Cretaceous palaeogeography of southern Gondwana from the distribution of the marine ostracod Majungaella Grekoff: New data and review

Cretaceous Research 37 (2012) 127e147

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Cretaceous palaeogeography of southern Gondwana from the distribution of the marine ostracod Majungaella Grekoff: New data and review Enelise Katia Piovesan a, *, Sara Ballent b,1, Gerson Fauth a a b

Laboratório de Micropaleontologia, Universidade do Vale do Rio dos Sinos-UNISINOS, Avenida Unisinos, 950, CEP 93022-000, São Leopoldo-RS, Brazil Departamento Paleontología Invertebrados, Museo de Ciencias Naturales de La Plata, Paseo del Bosque s/n, La Plata 1900, Argentina

a r t i c l e i n f o

a b s t r a c t

Article history: Received 16 November 2011 Accepted in revised form 14 March 2012 Available online 6 April 2012

Majungaella is a survivor genus of a long lineage of the family Progonocytheridae, ranging from the Jurassic to the Pliocene. In this paper new species from Argentina and Brazil are described and appropriately illustrated, comprising a systematic review of 16 species, including two new ones. The Jurassic and Early Cretaceous species seem to have inhabited mainly shallow, warm waters, whereas from the mid-Late Cretaceous and especially in the Cenozoic, a retrothermal propensity is observed in the genus. The records of Majungaella are revised and its distribution is used to demonstrate the prior juxtaposition of subsequently separated Gondwana continental blocks and the opening of new seaways. The data have allowed a deeper understanding of the depositional sequences and supplied information on the early geological history and subsequent palaeoceanographic evolution of the South Atlantic Ocean. The affinities of the ostracod faunas from South America and other gondwanan localities are used to analyze the evolution of seaways and oceanic barriers. Ó 2012 Elsevier Ltd. All rights reserved.

Keywords: Majungaella Ostracoda Systematics Palaeozoogeography Cretaceous Argentina Brazil

1. Introduction The genus Majungaella was described by Grekoff (1963) from the Upper Jurassic and Lower Cretaceous of Madagascar. Whatley and Ballent (1996) and Ballent et al. (1998) reviewed its taxonomic status, with special reference to its distribution in Argentina, where it is confined to Patagonia and ranges from Upper Jurassice Berriasian to Maastrichtian. Majungaella possesses the biological and palaeontological virtues of benthonic ostracods (i.e., long fossil history, abundance, diversity, control on their spatial distribution by depth, temperature, substrate, among others, and lack of pelagic larvae), which render them valuable tools in palaeogeographic reconstruction. The genus had a northeastern gondwanan distribution throughout the Jurassic, and by the early Neocomian, it had migrated to South America. The continuing southward migration resulted in its eventual arrival in Australia in the Early Cretaceous. It was present on the Antarctic continental shelf during the Late Cretaceous and also on the Antarctic Peninsula in the Eocene, Oligocene and late Pliocene (see Whatley et al., 2005 for a summary of references and a complete list of

* Corresponding author. Tel.: þ55 51 3037 1000 E-mail address: [email protected] (E.K. Piovesan). 1 Deceased 0195-6671/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2012.03.013

the known species and all known citations of valid species of the genus at the time). Cenozoic records from Antarctica demonstrate the adaptation of the genus to living in high latitudes with water temperatures very much lower than those of its Mesozoic forebears. This adaptation is advanced as the prime reason why the genus, alone among the progonocytherids, was able to survive both the post-Cretaceous global cooling and Antarctic palaeoenvironments into the late Neogene (Whatley et al., 2005). Considering that Majungaella is restricted to the Southern Hemisphere (or areas such as India which subsequently migrated north to the Equator), it has become a classical element on which to establish affinities among gondwanan localities during the Mesozoic. The works of Sigal et al. (1970) and Krömmelbein (1976, 1979), were pioneering in using species of Majungaella to indicate “gondwanide affinities” among South America, South Africa, Madagascar, India and Australia. The subject has also been treated several times by Dingle (1982, 1988). More recent antecedents are, among others, the publications of Babinot and Colin (1992), Seeling et al. (2004), Ballent (2009), Dingle (2009) and Guzel (in press). There are some references on the occurrence of Majungaella along the Brazilian continental margin (Krömmelbein, 1979; Miller et al., 2002; Ceolin et al., 2011). However, there have been no studies published on its taxonomy or its stratigraphical distribution hitherto.

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The purpose of this contribution is to review the present state of knowledge of Majungaella in Argentina and Brazil, to describe and illustrate appropriately new species from the Cretaceous Brazilian continental margin and to discuss the value of the genus and species in correlation within localities along the southern margins of Gondwana. It comprises a systematic review of 16 species, including the description of two that are new. This study complements others by the authors concerning the distribution of Mesozoic gondwanide ostracods (Ballent et al., 1998; Ballent and Whatley, 2000, 2006, 2007; Whatley et al., 2005; Piovesan et al., 2009, 2010). 2. Geological setting Fig. 1 shows the location of the basins in Argentinian Patagonia and along the Atlantic Brazilian margin, from which the described species have been recovered.

relatively undeformed (Howell et al., 2005). This is in contrast to the Andean region where Late CretaceouseCenozoic deformation has resulted in the development of a series of NeS-oriented fold and thrust that provide excellent outcrops of the Mesozoic successions. The geotectonic framework and the highly complex history of the basin are largely controlled by changes in the tectonics on the western margin of Gondwana (Riccardi, 1987; Howell et al., 2005, and references therein). Towards the end of the Cretaceous, continental sedimentation was widespread and the Neuquén Basin merged with other basins to the south (e.g., the San Jorge Basin) to produce a unique giant depocentre (Franzese et al., 2003; Howell et al., 2005). In the latest Cretaceous very high global sea levels resulted in the first marine transgression from the Atlantic, with shallow-marine deposits occurring over wide areas of the basin (Barrio, 1990; Malumián and Náñez, 2011). 2.2. Austral Basin, Argentina

2.1. Neuquén Basin, Argentina The Neuquén Basin is located in west-central Argentina and eastern Chile, between latitudes 34 S and 41 S. It is developed in Argentine territory in the provinces of Neuquén (from which takes its name), Mendoza, Río Negro and La Pampa. The succession exceeds 5000 m of marine and continental sedimentary rocks, which range from late Triassic to Paleocene in age. The basin has a broadly triangular shape (Fig. 1A, a) and two main regions are commonly recognized: the Neuquén Andes to the west, and the Neuquén Embayment to the east and southeast. The majority of the hydrocarbon fields in the basin are located in the Neuquén Embayment where most of the Mesozoic sedimentary record is in the subsurface and the strata are

The Austral or Magallanes Basin stretches with a NWWeSEE direction in almost all the south of the Patagonia to the south of parallel 47 S and involves the provinces of Santa Cruz, Tierra del Fuego and a small part of Chubut in Argentina and the province of Magallanes in Chile (Fig. 1A, b). It is a wide sedimentary recipient, generated from diastrophic movements from the Late Jurassic and filled with sediments from the Jurassic, Cretaceous and early Cenozoic. Its evolution and development is a product of interactive movement between the South American and Antarctic plates and has been interpreted as a back-arc marine basin (Riccardi, 1987). The subsidence probably began in the south in the latest Jurassic and continued throughout the Cretaceous and Paleocene to Pliocene, with some intermittent short events that left hiatuses and

Fig. 1. Study area. A, Argentina: a, Neuquén Basin; b, Austral Basin (main sedimentary basins of southern South America, modified from Malumián and Ramos, 1984). B, Brazil: sedimentary basins studied for this work (modified from Moreira et al., 2007).

