationships between the “Brazil-Central focused on the oil- and gas-producing strata of the basin, i.e., West Africa” biological province proposed by Tambareau the Carmópolis oil fi eld (Late Jurassic-Aptian) and the Calumbi (1982) with its counterparts worldwide, through comparison Formation (Santonian-Cenozoic) (Marques, 1965; Souza-Lima with previous works addressing the same issue. et al., 2002; Campos Neto et al., 2007; Graddi et al., 2007). A B C Figure 1. Location of the Sergipe-Alagoas Basin in South America (A) and Brazil (B), along with the outcrops and wells studied in the present work (C). ANTONIETTO ET AL. – REVIEW OF UPPER APTIAN-ALBIAN OSTRACODS FROM NE BRAZIL 3 Since the fi rst studies on ostracods from the Sergipe- STUDY AREA Alagoas basin conducted in the 1960s (Krömmelbein, 1962, 1964), the Riachuelo Formation has been at the heart of The Sergipe-Alagoas basin was originally described important discoveries. Viviers et al. (2000) established the by Schaller (1969) as a single basin. However, a later fi rst ostracod-based biostratigraphy of the basin (Figure 2), stratigraphic revision by Feijó (1994) considered two integrating it with previous cephalopod and foraminifer separate basins, the Sergipe basin and the Alagoas Basin, data (Koutsoukos & Bengtson, 1993) and, in this process, separated by the Japoatã-Penedo high. Campos Neto et al. identifying several new species in the aforementioned (2007) recently dismissed the Japoatã-Penedo high as a formation. Do Carmo et al. (2008, 2012), Poropat & Colin divisor between them, as it is not observable throughout the (2012) and Antonietto et al. (2013) presented subsequent entire basin. It is noteworthy that the sedimentary deposits taxonomic updates of some species. and tectonic style in the basin vary in the states of Sergipe Figure 2. Biostratigraphy of the Albian and early Cenomanian sections of the Sergipe-Alagoas Basin, Brazil, based in ostracodes, foraminifers and cephalopods (modified from Koutsoukos & Bengtson, 1993; Viviers et al., 2000). 4 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 and Alagoas, and they are therefore still considered to be 1-US-1; however, these wells were previously illustrated in separate sub-basins. maps by Viviers et al. (2000). The basin consists of 23 formations, which are correlatable Point ESTRE (713561 N, 8816808 E) in the Municipality to the evolutionary stages of the formation and the expansion of Rosário do Catete begins with marly mudstones that are of the South Atlantic Ocean: syneclise, pre-rift, rift, gulf and rich in phytofossils, followed by interbedded carbonate/clay drift (Ponte & Asmus, 1976; Feijó, 1994). In the beginning levels. Bioturbed creamy shales with abundant impregnations of the drift phase, initially restricted marine deposits and fossil-rich siliceous concretions on the top make up the evolved into open sea environments. The early Albian-late next strata. Immediately above it lie creamy-to-gray carbonate Cenomanian Riachuelo Formation was established during this mudstones that are abundant in phytofossils, especially from phase, due to a transgression that allowed the sedimentation angiosperms. In the same municipality, the Fazenda Santa of openly marine sediments. There are three members of Bárbara outcrop occurs (716635 N, 8821887 E), which is this formation: at the basin margins and grabens, deltaic considered to be the type section of the Taquari Member of the fans were formed, and the coarse sediments they carried Riachuelo Formation (Schaller, 1969). The analyzed section constitute the Angico Member. In lower sedimentation areas, comprises reddish calciferous mudstones, with a fossil record the Maruim Member dominates, with carbonate ramps and consisting of gastropod shells (Family Turritellidae Loven, dolomitized oolite/oncolite banks deposited under fl uctuating 1847) as well as possible brachiopods. sea level conditions. The Taquari Member, on the other hand, The Porto dos Barcos 3 location is situated in the corresponds to calcareous mudstone and shale from lagoonal municipality of Riachuelo (699059 N, 8814030 E). From and slope environments. Two third-order sequences are the base to the top, this slightly saprolitic sequence shows observed in the formation, which are characterized by a highly levels of creamy calcareous siltstone, followed by alternating organic clay basal interval and retrogradational stacking and creamy and gray calciferous siltstones. Upward, there is are interpreted as transgressive systems. A third sequence, a layer of gray-to-yellowish mudstones that become very with predominantly calcareous deposits and a progradational fossiliferous near the top. Over this mudstone, creamy pattern, is interpreted as a highstand system (Campos Neto calciferous siltstones are observed, and the outcrop ends with et al., 2007). creamy micaceous mudstones. The fossil record of the outcrop includes echinoderm spines, gastropods and bivalves. Point Regional settings Penha (692726 N, 8812604 W), also located in Riachuelo, The material studied in the present work comprised 147 presents a sequence of yellow, calcareous fi ne sandstones samples