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both local and regional unconformities (Riccardi, 1987; Robbiano et al., 1996). 2.3. Brazilian basins The geological evolution of eastern Brazilian marginal basins is related to the breakup of the Western Gondwana supercontinent and the Atlantic Ocean opening (e.g., Cainelli and Mohriak, 1999; Mohriak et al., 2008). Thirteen marginal basins are recognized where the oldest sedimentary sequences were deposited during Late Jurassic/Early Cretaceous times (Fig. 1B). Five tectonic phases are recognized for the development of sedimentary basins along the eastern Brazilian margin (Cainelli and Mohriak, 1999; Mohriak et al., 2008; Contreras et al., 2010). The material analyzed in this study is restricted to the fifth phase that extends in the Cretaceous from Albian to Maastrichtian and is characterized by open marine ocean conditions. The offshore Santos and Espírito Santo basins have received special attention from the oil industry owing to giant hydrocarbon reserves recently discovered. The Pelotas Basin is the southernmost Brazilian marginal basin. The boundaries are between 28 S and 34 S and coincide with the Polonio High in the northern Uruguay and Florianópolis High at the southern boundary of Santos Basin. The basin covers an area of 210,000 km2 of which 40,000 km2 are offshore. Between the Albian and Oligocene a trangressive sequence composed of thick sections of pelitic and clastic sedimentations is represented by the Atlântida, Tramandaí, Imbé and Cidreira formations (Bueno et al., 2007). The Santos Basin covers an area of 350,000 km2 completely offshore, located between the parallels 23 S and 28 S. The basin is bounded by Florianópolis and Cabo Frio highs, in the south and north respectively (Moreira et al., 2007). The sedimentary sequences were strongly influenced by the uplift of the Serra do Mar coastal mountain chain in the Late Cretaceous and subsequent organization of the coast-parallel Paraiba do Sul drainage system. The ancestral Paraiba do Sul tended to focus clastic influx into the northern and central Santos Basin during the Late Cretaceous and Paleogene (Modica and Brush, 2004). The Santonian and Campanian sedimentary successions have a thick progadational trend and are represented by the Juréia and Itajaí-açu formations (Moreira et al., 2007; Mohriak et al., 2008). During the Late Campanian and Maastrichtian a regressive pattern was established which resulted in the deposition of the conglomerates of Santos Formation (Moreira et al., 2007). The Espírito Santo Basin has an area of 46,000 km2 of which 4000 km2 is onshore. The basin is limited by the Vitoria Arch to the south and Abrolhos volcanic complex to the north (Cainelli and Mohriak, 1999). The Albian sedimentary sequence is characterized by a shallow calcareous platform represented by Regencia Formation. The CenomanianeEocene is characterized by a deep marine sequence corresponding to the pelitic packages from the Urucutuca Formation (França et al., 2007). The SergipeeAlagoas Basin has an approximate area of 33,000 km2 (Feijó, 1994). The northeast boundary of the basin is limited with Pernambuco-Paraiba Basin by the Maragogi High and the southwest limit is the Jacuipe Basin (França et al., 2007). During the AlbianeCenomanian the first open marine sequence in the basin was deposited, represented by the carbonates of Riachuelo Formation. The late CenomanianeConiacian Continguiba Formation represents a transgressive event deposited in a carbonatic ramp (França et al., 2007). The maximum transgressive event is registered in the early Campanian by the siliciclastic Calumbi Formation. During the late CampanianeMaastrichtian the sedimentary sequence is progradational and represented by the sandstones of Marituba Formation (França et al., 2007).

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3. Systematic palaeontology The suprageneric classification adopted is that proposed by Liebau (2005); for the genus, we follow the review by Whatley and Ballent (1996). In the systematic descriptions, the following conventions are employed: L, length; H, height; W, width; very small, <0.40 mm; small, 0.40e0.50 mm; medium, 0.51e0.70 mm; large, 0.71e0.90 mm; very large, >0.90 mm. Type and figured specimens are deposited in the collections of Museu de Paleontologia da Universidade do Vale do Rio dos Sinos (ULVG); Museu de Paleontologia, Ostracode section (MP-O), Universidade Federal do Rio Grande do Sul; Museo de La Plata, Argentina, sección Micropaleontología (MLPMi); Servicio Geológico Nacional, Argentina (SGN) and Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina, Laboratorio Micropaleontología (FCEN-LM) under their respective catalogue numbers. Subclass Ostracoda Latreille, 1806 Order Podocopida Sars, 1866 Suborder Cytherocopina Gründel, 1967 Superfamily Cytheroidea Baird, 1850 Family Progonocytheridae Sylvester-Bradley, 1948 Subfamily Progonocytherinae Sylvester-Bradley, 1948 Genus Majungaella Grekoff, 1963 Type species. Majungaella perforata Grekoff, 1963 Majungaella alta sp. nov. Fig. 2AeD Derivation of name. Portuguese, alta, high, derived from the shape of the carapace. Type and other material. Holotype ULVG-8736, L: 0.940, H: 0.720, W: 0.640 (Fig. 2AeC); paratype: ULVG-8737, L: 0.920, H: 0.640, W: 0.600 (Fig. 2D); eight carapaces. Type locality. Holotype, well SAN-04, 2952 m, Santos Basin; paratype, well SAN-04, 2907 m, Santos Basin, Brazil. Diagnosis. Very large and robust species, subtriangular carapace. Greatest height at mid-length. Ventral margin strongly convex. Ornamented with a reticulation and ribs parallel to the ventral and posterior margins. Description. Robust and very large carapace, subtriangular in lateral view. Maximum height and maximum width in the middle of the carapace. Anterior margin rounded; posterior margin short and truncated. Dorsal margin strongly convex. Ventrolateral margin inflated, overhanging ventral surface. Left valve larger than right, overlapping the dorsal margin and half of the anterior margin. Anterior marginal area slightly compressed. Biconvex in dorsal view. Conspicuous ribs parallel to posteroventral margins. Lateral surface reticulate, coarser in the middle of the carapace. Fine reticula disposed parallel to the anterior margin. Remarks. Majungaella alta sp. nov. differs from M. santosensis sp. nov., described below, in that the former is shorter and proportionally higher and wider. In addition, the greatest height is at midlength and the overlap in the middle of the carapace is more pronounced in M. alta. The new species shares with Majungaella sp. 1 of Miller et al. (2002, fig. 9g), from the Campanian of the Santos Basin, the development of the ribs, which are very conspicuous along the ventrolateral area; however, both differ in their lateral outline.