collected from outcrops of the Riachuelo Formation, with levels of brachiopod-coquina and sparse phytofossils in the State of Sergipe, Brazil, plus 189 samples from cores at its base. This sequence is followed by gray calciferous drilled by Petróleo Brasileiro S.A. (PETROBRAS) (Figures shales, laminated calcareous siltstones and laminated fi ne 1, 3), totaling 336 samples (Table 1). The UTM quadrant sandstones. Abundant occurrences of bivalves and echinoids reference for coordinates is 24L, in the SAD69 default system. are also recorded in these strata. The majority of the outcrops (ESTRE, Fazenda Santa Bárbara, In the same municipality, the São José 1 outcrop (693824 Porto dos Barcos 3, Penha, São José 1 and P384) is composed N, 8811279 W) presents the most complex lithology among the of alternate strata of the Taquari and Angico members. The sampled sites. Basal siltstones with calcareous impregnations Maruim Member is only present in the upper part of the are superposed with yellow silty fi ne sandstones and calciferous Massapê outcrop. No lithology or GPS coordinates were yellow siltstones with limestone levels. After a non-observable provided by PETROBRAS for wells 1-CPB-1, 7-CP-252 and interval, massive yellow sandstones are recorded, followed Table 1. Sampled depths analyzed by well in the present work. Well Sampled depth (m) 1-CPB-1 198, 222, 345.5, 348, 348.5, 35, 354.5, 355.5, 356.5, 358.5, 359.5, 360.5, 364.5 0, 12, 30, 60, 90, 98.5, 99.5, 100.5, 101.1, 120, 150, 180, 210, 240, 246, 261, 270, 270.05, 276, 291, 300, 306, 321, 330, 336, 351, 360, 366, 381, 390, 396, 399, 399.5, 400.5, 411, 420, 426, 441, 450, 456, 471, 480, 486, 501, 510, 516, 531, 540, 541, 546, 552, 1-US-1 552.5, 553.5, 554.5, 555.5, 561, 570, 576, 591, 600, 606, 621, 630, 636, 651, 660, 666, 678, 679.55, 681, 690, 696, 711, 720, 726, 741, 750, 756, 771, 780, 786, 801, 804, 810, 816, 831, 840, 846, 861, 870, 876, 891, 900, 906, 921, 930, 936, 951, 953, 953.1, 960, 966, 981, 990, 996, 1011, 1020, 1026, 1041, 1050, 1056, 1067, 1067.25, 1071, 1080, 1086, 1101, 1110, 1116, 1131, 1139.2, 1139, 1140, 1146, 1161, 1170, 1200, 1230, 1260, 1290, 1320, 1350, 1380, 1395, 1410, 1425 466, 466.2, 466.7, 467.5, 467.55, 467.9, 468.7, 469.2, 469.75, 469.8, 470.15, 470.6, 470.65, 471.25, 471.8, 472.25, 472.3, 472.6, 473.3, 473.8, 474.45, 474.5, 474.85, 475.3, 475.85, 476.25, 476.3, 476.5, 476.7, 477.1, 477.5, 477.95, 478.45, 478.85, 479.5, 480.3, 7-CP-252 480.55, 480.95, 481.6, 481.8, 482.9, 483.5, 483.55, 484.15, 484.4, 484.75, 485.6, 486.05, 486.45, 487, 488.15, 488.4, 489.15, 489.2, 489.7, 490, 490.1, 490.35, 492.95, 493, 493.45, 493.5, 494.3, 494.75, 495.85, 495.9, 496.55, 496.6, 497.7, 498.3, 498.6, 502.4, 503.35, 503.53, 506.3, 507.15, 508.4, 508.75, 509.3, 511.35, 511.4, 511.9, 512.7, 514, 514.05, 515, 516, 517.45, 518.3, 518.8, 521, 521.4, 523.9, 527.1, 534.8, 535.55, 540.05, 552, 553.4, 558.9, 560.3, 564.8, 567.15, 568.15, 568.95, 581.35 ANTONIETTO ET AL. – REVIEW OF UPPER APTIAN-ALBIAN OSTRACODS FROM NE BRAZIL 5 by limestones with abundant bivalves, cephalopods and RESULTS AND DISCUSSION bioturbations. Towards the top, marl levels with varying amounts of limestone occur, and above them, there are very The revision of the ostracod biostratigraphy of the fossiliferous solid limestones, immediately under a non- Riachuelo Formation’s Albian strata demonstrates the rhythmic intercalation of creamy laminated siltstone marls necessity of the present changes in range and nomenclature and mudstones, rich in turritelid gastropods. The section ends of the zones and subzones established by Viviers et al. (2000). with the deposition of creamy-to-yellowish, medium-to-thinly New information was added to the Aptian-Albian ostracod laminated calcareous sandstones, with many phytofossils. zones, in order to improve application and resolution of the Locality P384 (695625 N, 8814175 E), also in Riachuelo analyzed units. In accordance to the “Código Brasileiro de Formation, comprises, from the base to the top, creamy Nomenclatura Estratigráfi ca” (Murphy & Salvador, 1999), calciferous laminated mudstones with bioturbation levels, a new code name for these units, “MSA” (“Marinho de rich in brachiopods, bivalves and phytoclasts; interbedded Sergipe-Alagoas”), was established in substitution for the yellowish calciferous siltstones and creamy sandstones; and original “OSE”. A new interval zone was also formalized in creamy calciferous laminated mudstones. The whole section the uppermost Albian of the Riachuelo Formation. is abundant in turritelid shells. Point Massapê (700764 N, Some specimens identifi ed as Veenia guianensis by Viviers 881233 L), located at the homonymous village in Riachuelo, et al. (2000) were guide fossils to one of the subzones of the is the only section addressed in the present work identifi ed as Albian of the Riachuelo Formation. However, they belong in part of the Maruim Member. The lithology of this extensive fact to an entirely new species, Reticulocosta edrianae n. sp., outcrop begins with yellow-creamy laminated calciferous which is herein described. A paleozoogeographic revision is and micaceous siltstones. Over these siltstones, fi ne-to-very also performed in this work, expanding on the correlations fi ne yellowish calciferous/micaceous sandstones occur. These with paleozoogeographic provinces in the northern and are followed by (in this order) intercalations of yellow-to- southern regions along the Proto-Atlantic Ocean. creamy siltstones and mudstones; gray greywacke with dark impregnations; and, fi nally, interposed levels of oolitic Systematic paleontology calcarenites and yellow sandstones with smoky quartz grains, The suprafamiliar classifi cation used here follows Liebau sparse ooids and bioturbations. (2005), while the taxonomy of lower ranks was based on Gründel (1974). The morphological terminology is the same MATERIAL AND METHODS from Sylvester-Bradley & Benson (1971), except for the usage of the term “ridge”, in which Kesling (1951) is followed. One hundred forty-seven samples collected during Terms used to describe the curvature of anterior and posterior fi eldwork were listed, prepared and packaged according ends – infra-curvate, equi-curvate and supra-curvate – were to the methodology developed by the Laboratory of adopted from Lüttig (1962). Type materials were housed Micropaleontology of the University of Brasilia (LabMicro- at the research collection of the Museum of Geosciences, UnB), Brazil. Sixty grams of each sample was prepared, University of Brasilia (UnB), under the prefi x CP (“Coleção and the remainder was packed in double plastic bags with de Pesquisa”). catalogue-record sheets. The sample contents separated for preparation were treated as follows: fi rst, they were treated Subclass OSTRACODA Latreille, 1802 with a solution containing 30% hydrogen peroxide (H2O2), Superorder PODOCOPOMORPHA Kozur, 1972 with the aim of fragmenting rock particles. Ethanol at 92 GL Order PODOCOPIDA Sars, 1866 was added to interrupt this reaction, after which separation of Suborder CYTHEROCOPINA Gründel, 1967 grains by size was performed, using mesh sieves of 630, 250, Infraorder NOMOCYTHERININA Liebau, 1991 160 and 80 μm. For samples MP-1430 to MP-1434, MP-1493, Superfamily TRACHYLEBERIDOIDEA Sylvester- MP-1494 and MP-1678, the 80 μm sieve was replaced by a Bradley, 1948 90 μm-56 μm dual set. Family CYTHERETTIDAE Triebel, 1952 The carapaces and valves of several species were studied Subfamily PALAEOCYTHERIDEINAE Ljubimova, 1955 at LabMicro-UnB and the Smithsonian National Museum of Tribe CYTHERETTINI Triebel, 1952 Natural History (NMNH), Washington D.C., United States. The analysis of the additional 189 previously processed Reticulocosta Gründel, 1974 samples from PETROBRAS was conducted at the “Centro de Pesquisas Leopoldo Américo Miguez de Mello” (CENPES), Type species. Reticulocosta ornatoreticulata Reyment, 1963, Rio de Janeiro, Brazil. After taxonomic analysis of the by subsequent designation of Gruendel (1974). ostracods present in the samples, a new biozonation scheme for the Albian of the Sergipe-Alagoas Basin was obtained. The Reticulocosta edrianae n. sp. ostracods were also used to revise the paleozoogeographic (Figures 4A-L) affi nities of the Sergipe-Alagoas basin with other basins during the Albian-early Cenomanian through comparison 2000 Veenia guianensis Swain. Viviers, Koutsoukos, Silva- with previous results from the current literature on this topic. Telles Jr. & Bengtson, p. 413, fi g. 22.10. 6 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 2000 Veenia sp. P1. Viviers, Koutsoukos, Silva-Telles Jr. & view, Paramphidont hinge, composed of an elliptical, Bengtson, p. 410, fi gs. 23.3, 23.4. four-lobed tooth in the right valve, followed by an anterior round socket, a smooth bar and a posterior, also four-lobed Etymology. Latinized form of “Edriana”, first name of tooth; elements in the left valve are complementary. Well- Edriana Araújo de Lima. The name of this species is a tribute developed inner lamela, not visible in the oral region, where to her. the valve margin bends inwards expressively. Selvage is not Holotype. Female carapace (CP-716). visible at the postero-ventral and postero-central regions. Alotype. Male carapace (CP-717). Central muscle scars group consisting of a slightly concave Paratypes. Female carapace (CP-718), female carapace set of four adductor scars, the second from dorsum to venter (CP-719), female left valve (CP-720) and male right valve subdivided in two, and two elliptical frontal scars. Strong (CP-768). sexual dimorphism: females are shorter and more oblong Type locality and stage. Riachuelo Formation, Sergipe- in dorsal view than males; thicker ridges and less reticulate Alagoas Basin, Riachuelo Municipality, Sergipe State, Brazil, ornamentation in females than males. Interspecifi c variation approximate coordinates: 699059N, 8814030L (UTM 24L observed as follows: degree of reticulation in the carapace quadrant, SAD69 geodesic reference system), Porto dos lateral surface varying from widespread to concentrate along Barcos 3 outcrop, Sample MP-1461. middle Albian, Aracajuia its central region, and height/length ratios of the specimens benderi Zone (MSA-0), Praebythoceratina amsittenensis resulting in sturdier or more elongate forms. Subzone (MSA-0.3). Dimensions. CP-716 (holotype), length 0.54 mm, height Diagnosis. A species of Reticulocosta distinguished by the 0.33 mm, width 0.27 mm; CP-717 (alotype), length 0.57 mm, following features: carapace ornamentation consisting of height 0.31 mm, width 0.26 mm; CP-718 (paratype), length three highly acuminate ridges, quite varied in thickness, at 0.52 mm, 0.32 mm, width 0.29 mm; CP-719 (paratype), length dorsal, central and ventral position. Secondary ornamentation 0.52 mm, height 0.31mm, width 0.28 mm; CP-720 (paratype), includes rounded to foveolate reticulation in several degrees length 0.55 mm, height 0.33 mm; CP-768 (paratype), length of extension, but always present at the centro-anterior region. 