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Fig. 2. AeD, Majungaella alta sp. nov. ULVG-8736. A, holotype, right lateral view. B, left lateral view. C, dorsal view. D, paratype, ULVG-8737, right lateral view. E, F, Majungaella australis (Bertels, 1975a), FCEN-LM 858. E, RV, external lateral view. F, same valve, internal view. G, H, Majungaella hemigymnae Brenner and Oertli, 1976, MLP-Mi 892. G, left lateral view. H, dorsal view. IeK, Majungaella nematis Grekoff, 1963, MLP-Mi 894. I, right lateral view. J, dorsal view. K, detail of the ornamentation. LeN, Majungaella pavta Ballent, Ronchi and Whatley, 1998, MLP-Mi 886. L, left lateral view. M, dorsal view. N, detail of the ornamentation. OeQ, Majungaella praehemigymnae Valicenti and Stephens, 1984, MLP-Mi 893. O, left lateral view. P, dorsal view. Q, detail of the ornamentation in the anterodorsal region. RV ¼ right valve. Scale bars represent 100 mm.

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Distribution. CampanianeMaastrichtian, Santos Basin, Brazil. Majungaella australis (Bertels, 1975a) Fig. 2E, F 1975 Tumidoleberis australis Bertels, 1975a, pl. 6, figs. 4e6. 2005 Majungaella australis (Bertels, 1975a); Whatley et al., p. 312, fig. 2, 1e4. Brief description. Large, subtrapezoidal in lateral view, with the greatest length at mid-height and the greatest height at anterior cardinal angle. Ornamented by a coarse reticulum consisting of rows of very deep pits arranged subconcentrically and subparallel to the free periphery of the valve, and radiating mostly from a point near the anterior cardinal angle. Eye-swelling present. Dimensions (mm) of figured specimen: FCEN-LM 858, right valve, L: 0.840, H: 0.540. Remarks. Bertels (1975a) included this species in Tumidoleberis Deroo, but without doubt it is a species of Majungaella. As the trivial name had been subsequently used by Rossi de García and Proserpio for M. australis, so M. pseudonymos was proposed as replacement name (see Dingle and Majoran, 2001, p. 380; Whatley et al., 2005, p. 313). In external appearance and lateral outline M. australis, resembles M. pseudonymos described herein, but differs in possessing thicker ornamentation and in lacking secondary ornamentation and papillae over the external surface. Distribution. Middle Maastrichtian, Neuquén Basin, Argentina. Majungaella hemigymnae Brenner and Oertli, 1976 Fig. 2G, H 1976 Majungaella hemigymnae Brenner and Oertli, pl. 6, figs. 1e4; pl. 8, fig. 5. 1998 Majungaella hemigymnae Brenner and Oertli; Ballent et al., pl. 1, fig. 6. Brief description. Medium to large, subtriangular to subtrapezoidal in lateral view, with the greatest height just behind the anterior cardinal angle, very truncated posterior margin and distinctive posterior cardinal angle. Ornamentation with small ribs extending parallel to the ventral outline, mainly in the posterior part of the carapace and irregular ornamentation in the central area. Dimensions (mm) of figured specimen: MLP-Mi 892, L: 0.740, H: 0.490, W: 0.500. Remarks. The truncated posterior margin which gives the carapace a stout and angular appearance distinguishes M. hemigymnae from other species of the genus. The record from the Aptian of South Africa in Dingle (1984, fig. 17F) as Majungaella? hemigymnae is doubtful. It is only a fragmentary carapace, and it is not possible, therefore, to assign it precisely to any genus. Distribution. Upper part of the Hauterivian, Algoa Basin, South Africa; ValanginianeHauterivian, Austral Basin, Argentina. Majungaella nematis Grekoff, 1963 Fig. 2IeK 1963 Majungaella nematis Grekoff, pl. 5, figs. 141e145; pl. 9, figs. 213e232. 1998 Majungaella nematis Grekoff; Ballent et al., pl. 1, fig. 6. Brief description. Robust and large to very large, subtrapezoidal in lateral view. The ornamentation consists of a coarse, concentrically

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arranged reticulated pattern which extends ventrally parallel to the outline. Post-ocular sulcus well developed. Anterior and posterior regions compressed. Dimensions (mm) of figured specimen: MLP-Mi 894, L: 1.010, H: 0.655, W: 0.650. Remarks. Although the specimen recovered from Argentina is larger, it is very similar to those described by Grekoff from the Neocomian of Madagascar. The specimens from the Hauterivian of the Algoa Basin (South Africa) of Brenner and Oertli (1976) are more elongate and posteriorly acuminate compared to the original description by Grekoff. Distribution. TithonianeHauterivian, Madagascar; Late Valanginiane Hauterivian and late Aptianeearly Cenomanian, South Africa; Late Jurassic, India; Neocomian, DSDP site 249, Mozambique Ridge; BerriasianeValanginian, offshore NW Australia; Valanginiane Hauterivian, boreholes, Austral Basin and Hauterivian at Fontana Lake, Chubut Province, Argentina (Musacchio and Simeoni, 2008). Majungaella aff. nematis is cited (not figured) from probable Valanginian strata in southern Chile (Sigal et al., 1970). Majungaella pavta Ballent, Ronchi and Whatley, 1998 Fig. 2LeN 1998 Majungaella pavta Ballent et al., pl. 1, figs. 1e5. Brief description. Medium and tumid, triangular to subtriangular in lateral view, with the greatest length in the lower third and the greatest height almost coincident with the mid-length of the valve. Convex dorsal margin, an almost umbonate apex and a steep decline towards the posterior. Ornamented by fine concentrically arranged ribs, subparallel to margins. Primary reticulation situated between ribs. Second-order reticulation occurs in the solum of the primary reticule; four divisions in each solum are observed with SEM. Dimensions (mm) of figured specimen: MLP-Mi 886, L: 0.640, H: 0.400, W: 0.410. Remarks. The subtriangular outline in lateral view distinguishes M. pavta from others of the genus. It is close to M. uitenhagensis (Dingle) as emended by Valicenti and Stephens (1984, p. 186) from the late Valanginian and Hauterivian of South Africa; however, M. pavta is smaller, with the greatest height almost coincident with the mid-length, compressed periphery and with the posterior margin subangular with the bluntly apex pointed downwards. Distribution. Late TithonianeBerriasian and Valanginian, Neuquén Basin, Argentina. Majungaella praehemigymnae Valicenti and Stephens, 1984 Fig. 2OeQ 1984 Majungaella praehemigymnae Valicenti and Stephens, pl. 10, figs. 6e11; pl. 11, fig. 1. 1998 Majungaella praehemigymnae Valicenti and Stephens; Ballent et al., pl. 1, fig. 7. Brief description. Medium size and subtrapezoidal in lateral view, with the greatest length in the lower third and the greatest height immediately in front of the mid-length of the valve. Ventrolateral expansion regularly developed. Surface ornamented with large puncta, faintly developed ribs and weak reticulation. Dimensions (mm) of figured specimen: MLP-Mi 893, L: 0.615; H: 0.400; W: 0.330. Remarks. It closely resembles M. hemigymnae Brenner and Oertli from the same South African basin and from the Valanginiane

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Hauterivian of the Austral Basin in Argentina (see above). Nevertheless, it is distinguished by its smaller size, smoother surface and proportionally higher carapace, which, in addition, has a more rounded anterior margin and the greatest height situated more anteriorly. Distribution. Late Valanginian, South Hauterivian, Austral Basin, Argentina.

Africa;

Valanginiane

Majungaella pseudonymos Whatley, Ballent and Szczechura, 2005 Fig. 3A, B 1980 Majungaella australis Rossi de García and Proserpio, pl. 3, figs. 3, 4, 7. 2005 Majungaella pseudonymos Whatley, Ballent and Szczechura nov. nom., p. 313. Brief description. Very large and robust, subtrapezoidal in lateral view. Greatest length at mid-height and greatest height at cardinal anterior angle. The lateral surface with a distinct polygonal reticulation in a concentric pattern; small papillae in the intersection of the primary reticula are also present. Dimensions (mm) of figured specimen: SGN 2240 , left valve L: 0.920, H: 0.400. Remarks. Majungaella pseudonymos was proposed as a new name for Majungaella australis Rossi de García and Proserpio, 1980 (see Whatley et al., 2005, p. 313), which is a junior secondary homonym of Majungaella australis (Bertels, 1975a). In the former species, the presence of a distinct polygonal reticulation in a concentric pattern and a small papilla in the intersection of the primary reticula, distinguish it from Majungaella australis (Bertels) in which the reticulation is notably coarser and lacks small intercostal papillae. Distribution. Late Campanian?eMaastrichtian, Chubut Province, Argentina; middleelate Campanian, James Ross Island, Antarctic Peninsula (Fauth et al., 2003). Also, a partly crushed specimen of uncertain identification from the late Campanian, Snow Hill Island, Antarctic Peninsula (Dingle, 2009). Majungaella aff. pseudonymos is recorded from the SantonianeCampanian, northwestern James Ross Island (Florisbal et al., 2010; Florisbal, 2011, pers. comm. 2011). Majungaella santacruziana (Rossi de García, 1972) Fig. 3CeF 1972 Novocythere santacruziana Rossi de García, pl. 1, fig. 7aec. 1975 Tickalaracythere scheibnerovae Krömmelbein, pl. 4, figs. 12, 13, text-figs. 5, 6. 1998 Majungaella santacruziana (Rossi de García); Ballent et al., pl. 1, figs. 9e11. Brief description. Large and robust, pyriform, subtrapezoidal in lateral view and posterodorsally upturned. Lateral surface ornamented by a coarse and polygonal reticulation in concentric pattern. Secondary reticulation and punctation occur in the sola of primary reticula. Anterior region compressed. Dimensions (mm) of figured specimens: MLP-Mi 899, L: 0.900, H: 0.590, W: 0.540; MLPMi 895, L: 0.890, H: 0.575, W: 0.530. Remarks. As already pointed out by Rossi de García, 1977 (p. 120), Tickalaracythere scheibnerovae Krömmelbein is a junior synonym of M. santacruziana (ex Novocythere santacruziana Rossi de García). In this respect, Whatley and Ballent (1996) emended the diagnosis of Majungaella to accommodate the species of Tickalaracythere and Novocythere. Ballent et al. (1998, pp. 49, 50) have provided a complementary description of M. santacruziana in order to clarify its status. They have also explained the differences between

M. nematis Grekoff sensu Brenner and Oertli (1976, pl. 5, figs. 11, 12) and sensu McLachlan et al. (1976, pl. 15, figs. 14, 15), both from the Hauterivian of wells in the Algoa Basin and synonymized with M. santacruziana. Distribution. AlbianeCenomanian, Great Australian Basin; Albian, Austral Basin, Argentina. Majungaella santosensis sp. nov. Fig. 3GeM Derivation of name. After the Santos Basin where the type material was recorded. Type and other material. Holotype: ULVG-8738, L: 1.08, H: 0.720, W: 0.640 (Fig. 3GeK); paratype: ULVG-8739, L: 1.050, H: 0.675, W: 0.600 (Fig. 3LeM); 29 carapaces. Type locality. Holotype: well 1-BBS-76, 2644 m, Santos Basin; well SAN-04, 3357 m, Santos Basin, Brazil. Diagnosis. Very large and robust species, pyriform carapace outline, posterodorsally upturned. Ornamented with a concentric pattern of ribs, parallel to the periphery of the valves. Inter-rib areas covered with irregular reticula, more prominently developed in the mid part of the valves. Description. Robust and very large carapace, pyriform. Greatest height positioned at the anterior cardinal angle. Anterior margin rounded and oblique in the upper part; posterior margin upturned dorsally. Dorsal margin almost straight and gently sloping to the posterior. Ventrolateral margin inflated, overhanging ventral surface. Anterodorsal and posterodorsal extensions, more prominent in the left valve. Subelliptical and inflated in dorsal view. Ribs in concentric pattern, parallel to free margins of the valve. The primary ribs bifurcate in the anteroventral part to produce fine secondary ribs. Inter-rib areas covered with an irregular reticulum, more prominently developed in the mid part of the carapace. Fine reticulation disposed in rows parallel to the anterior margin. Small, but distinct eye-swelling. Sieve-type normal porecanals near the muri of the reticula. Remarks. The reticulum over the anterior part of the valve shows a variable degree of development, being distinct in some specimens and thinner and densely punctuate in the anterior regions of others. Distribution. CampanianeMaastrichtian, Santos and Espírito Santo basins, Brazil. Majungaella sp. of Ballent, Ronchi and Whatley, 1998 Fig. 3NeP 1998 Majungaella sp. Ballent et al., pl. 1, fig. 2. Brief description. Very large, subtriangular in lateral view. Dorsal margin sloping strongly to the posterior. Ventral margin very convex. Greatest height just anterior of mid-length. Low, elongate eye swelling. Coarse concentric ornament with an irregular reticulum parallel to valve margin; secondary reticulation poorly developed. The anterior and posterior peripheral areas bear an almost celate ornamentation. Dimensions (mm) of figured specimen: MLP-Mi 901, L: 0.930, H: 0.660, W: 0.530. Remarks. Majungaella sp. strongly resembles M. santacruziana (Rossi de García) from the Albian of the Austral Basin, from which it may have evolved. Nevertheless, it differs in details of the