0.56 mm, height 0.28 mm. Description. In lateral view, subtriangular to sub-rectangular Occurrence. ESTRE outcrop, samples MP-1423 and MP- carapace, with greatest height at the antero-central and 1424; Fazenda Santa Bárbara outcrop, samples MP-1419 greatest length at the central regions. Left valve is larger and MP-1420; P384 outcrop, samples MP-1496, MP-1497, than the right, overlapping it through the entire free margin, MP-1499, MP-1506 and MP-1509 to MP-1513; Porto dos but more intensely at the antero-dorsal, antero-ventral and Barcos 3 outcrop, samples MP-1460 to MP-1466, MP-1468 postero-dorsal margins. Round anterior end, becoming to MP-1472, MP-1474 to MP-1477, MP-1481 and MP-1482. slightly rectilinear close to the dorsal margin, where an Remarks. Reticulocosta edrianae n. sp. is similar to obtuse cardinal angle is formed. Round, supra-curvate Reticulocosta tarfayensis (Reyment, 1978), even presenting posterior end, becoming slightly rectilinear close to the some degree of intraspecific variation on the reticulate dorsal margin, where they form an obtuse cardinal angle; ornamentation, like the latter (see Reyment, 1978; Andreu- a small caudal process is present. Rectilinear to slightly Boussut, 1991; Majoran, 1991). Even so, the aforementioned concave dorsal margin. Rounded ventral margin, with a mild species, plus Reticulocosta vitiliginosa (Apostolescu, 1961), concavity at the oral region. Ornamentation consisting of differ from the present species by its overall lesser presence three highly acuminate ridges, quite varied in thickness, at of reticulation and sharper ridges, when compared to them dorsal, central and ventral position: the generally smooth, (see Apostolescu, 1961). In addition, R. edrianae n. sp. has its curved dorsal one runs from the postero-dorsal to the antero- occurrence restricted to the Lower Cretaceous (Albian), while dorsal region, projecting behind the dorsal margin; the lightly the others are Upper Cretaceous species (upper Cenomanian- posteriorly nodulate and undulate central one runs from the Santonian). postero-central to the central region, where it connects to the reticulate centro-anterior tubercle; and the lightly posteriorly Biostratigraphy nodulate and curved ventral one runs from the postero- According to Viviers et al. (2000), the Albian Stage in ventral to the antero-ventral region. Two riblets also occur, the Sergipe-Alagoas basin originally comprised one zone, one originating from the eye tubercle and running along the Amphycytherura benderi, or OSE-1, later renamed Aracajuia anterior end, right below it, and the other, irregularly shaped, benderi by Antonietto et al. (2013). This zone was subdivided runs along the ventro-central area, partially covering the into six subzones from bottom to top: Pattersoncypris ex. ventral margin. Additionally, a series of teeth radiates from gr. angulate, OSE-1.1; Sergipella transatlantica, OSE-1.2, the anterior and posterior ends. Secondary ornamentation renamed as Sergipella viviersae Do Carmo et al., 2012 by Do includes rounded to foveolate reticulation in several degrees Carmo et al. (2012); ‘Patellacythere’ sp. GA E 27, OSE-1.3; of extension, but always present at the centro-anterior Veenia guianensis Swain, 1976, OSE-1.4; Metacytheropteron region; Well-developed rounded eye tubercle. In dorsal aff. M. sp. GA C 24, OSE-1.5; and Amphycytherura benderi, view, elliptical, with greatest width at the centro-posterior OSE-1.6 – also Aracajuia benderi, after Antonietto et al. (2013). region; a fl at surface is formed between the anterior and Between subzones OSE-1.3 and OSE-1.4, there was an interval posterior cardinal angles and the dorsal ridges. In internal where no zonation based on ostracods could be established, ANTONIETTO ET AL. – REVIEW OF UPPER APTIAN-ALBIAN OSTRACODS FROM NE BRAZIL 7 although a transition is observable in the planktic (ending of Sergipella viviersae Latest Occurrence Interval Subzone Ticinella bejaouensis Zone, Ticinella bejaouensis Subzone) and (MSA-0.2) benthic (ending of Epistomina carpenteri Zone, Osangularia Defi nition. Lower and upper boundaries of the subzone schloenbachi-Pseudogaudryinella/Spiroplectinata ex gr. defi ned by the last occurrences of Harbinia sinuata? and dividens Subzone) foraminiferal faunas as well as between Sergipella viviersae, respectively. cephalopods (ending of Douvilleiceras Zone ) (Figure 2). Characterization. Other species observed