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Fig. 3. A, B, Majungaella pseudonymos Whatley, Ballent and Szczechura, 2005, SGN 2240 . A, RV, external lateral view. B, same valve, internal view. CeF, Majungaella santacruziana (Rossi de García, 1972), MLP-Mi 899. C, right lateral view. D, left lateral view, MLP-Mi 895. E, dorsal view. F, detail of the ornamentation. GeM, Majungaella santosensis sp. nov. GeK, holotype ULVG-8738. G, right lateral view. H, left lateral view. I, dorsal view. J, detail of the ornamentation. K, detail of the sieve porecanals. L, M, paratype ULVG-8739. L, right lateral view. M, ventral view. NeP, Majungaella sp. of Ballent, Ronchi and Whatley (1998), MLP-Mi 901. N, right lateral view. O, dorsal view. P, detail of the ornamentation. Q, Majungaella sp. of Ceolin, Fauth and Coimbra (2011), MP-O-2138, right lateral view. RV ¼ right valve. Scale bars represent 100 mm.

ornamentation. A characteristic feature of the former species is the irregular concentric reticulation, poorly developed secondary reticulation and anterior and posterior peripheral celate areas, which are absent or weaker in the latter. Specimens from the mid-

late Campanian of Antarctica externally appear more robust and thicker, egg-shaped in lateral view. The present species is closest to M. ticka (Krömmelbein), from the mid-Cretaceous of central Queensland, Australia (see Whatley et al., 1994). However, the latter

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is proportionally higher and the coarse concentric ornament is more regular than in the Argentinian species. For its part, M. wilsoni Dingle, 2009, confined to the Maastrichtian of New Zealand and with similar ornamentation, is proportionally lower and more elongate, and more compressed anteriorly than the present species. Majungaella sp. of Ballent et al. differs from M. pseudonymos (see above) in having a different lateral outline and ornamentation pattern. Although probably a new species, with a few specimens from Argentina and the Antarctic Peninsula, its identification open nomenclature is maintained. Distribution. SantonianeCampanian, boreholes, and late Campanian, Cerro Cazador Formation (Malumián et al., 2000, p.11), Santa Cruz Province, Austral Basin, Argentina; mid-late Campanian, James Ross Island, Antarctic Peninsula (Fauth et al., 2002, 2003). Majungaella sp. of Ceolin, Fauth and Coimbra, 2011 Figs. 3Q and 4A 2011 Majungaella sp. of Ceolin et al., fig. 5f. Brief description. Large, sub-pyriform in lateral view, with maximum height positioned anteriorly. In dorsal view, it is very compressed in the anterior region. Anterior margin rounded, posterior margin truncated. Ventral margin gently convex, dorsal margin convex and inclines towards the posterior end. Surface ornamented with weak concentric ribs disposed in the ventral, anterior and posterior regions. Dimensions (mm) of figured specimen: MP-O-2138, L: 0.920; H: 0.500; W: 0.570. Remarks. The outline of this species resembles Majungaella sp. of Ballent et al., 1998, which is, however, higher and more strongly ornamented (see above). It is likely to be a male owing to its elongated carapace. Despite the fact that the present specimen is very poorly preserved, it is undoubtedly referable to Majungaella. Distribution. Maastrichtian, Pelotas Basin, Brazil (Ceolin et al., 2011). Majungaella sp. 1 Fig. 4BeD Description. Very large, subtrapezoidal in lateral view, greatest height at anterior cardinal angle. Anterior margin asymmetrically rounded; anterior marginal area moderately broad and compressed; posterior margin short and slightly upturned. Dorsal margin almost straight and sloping gradually to the posterior margin. Ventral margin broadly convex, posteroventrally upturned. Subelliptical in dorsal view, with very compressed anterior part. An anterodorsal extension, more prominent in the left valve is present. Concentric ribs well-marked along the ventral, posterior and dorsal margins. An inter-rib reticulation is present, but not clearly defined. Rows of conspicuous papillae are disposed over the ribs. Dimensions (mm) of figured specimen: ULVG-8734, L: 0.930; H: 0.660; W: 0.450. Remarks. The outline of the carapace and the presence of rows of papillae on the ribs distinguish Majungaella sp.1 from other representatives of the genus. Although probably a new species, with only a single specimen, it is retained under open nomenclature. Distribution. Santonian, Santos Basin, Brazil. Majungaella sp. 2 Fig. 4EeG Description. Very large and tumid, subtrapezoidal in lateral view, with the greatest height at mid-length. Anterior margin broad and

symmetrically rounded; posterior margin with truncated apex at the middle third of the height. Dorsal margin somewhat convex and downwardly curved to the posterior margin. Ventral margin strongly convex. In dorsal view subovoidal, with the greatest width in the middle of carapace and moderately compressed anteriorly. Surface ornamented by concentric ribs in an almost circular arrangement and parallel to the valve margins. These ribs are more developed ventrolaterally and are thinner at anterior and posterior areas of the valves. Inter-rib areas moderately reticulate. Internal features not seen. Dimensions (mm) of figured specimen: ULVG8735, L: 1.000; H: 0.680; W: 0.680. Remarks. The outline of the carapace with the dorsal margin markedly convex and the almost circular pattern of the ribs differentiate this species from the others. Because only one specimen has been recovered this species is left in open nomenclature. Distribution. Early Campanian, Santos Basin, Brazil. Majungaella sp. 3 Fig. 4HeJ Description. Large and robust, subtrapezoidal in lateral view. Anterior margin broad and asymmetrically rounded with apex below mid-height; anterior marginal area moderately wide and compressed. Posterior margin obtusely angulate. Dorsal margin slightly convex in the middle part, sloping posteriorly. Cardinal angles well marked. The greatest height positioned at anterior cardinal angle. Ventral margin almost straight to slightly convex in the anterior and mid third of its length and posteroventrally upturned. In dorsal view the carapace is subelliptical and inflated, compressed anteriorly and with the greatest width at mid-length. Ornamented with ribs, the most distal one being parallel to the free margins. Inter-rib areas covered by pentagonal reticulation; abundant papillae over the muri. Dimensions (mm) of figured specimen: MLP-Mi 1919, L: 0.850, H: 0.540, W: 0.505. Remarks. The subtrapezoidal lateral outline with well-marked cardinal angles, and the inter-rib areas covered by pentagonal reticulation with abundant papillae over the muri characterize this species and distinguishes it from M. pseudonymos. The ornamentation resembles that of Parahystricocythere ericea Dingle, 2009 (pl. 2, figs. 12e16), a reticulate spinose progonocytherid known only from the uppermost Maastrichtian sediments of Waipara Gorge, New Zealand. Given the close similarity and the known stratigraphic distribution of Majungaella and Parahystricocythere in Australasia, Dingle (2009, p. 164) suggested that the former “seems the most obvious progenitor of Parahystricocythere”. The New Zealand species is, however, ovate to piriform, with a distinctive upturned posteroventral outline and a low but prominent elevation at the posterior end of the dorsal margin, which is surmounted with small spines. Although probably a new species, with only a single specimen, it is retained under open nomenclature. Distribution. Campanian, Austral Basin, Argentina. Majungaella sp. 4 Fig. 4KeM Description. Medium, subpyriform in lateral view. Anterior margin very broadly, but asymmetrically rounded; anterior region very compressed. Posterior margin somewhat truncated, upturned dorsally. Dorsal margin straight and strongly sloping posteriorly. Ventral margin slightly convex and medianly obscured by valve tumidity. Greatest height, which coincides with the anterior

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Fig. 4. A, Majungaella sp. of Ceolin, Fauth and Coimbra (2011), MP-O-2138, dorsal view. BeD, Majungaella sp. 1. ULVG-8734. B, right lateral view. C, left lateral view. D, dorsal view. EeG, Majungaella sp. 2. ULVG-8735. E, right lateral view. F, left lateral view. G, dorsal view. HeJ, Majungaella sp. 3, MLP-Mi 1919. H, left lateral view. I, dorsal view. J, detail of the ornamentation. KeM, Majungaella sp. 4, MLP-Mi 1920. K, left lateral view. L, dorsal view. M, detail of the ornamentation. N, O, Majungaella sp. 5, lost. N, right lateral view. O, dorsal view. Scale bars represent 100 mm.

cardinal angle, is just over 70% of the greatest length. Carapace is subelliptical in dorsal view, anterior area more compressed than the posterior. Well-defined ribs, concentrically disposed and parallel to the free margins. Inter-rib areas clearly reticulate.

Second-order reticulation occurs in the sola of primary reticula. Each solum is generally divided into four small fossae. Anterior marginal area is densely punctate. Dimensions (mm) of figured specimen: MLP-Mi 1920, L: 0.620, H: 0.450, W: 0.400.

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Remarks. This species is closest to Majungaella sp. of Ballent et al. (see above) which is, however, larger, more robust and ornamented. Despite the fact that the present specimen has the appearance of a juvenile, it is clearly a bona fide member of Majungaella, but is retained under open nomenclature. Distribution. Probably CampanianeMaastrichtian, Austral Basin, Argentina. Majungaella sp. 5 Fig. 4N, O 1979 Tickalaracythere sp. nov. Krömmelbein pl. 3, fig. 3.5. Brief description. Robust and large carapace, pyriform. Greatest height positioned at the anterior cardinal angle. Anterior margin broadly rounded; posterior margin upturned. Dorsal margin almost straight and sloping to the posterior. Ventrolateral margin convex, overhanging ventral surface. Subelliptical and inflated in dorsal view; anterior marginal area strongly compressed. Shell surface ornamented by ribs in a concentric pattern, parallel to free margins of the valve. Abundant small papillae are disposed over the ribs and the muri. Strongly punctate in the mid region and finely and densely in the anterior. Dimensions (mm) of figured specimen: lost, L: 0.880, H: 0.680, W: 0.560. Remarks. Majungaella sp. 5 coincides with the species figured by Krömmelbein in having a similar lateral outline and pattern of ornamentation, including the presence of small papillae disposed over the ribs and the muri. Distribution. CampanianeMaastrichtian, Sergipe Basin, Brazil. 4. Records of Majungaella 4.1. India Neale and Singh (1986, pl. 2, figs. 13e16) described M. biswasi, a long ranging form from the BathonianeCallovian (scarcely represented) to Tithonian, Kachchh region of northern peninsular India, which probably corresponds to M. perforata (see Khosla et al., 1997). This record comes from only one core sample (Banni well N 2) and seems to be the earliest record of the genus in this region. From different localities in the same Indian region, M. perforata, M. perforata kachchhensis Khosla et al. (1997, pl. 4, fig. 11 and figs. 12e14, respectively) and M. rasilis Khosla et al. (1997, pl. 4, figs. 15e17) are frequent species through mid-late Callovian deposits (Khosla et al., 1997, 2003, 2005, 2006). M. mundula (Grekoff) was identified from the CallovianeOxfordian by Kulshreshtha et al. (1985) and M. perforata from the Upper Jurassic by Jain and Mannikeri (1975). M. spitiensis (Jain and Mannikeri, 1975, fig. 1, FeH) is certainly a junior synonym of M. perforata while M. brentonensis (Dingle) sensu Guha, 1976 (pl. 3, fig. 16a, b) from the Upper Jurassic of Kutch, is a representative of Fastigatocythere Wienholz. Latest Early Cretaceouseearly Late Cretaceous (Albiane Coniacian) ostracods from subsurface of Jaisalmer Basin were referred to by Singh (1997) and Andreu et al. (2007). M. manheratibbaensis Singh, 1997 (pl. 7, figs. 8e13; pl. 8, figs. 1, 3) from the upper member (Cenomanian) of the Goru Formation is a valid member of Majungaella. Other species, retained in open nomenclature (see Andreu et al., 2007, pl. 9, figs. 7e21), were subsequently reallocated to the genus Malagaspyella Babinot et al., 2009 (p. 11), a new Bracychytherinae described from the CenomanianeTuronian of Magadascar. Majungaella indrai Singh,