in this At the present revision of the upper Aptian-Albian zone include Cytherella besrinensis (Bischoff, 1964), stratigraphy of the basin, several changes were made in Cytherelloidea btaterensis Bischoff, 1964, Bairdoppilata previously established ostracod zones and subzones, based on pseudoseptentrionalis Mertens, 1956, Cetacella sp. 1, the correlation of outcrops and wells analyzed (Figure 5). The Paracypris eniotmetos Nicolaidis & Piovesan, 2013, OSE-1 Zone and its subzones, OSE-1.1, OSE-1.2, OSE-1.3 and Harbinia crepata Do Carmo et al., 2013, Liasina sp. 1, OSE-1.4, are renamed and redescribed below, while OSE-1.5 Praebythoceratina amsittenensis, Praebythoceratina sp. 1, had only its name changed (MSA-0.5). An entirely new zone, Praebythoceratina reducta (Gründel, 1964), Patellacythere MSA-1, is created in the uppermost Albian of the Riachuelo shimonensis (Rosenfeld & Raab, 1983), Xestoleberis? sp. Formation, susbstituting the form OSE-1.6 Subzone. 1, Xestoleberis? sp. 2, Dico, Aracajuia antiqua (Rosenfeld & Raab, 1983), Aracajuia benderi, Eocytheropteron sp. 1, Aracajuia benderi Range Zone (MSA-0) Metacytheropteron aff. M. sp. GA C 24 and Metacytheropteron Defi nition. Stratigraphic interval corresponding to the local minuta? (Swain, 1976). range of Aracajuia benderi Krömmelbein, 1967. Stratotype. 1-CPB-1 well, between depths of 348 and 354.5 Characterization. From the base to the top, the following m, Municipality of Carmópolis, Sergipe State, Brazil. subzones are recognized: the Harbinia sinuata? (Krömmelbein Stage. Uppermost Aptian-lowermost Albian. & Weber, 1971) (MSA-0.1), Sergipella viviersae (MSA- Observations. A detailed discussion on the MSA-0.2 0.2), Praebythoceratina amsittenensis Andreu-Boussut, Subzone (formerly OSE-1.2) can be found in Viviers et al. 1991 (MSA-0.3), Reticulocosta edrianae (MSA-0.4) and (2000). The present work formalizes the taxonomic changes Metacytheropteron aff. M. sp. GA C 24 (MSA-0.5). made to this subzone by Do Carmo et al. (2012), while Stage. UpperAptian-upper Albian. proposing a stratotype for it. Originally, this subzone was Stratotype. 1-US-1 well, between depths of 690 and 1161m, restricted to the Taquari Member; however, present data Municipality of Laranjeiras, Sergipe State, Brazil. show it also occurs in the Angico Member. The occurrence Observations. A detailed discussion on the Zone MSA-0 of Harbinia crepata, a characteristic species of the Aptian (formerly OSE-1) can be found in Viviers et al. (2000). A of Brazil (Do Carmo et al., 2013), at the lowest portion of stratotype, established in core sections of the 1-US-1 well, MSA-0.2, associated with other Aptian species (Cytherella is also proposed for MSA-0 for the fi rst time. Although the besrineensis, Cytherelloidea btaterensis, Patellacythere zone is not entirely represented in this well (it lacks MSA-0.1, shimonensis and Aracajuia benderi) led to repositioning MSA-0.3 and MSA-0.4 subzones), this is is the most complete the lower limit of this subzone from the lower Albian to the section of MSA-0 so far analyzed. uppermost Aptian. Conchoecia? sp. Se1 was not observed Harbinia sinuata? Range Subzone (MSA-0.1) in the present work. Defi nition. Stratigraphic interval corresponding to the local range of Harbinia sinuata?. Praebythoceratina amsittenensis Latest Occurrence Characterization. Interval containing a characteristic Interval Subzone (MSA-0.3) association of Harbinia sinuata?, Aracajuia benderi, Defi nition. Lower and upper boundaries of the subzone Sergipella viviersae and Dicrorygma? sp. 1. defi ned by the last occurrences of Sergipella viviersae and Stratotype. 7-CP-252 well, between depths of 478.45 and Praebythoceratina amsittenensis, respectively. 484.4m, Municipality of Santo Amaro das Brotas, Sergipe Characterization. The ostracod assemblage of the subzone State, Brazil. also comprises Cytherella besrinensis, Cytherelloidea Stage. Upper Aptian. btaterensis, Bairdoppilata pseudoseptentrionalis, Robsoniella Observations. A detailed discussion on the MSA-0.1 Subzone falklandensis Dingle, 1984, Paracypris eniotmetos, (formerly OSE-1.1) can be found in Viviers et al. (2000). Praebythoceratina amsittenensis, Patellacythere parva? This subzone was established using Pattersoncypris ex. gr. Weaver, 1982, Xestoleberis? sp. 2, Apatocythere? sp. 1, angulata (Krömmelbein & Weber, 1971) as its fossil guide. Dicrorygma? sp. 1, Neocythere tenuis, Aracajuia benderi, However, Harbinia sinuata? (Krömmelbein & Weber, 1971) is Microceratina azazoulensis Andreu & Colin, 2005, more abundant in the studied samples (including the reviewed Microceratina? sp. 1, Reticulocosta edrianae, Algeriana? well material), which justifi es its preference as a marker for sp. 1 and Quasihermanites? sp. 1. Among species from the interval. A stratotype is also proposed for the subzone for Viviers et al. (2000) not observed in the MSA-0.3 Zone in the the fi rst time. Additionally, several species from Viviers et al. present revision are Cytherella sp. Se4, Liasina sp. 1, Veenia (2000), such as Ovocytheridea? aff. Ovocytheridea? sp. GA guianensis (Swain, 1976), Sergipella aff. S. transatlantica D 6, Pattersoncypris ex. gr. angulata, Conchoecia? sp. Se1 Krömmelbein, 1967, Cetacella aff. C. sp. GA D 24 and were not observed in the present study. Metacytheropteron aff. M. sp. GA C 24. 8 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 Figure 3. Lithology of sampled Riachuelo Formation outcrops in the Sergipe-Alagoas basin, Brazil. The samples are numbered from MP-1421 to MP-1698. Abbreviations: MP, Micropaleontological collection of the “Museu de Geociências” of the “Universidade de Brasilia”. Stratotype. 1-CPB-1 well, between depths of 345.5 and 348 m, Characterization. This subzone fauna also includes Municipality of Carmópolis, Sergipe State, Brazil. Cytherella icknieldensis?, Paracypris eniotmetos, Neocythere Stage. Lower-middle Albian. tenuis, Aracajuia benderi and Aracajuia fragilis (Piovesan Observations. The species which originally named the & Nicolaidis, 2013). MSA-0.3 (formerly OSE-1.3) subzone, ‘Patellacythere’ Stratotype. Porto dos Barcos 3 outcrop (699059 N, 8814030 sp. GA E 27, was found to actually belong to a group E, 24L UTM quadrant reference, SAD69 default system), of species in the Family Bythocytheridae Sars, 1926. between 6.7 and 7m high from the outcrop base, Municipality The figured specimen in Viviers et al. (2000) was of Riachuelo, Sergipe State, Brazil. reclassified as Patellacythere shimonensis, a species Stage. Middle Albian. restricted to MSA-0.2 subzone, and most of the material Observations. The specimens identifi ed by Viviers et al. (2000) assigned to this species in fact belonged to another taxon, as Veenia guianensis, the fossil guide for MSA-0.4 (formerly Praebythoceratina amsittenensis. This was particularly OSE-1.4), belong to Reticulocosta edrianae leading to the true for specimens occurring after the end of MSA-0.2. renaming of this subzone. Nevertheless, Veenia guianensis Therefore, Praebythoceratina amsittenensis is considered was also found in the present material, though not associated to be the true MSA-0.3 marker. A detailed discussion on this with R. edrianae Between subzones MSA-0.3 and MSA-0.4, subzone can be found in Viviers et al. (2000). The MSA-0.3 Viviers et al. (2000) observed an absence of any clear biozone subzone was originally restricted to the Angico Member of marking species, and the interval between these subzones was the Riachuelo Formation; its occurrence is herein expanded left open for biozonation. The present work established the to the Taquari Member. Species not observed in the present contact between the two zones, although it was not possible work include Cytherella sp. Se4, Liasina sp. 1, Veenia to determine when in time it occurred. Therefore, the authors guianensis, Sergipella aff. S. transatlantica, Cetacella aff. chose to maintain along the top limits of MSA-0.3, Ticinella C. sp. G D 24 and Metacytheropteron aff. M. sp. GA C 24. bejaouaensis Zone (Al2) of planktic foraminifera and Douvilleiceras Zone of ammonites. A detailed discussion on Reticulocosta edrianae Latest Occurrence Interval this subzone can be found in Viviers et al. (2000). Originally, Subzone (MSA-0.4) there was no information about which members of the Riachuelo Defi nition. Lower and upper boundaries of the subzone Formation MSA-0.4 was present at; therefore, according to defined by the last occurrences of Praebythoceratina our results, it is restricted to the Angico and Taquari members. amsittenensis and Reticulocosta edrianae, respectively. Ovocytheridea? sp. Se1 was not observed in the present study. ANTONIETTO ET AL. – REVIEW OF UPPER APTIAN-ALBIAN OSTRACODS FROM NE BRAZIL 9 Aracajuia antiqua Latest Occurrence Interval Zone 0 Zone (Viviers et al., 2000), were reclassifi ed as Aracajuia (MSA-1) antiqua, creating an entirely new zone in the latest Albian Defi nition. Lower boundary of the zone defi ned by the last of the Riachuelo Formation, the Aracajuia antiqua Latest occurrences of Aracajuia benderi and Metacytheropteron aff. Occurrence Interval Zone, after its guiding species, and coded M. sp. GA C 24, while the upper boundary is defi ned by the as MSA-1. Although its lower limit is established based in last occurrence of Aracajuia antiqua. two species, Aracajuia benderi and Metacytheropteron aff. Characterization. The MSA-1 Zone spans areas nearby M. sp. GA C 24, these two disappear from the fossil record the Massapê outcrop and the 1-US-1 well, between the within a slightly different interval between them. Viviers et towns of Riachuelo and Laranjeiras, Sergipe State, Brazil. al. (2000) did not discuss the distribution of this zone along It is observable in Massapê in all members of the Riachuelo the Riachuelo Formation; according to present results, it is Formation. Its thickness varies between 11 (Massapê observable at all members of that formation. outcrop) and almost 300m (1-US-1 well). Other taxa present in this zone include Cytherella besrinensis, Cytherelloidea Paleozoogeography btaterensis, Bairdoppilata comanchensis (Alexander, 1929), Original attempts to establish relationships based Bairdoppilata sp. 1, Bairdoppilata sp. 2, Cetacella sp. 1, on the composition of taxa between the upper Aptian- Paracypris eniotmetos, Xestoleberis? sp. 1, Dicrorygma? Albian ostracodefaunas of the Riachuelo Formation and sp. 1, Neocythere pseudovanveeni (Gründel, 1966), coeval lithologic units from western Africa indicated Eocytheropteron? sp. 1, Microceratina azazoulensis, Veenia great similarity between the latter units and the Madiéla guianensis and Brachycythere sp. 1. Formation in the Gabon basin, Gabon (Krömmelbein, Stratotype. 