1997 (pl. 8, fig. 2) from the Cenomanian of Rajasthan certainly is not a representative of Majungaella. 4.2. East Africa (Madagascar, Tanzania, Somalia) The first record of the genus is in the south-west Madagascar and represented by M. mundula in early Callovian deposits of the Morondava Basin (Mette and Geiger, 2004b). From the Oxfordian of the same basin, Mette and Geiger (2004c) recognized four species: M. microperforata (pl. 2, figs. 7e10; pl. 3, figs. 1, 2, restricted to the middle Oxfordian), M. ventriosa (pl. 2, figs. 1e6), M. glabra (pl. 3, figs. 3e10; pl. 4, figs. 1e3) and Majungaella sp. 1 (pl. 4, fig. 8, possibly a juvenile of M. perforata kachchhensis), and M. perforata (pl. 4, figs. 4e7) from the middle Oxfordian to lower Kimmeridgian. Grekoff (1963) and Rafara (1990) recognized M. mundula (mid Callovian), M. perforata (Tithonian) and M. nematis (TithonianeHauterivian) from the Majunga Basin, including the Antsiranana region (ex Diégo Suarez) in northern Madagascar. The latter author also identified Tickalaracythere? sp. A (pl. 2, fig, 8e10) from the Hauterivian of the Majunga Basin, which is certainly very close to M. pyriformis Bate and Bayliss. Grekoff (in Collignon et al., 1979, pl. 5, fig. 10) and more recently Babinot et al. (2009, pl. 1, figs. 16e18) recorded M. pyriformis, from the middle-lowermost upper Albian of the same region Sigal (1974, pl. 1, fig. 4a,b) recognized M. nematis from the “Neocomian” of the DSDP site 249, Leg 25, Mozambique Ridge, Indian Ocean. The genus first appears in Tanzania represented by M. mundula from the mid Callovian of Mandawa Anticline (Bate, 1975, pl. 5, figs. 10e13). From the same region, Bate also recongnized M. perforata (pl. 3, figs. 1e3, 7) from early?elate Tithonian deposits, and described three new species: M. oxfordiana (pl. 5, figs. 4e9), late Oxfordian; M. kimmeridgiana (pl. 4, figs. 4e11; pl. 5, figs. 1e3), early Kimmeridgian; and M. praeperforata (pl. 3, figs. 4e6, 8e10, pl. 4, figs. 1e3), mid to late Kimmeridgian. Bate and Bayliss (1969, pl. 5, fig. 13) described M. pyriformis from the Albian of the Wami river area. Mette (1993) recognized three species in northern Somalia: M. vertireticulata Mette (pl. 4, figs. 14e16; pl. 5, figs. 1e13) from the loweremiddle? Oxfordian; M. cf. praeperforata Bate (pl. 4, figs.5e13), and Majungaella sp. 1 (pl. 3, figs. 21e23; pl. 4, figs. 1e4) from the upper Oxfordianelower Kimmeridgian. 4.3. Middle East Rosenfeld and Raab, 1984 (pl. 8, figs. 23, 24) identified M. sp. cf. M perforata from the ValanginianeHauterivian Helez Formation (subsurface, Coastal Plain, Jerusalem). 4.4. Argentina/Chile Two species have been recognized in the Neuquén Basin. M. pavta, although scarcely represented from the uppermost TithonianeBerriasian (wells from the Entre Lomas area, eastern part of the basin), is a frequent species in the Valanginian of several outcrops and boreholes drilled in the eastern part of the basin (Ballent et al., 1998; Ballent in Schwarz, 2003; Concheyro et al., 2009). The other species, Majungaella australis (Bertels) is known from the middle Maastrichtian of Río Negro Province (Bertels, 1975a). Eight species of Majungaella have been recognized in the Argentinian part of the Austral Basin and from outcrops in the Chubut Province. Majungaella praehemigymnae and M. hemigymnae were identified during the ValanginianeHauterivian interval. Majungaella nematis, with a wider distribution, was recorded from

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the ValanginianeHauterivian of boreholes (Ballent et al., 1998), Valanginian, YPF.SCA. CSO.39 (Cañadón Salto), Santa Cruz (as Majungaella sp. A of Kielbowicz et al., 1983, pl. 6, fig. 13) and Hauterivian at Fontana Lake, Chubut Province, Argentina (Musacchio and Simeoni, 2008). Majungaella aff. nematis has been reported (not figured) from probable Valanginian strata in southern Chile (Sigal et al., 1970). M. santacruziana is frequent in Albian samples of the YPF.SC. ALEP e-2 (Laguna El Palo) borehole drilled in the southeast part of the Austral Basin (Ballent, 1998; Ballent et al., 1998). Late Cretaceous species are Majungaella sp. of Ballent et al. from the SantonianeCampanian (Ballent et al., 1998) and late Campanian, Cerro Cazador Formation (Malumián et al., 2000, p.11), Santa Cruz Province and M. pseudonymos from the late Campanian?e Maastrichtian outcrops in Chubut Province (Rossi de García and Proserpio, 1980). Two new species, described in the present paper, are Majungaella sp. 3 from the Campanian and Majungaella sp. 4, from probable CampanianeMaastrichtian deposits. 4.5. Australia According to Guzel (in press), the right valve illustrated by Oertli, 1974 (pl. 7, fig. 11) as Metacytheropteron? from one sample at DSDP Site 261 (core 33, dated late Oxfordian by nannofossils), northwest of Perth, western Australia, is very similar to the male right valve of Majungaella oxfordiana of Bate, 1975 (pl. 5, fig. 4) from late Oxfordian deposits in Tanzania. This record, if confirmed, would be the earliest of the genus in Australia. Damotte (1992, pl. 2, fig. 7) recognized M. nematis in the BerriasianeValanginian Barrow Group on Exmouth Plateau. She also recognized Majungaella sp. 20R (pl. 2, fig. 8) and Majungaella sp. 40R (pl. 8, figs. 9, 10) in the Neocomian of Exmouth Plateau. The first is a small left valve, surely a juvenile, which doubtfully resembles M. bifurcata. The second, with a large subtrapezoidal carapace ornamented by strong ribs that are concentrically arranged, could be better accommodated in Centrocythere Mertens. By mid AlbianeCenomanian times, M. ticka (Krömmelbein) and M. scheibnerovae (Krömmelbein), which is a junior synonym of M. santacruziana (Rossi de García), were present in the Great Artesian Basin, southwestern Queensland, Australia (Krömmelbein, 1975, pl. 4, figs. 12, 13; Scheibnerová, 1980, pl. 14, fig. 1; Whatley et al., 1994). Bate (1972, pl. 5, figs 7e9; pl. 6, figs. 2e7; pl. 7, figs. 1e4) described M. annula from the SantonianeCampanian of the Carnarvon Basin; the same species was identified by Neale (1975, pl. 8, fig. 8) from the Santonian of Gingin, western Australia. 4.6. South Africa In South Africa, Majungaella is both abundant and diverse in the Neocomian. The first record is Majungaella uitenhagensis (Dingle), described as Neocythere (Neocythere) uitenhagensis Dingle (1969, fig. 11). Subsequent works by McLachlan et al. (1976), Brenner and Oertli (1976) and Dingle (1984) recognized M. nematis from the upper ValanginianeHauterivian of Zululand and the Algoa Basin and upper Aptianelower Cenomanian of Zululand, according the Dingle. Brenner and Oertli (1976) described two species from the upper ValanginianeHauterivian: M. hemigymnae (pl. 6, figs. 1e4; pl. 8, fig. 5) and M. bifurcata (pl. 5, figs. 13e21; pl. 8, fig. 4) and M. grekoffi (as Fastigatocythere? grekoffi, pl. 6, figs. 5e12; pl. 8, fig. 6) from the Valanginian, all from the Sundays River Formation, Algoa Basin. From the same age and basin, Valicenti and Stephens (1984, pl. 10, figs. 6e11) described M. praehemigymnae. The genus survives into the Santonian according to the mention of Majungaella sp. K9A3 of Dingle, 1996 (table I, p. 6) from the AlbianeSantonian interval in the Kudu boreholes on the Orange Shelf of southernmost Namibia (McMillan, 1990).