1-US-1 well, between depths of 480 and 681 m, 1966, 1975; Krömmelbein & Wenger, 1966; Bertels, Municipality of Laranjeiras, Sergipe State, Brazil. 1977; Grosdidier, 1979; Koutsoukos & Silva-Telles Jr., Stage. Uppermost Albian. 1993). Tambareau (1982) even suggested the existence of Observations. Specimens previously considered to be a “Central Brazil-West Africa” biological province. This Aracajuia benderi in the OSE-1.6 Subzone, at the top of MSA- correlation was later expanded by Koutsoukos & Dias- A G C D H B I F J E K L Figure 4. Reticulocosta edrianae n. sp., Riachuelo Formation, Sergipe-Alagoas Basin, Middle Albian, Brazil. A, CP-716, holotype, female carapace, right lateral view. B, CP-718, paratype, female carapace, left lateral view. C, CP-719, paratype, female carapace, dorsal view. D, E, CP-717, alotype, male carapace; D, dorsal view; E, right lateral view. F-J, CP-720, paratype, female left valve; F, internal view; G, detail of the hinge line; H, interpretation of the hinge line structures; I, detail of the central and anterior muscle scars; J, interpretation of the central and anterior muscle scars. K, L, CP-768, paratype, male right valve; K, dorsal view; L, interpretation of the hinge line structures. Scale bars = 100 μm. 10 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 Britto (1987) and Piovesan et al. (2013), who observed Switzerland (Charollais et al., 1977), the Persian Gulf and the co-occurrence of ostracod species and genera of the Israel (Rosenfeld & Raab, 1983). This observation was partly Riachuelo Formation in other Brazilian shore units, such confi rmed by Piovesan et al. (2013), following a review by as the northeasternmost Potiguar and the southeastern Andreu (2002), who identifi ed some genera distributed along Campos and Santos basins. the Brazilian coast (including the Sergipe-Alagoas Basin) and Dias-Brito (2000), based on the occurrence of pithonellids lithologic units in Morocco. (calcispheres), established the notion of the “Megatethys”, Reyment (1980) and Dingle (1999) studied the role of or the “Mid Cretaceous Tethyan Realm”. According to this the proto-Walvis ridge as a geographical barrier between the concept, the global distribution of pithonellids defi ned a central and southernmost Atlantic ostracodefaunas during realm occupying both hemispheres, approximately between the Berriasian-Santonian. After a taxonomic study of several latitudes 40ºN and 40ºS, in the Albian-Turonian time interval. Cretaceous stages of the Sergipe-Alagoas Basin, Viviers et This Megatethys Ocean reached its maximum extension in al. (2000) summarized the relationships of the Riachuelo the late Albian, when warm water reached temperate regions Formation with strata in several Early-Late Cretaceous and initially tropical organisms made brief incursions into realms along the Proto-Atlantic Ocean. These correlations high-latitude areas. The southern limit of this Tethyan arm was were made based in part on the chronologic and geographic coincident with the Walvis Ridge, an effective topographic distributions of the ostracod genera Conchoecia Dana, 1849, barrier at that time. Amphicytherura Butler & Jones, 1957 and Veenia Butler The first evidence of faunal similarity between the & Jones, 1957. A signifi cant faunal renewal was recorded Riachuelo Formation and strata outside of the central South approximately in the middle Cenomanian, with the fi rst Atlantic Ocean was provided by Colin & Andreu (1990), who appearance in the basin of genera such as Brachycythere were working with halocypridid ostracods. These authors Alexander, 1933 and Paracypris Sars, 1866. found the formations in eastern Brazil and western Africa Present results corroborate previous assumptions to be strongly correlated with those from the North Atlantic herein summarized about the paleozoogeography of the Ocean, from areas such as the Agadir basin in Morocco, Riachuelo Formation ostracods, both at specifi c and generic southern England (Kaye, 1965), the Jura Mountains in levels. Although the majority of the species identifi ed are Figure 5. Reviewed ostracod biostratigraphy of the Sergipe-Alagoas basin, along with the temporal distribution of its late Aptian-Albian ostracod species. Names of species in bold indicate fossil guides to each biozone. ANTONIETTO ET AL. – REVIEW OF UPPER APTIAN-ALBIAN OSTRACODS FROM NE BRAZIL 11 endemic to this province, some interchange is observed through their origins, greater paleozoogeographical with Tethyan faunas beginning in the early Albian (Figure proximity with the Tethysyan realm, such as Dicrorygma 6). Species such as Veenia guianensis and Bairdoppilata Poag, 1962 (see also Christensen, 1965), Neocythere comanchensis are shared with North-Central American Mertens, 1956, Veenia Butler & Jones, 1957, Reticulocosta shores (Veenia guianensis is also found in Africa), while Gründel, 1974, Robsoniella Kuznetsova, 1956 (see also Conchoecia? sp. 1, Cytherella besrinensis, Cytherelloidea Gramm & Kuznetsova, 1970) and Microceratina Swanson, btaterensis, Praebythoceratina amsittenensis, Patellacythere 1980 (see also Colin et al., 2005). Sergipella Krömmelbein, shimonensis and Aracajuia antiqua appear in both 1967 is the only endemic genus of the region, according Brazil-Central West Africa and the Levantine region in to the occurrences summarized in Do Carmo et al. (2012). the Middle East (Israel and Lebanon). Bairdoppilata The genus Aracajuia Krömmelbein, 1967 originated in pseudoseptentrionalis, Praebythoceratina reducta, the Austral province during the Berriasian, prior to becoming Neocythere pseudovanveeni and Neocythere tenuis co-occur very common in this realm during the Hauterivian. However, in the Riachuelo Formation and some of the southern-to- this genus showed its peak diversity and abundance at the western European basins in England, France and Germany. end of the Early Cretaceous (Barremian-Albian), along Two species shared with realms south of the Walvis Ridge, the shores of Brazil-Central West Africa and the South Microceratina azazoulensis and Robsoniella falklandensis, Mediterranean, extending to the Middle East (Antonietto et probably originated in Brazil-Central West Africa, where al., 2013). According to Dingle (1999), limited northward they surged in the middle Albian. These species reached the marine infl uxes across the Walvis Ridge are postulated for the former regions during the Cenomanian. middle Aptian age, while large-scale southward migrations The exchange of genera between the provinces of marine ostracods from the northern sector occurred in appears to follow the same pattern observed at the early Cenomanian and/or Turonian times. This observation specifi c level. Genera such as Cytherella Jones, 1849, is confi rmed by the occurrences of Aracajuia in the early-late Cytherelloidea Alexander, 1929, Bairdoppilata Coryell Albian of the Sergipe-Alagoas Basin. et al., 1935, Paracypris, Bythoceratina Hornibrook, Brachycythere appeared to have originated during 1952 and Patellacythere Gründel & Kozur, 1971 are both the Albian, somewhere close to the limits of the North geographically and chronologically cosmopolitan. However, American and African regions of the Tethysian realm (Colin some other genera present a more limited distribution, at & Babinot, 1996; Puckett, 2002), although occurrences in least during the Early Cretaceous, and seem to indicate, India (Austral realm) have also been reported for the same Figure 6. Paleozoogeography of species with occurrences in the Riachuelo Formation, Sergipe-Alagoas Basin, and coeval lithologies worldwide during the Albian and early Cenomanian. Paleomaps were adapted from ODSN (2011). 12 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 age (Andreu et al., 2008). The presence of Brachycythere (Project no. 401.794/2010-5). We are also grateful to UnB sp. 1 in the Brazil-Central West Africa province dates from for fi nancial support to the visit to PETROBRAS. Thanks the late Aptian (Grosdidier, 1979), much earlier than the to CENPES for technical support with core sample slides previous earliest record for this region, i.e., Brachycythere (J.V. Queiroz Neto and J.L.Grillo) and SEM photography sapucariensis Krömmelbein, 1964 from the Cenomanian of (R.S.M. Costa). Special thanks go to the “Agência Nacional several Brazilian and African basins (Viviers et al., 2000). do Petróleo, Gás Natural e Biocombustíveis”, PETROBRAS Therefore, it relocates the geographic origin of the Subfamily and the “Fundação de Empreendimentos Científicos e Brachycytherinae Puri, 1954 southwards, as it helps to explain Tecnológicos” (FINATEC) for fi nancial support to the fi eld the occurrence of the group in both the Austral realm and part work in the Sergipe-Alagoas basin, through the OSTRAKi of the Tethyan realm during the Albian. project (PETROBRAS/GEO-2007/03676). The fi rst author L.S. Antonietto wishes to thank the Coordination for the CONCLUSIONS Improvement of Higher Education Personnel (CAPES) for granting support that allowed the realization of most of the The present revision of the ostracod biostratigraphy and work in the Smithsonian NMNH from July 2013 to July paleozoogeography of the upper Aptian and Albian stages of 2014 (Process no. 3604-13-6), and the board of the Institute the Riachuelo Formation resulted in the description of a new of Geosciences-UnB, for all cooperation throughout the species, Reticulocosta edrianae. There were also changes in process. Last, but not least, the fi rst author thanks the NMNH one previously established zone and its fi ve subzones, along Paleobiology Department staff members, G. Hunt and C.E. with the proposition of an entirely new zone, MSA-1, created Sanford, for all of their help and great joyful moments during in detriment of OSE-1.6 and based on Aracajuia antiqua. The his stay at Washington, D.C., United States. 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