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Majungaella sp. 327/16 of Dingle (1984, fig. 19AeC) from the mid Albian at DSDP site 327 on the Falkland Plateau, which seems to have an antimerodont hinge, should be referred to the Cytherideidae (Whatley and Ballent, 1996). 4.7. Brazil The genus was recorded from Brazilian basins by Krömmelbein (1979) with the description of Tickalaracythere sp. nov. (Pl. 3, fig. 3.5), from CampanianeMaastrichtian, in the Sergipe Basin, which is certainly conspecific with Majungaella sp. 5, described herein. Two other records are Majungaella. sp. 1 of Miller et al. (2002, fig. 9f), from the Campanian of the Santos Basin and Majungaella sp. of Ceolin et al. (2011) (fig. 5f) from the Maastrichtian of the Pelotas Basin, the southernmost basin of Brazilian continental margin. Also, five species are presented in this paper: Majungaella sp. 1 (Santonian, Santos Basin); Majungaella sp. 2 (lower Campanian, Santos Basin), M. santosensis sp. nov. (CampanianeMaastrichtian, Santos and Espírito Santo basins); M. alta sp. nov. (Campaniane Maastrichtian, Santos Basin); and Majungaella sp. 5 (Campaniane Maastrichtian, Sergipe Basin). These undoubtedly enlarge the diversity of the genus during the Late Cretaceous in South America. 4.8. Antarctica Two species of Majungaella are known from the middleelate Campanian Hamilton Member in the upper part of the Santa María Formation at Hamilton Point on James Ross Island; they are, Majungaella sp. of Ballent et al. (1998) and Majungaella pseudonymos (Rossi de García and Proserpio), which were figured by Fauth et al. (2003) in their plates 1, figs. 19e21 and 2, figs. 1e3, respectively. Although of uncertain identification, Dingle (2009, pl. 2, fig. 10) recorded the last species from the upper Campanian of the lower part of the overlying Snow Hill Formation (Dingle, 2009, p. 163). Majungaella aff. pseudonymos has also been recorded from the SantonianeCampanian of northwestern James Ross Island (Florisbal et al., 2010; Florisbal, 2011, pers. comm., 2011). Szczechura (2001) described and illustrated Majungaella antarctica (pl. 2, figs. 5e8) from the early Eocene La Meseta Formation of Seymour Island (¼Isla Marambio) in the northern Antarctic Peninsula. Dingle and Majoran (2001, fig. 2A, B, D) recorded Majungaella sp. 4471 from early and late Oligocene glaciomarine sediments of the Cape Roberts boreholes in Victoria Land Basin, Ross Sea area. Szczechura and Blaszyk (1996) identified a species of Majungaella as ? Loxocythere sp. (pl. 40, figs. 4e6) from the Pecten Conglomerate (late Pliocene) of Cockburn Island. This species has been also identified as Tumidoleberis sp. nov. in the late Pliocene Cage Formation (sensu Lirio et al., 2003, p. 304), exposed on the south coast of Cape Cage, northern James Ross Island (Concheyro et al., 2007, Ballent, unpublished data). 4.9. New Zealand Dingle (2009) described three new species, two of which are confined to the uppermost Maastrichtian of the Mid-Waipara River Gorge, South Island; they are M. wilsoni (pl. 2, figs. 2e5) and M. waiparaensis (pl. 2, figs. 6e9). The third is Majungaella sp. 4978 (pl. 2, fig. 11) from the uppermost Maastrichtian of Pukehou, which is the only representative of the genus found in the outcrops on North Island, New Zealand. 4.10. Doubtful species of Majungaella Majungaella? minuta Swain, 1976 (pl. 1, figs. 19e21, 23) from the Aptian/Albian of DSDP sites in the north-west Atlantic, has

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a hemimerodont hinge and is very much smaller (L ¼ 0.360 mm) than all other species. This is possibly a Metacytheropteron Oertli. Bassiouni (2002, pp. 61e63, pl. 13) described five species of Majungaella from late Late Aptianelate Cenomanian deposits of Gebel Khutmiya, Sinai, Egypt. However, despite all of them having entomodont hinges and the specimens illustrated in his pl. 13, figs 1, 2 resembling M. pyriformis, they have a distinctive caudal process, which is not diagnosed for the genus. These species are more likely to belong to Neocythere Mertens. Fig. 5 summarizes the records, in stratigraphical order, of the valid species of Majungaella referred to in the text, including those newly described. 5. Abundance and depositional environment The ostracod faunas with Majungaella species recovered from MiddleeLate Jurassic deposits in India indicate a relatively shallow-water marine environment with shoreline fluctuations in a tectonically unstable shelf zone (Neale and Singh, 1986; Khosla et al., 1997). These conclusions are in broad agreement with the associated foraminifers, whose assemblages are dominated by species of Vaginulinidae and Nodosariidae (Gaur and Talib, 2009 and references therein). M. manheratibbaensis from the Cenomanian of Jaisalmer occurs associated with other ornamented cytheroids in a fluctuating inner neritic environment (Singh, 1997). In southwest Madagascar, the diverse CallovianeOxfordian ostracod assemblages, including species of Majungaella, have been assigned to warm and shallow open marine environments with a predominance of siliciclastic sedimentation (Mette and Geiger, 2004b). Ostracod faunas from the AlbianeCenomanian of northern Madagascar are diverse although numbers of specimens are low. In this case, M. pyriformis from the mid-lowermost upper Albian is associated with a wide variety of planktonic foraminifers (nearly a dozen species) probable indicating a depth greater than a shelf enviroment (Collignon et al., 1979). Majungaella mundula from the middle Callovian of Tanzania has been recovered from deposits of a shallow water shelf environment, in which the closeness of land is evidenced by the presence of freshwater ostracods and charophytes (Bate, 1975). In the same region, M. kimmeridgiana is one of the three dominant species of the early Kimmeridgian fauna. Althought normal marine conditions prevailed, this is a poor fauna in terms of number of species but excessively rich in number of individuals. Bate (1975) attributed its reduced diversity to a probable lowering of the water temperature. Species of Majungaella from the Algoa Basin in South Africa are associated with other highly ornamented ostracods (Sondagella Dingle, for example) with a robust carapace and a conspicuous eye tubercle; this usually develops in response to an environment typified by shallow water and, as a consequence, a rich food supply (Brenner and Oertli, 1976; Valicenti and Stephens, 1984). This agrees with the diverse associated foraminifers (McMillan, 2003), which indicate water depths probably typical of innermost shelf, and high oxygen levels. In the “Neocomian” of Israel, levels containing M. cf. perforata correspond to a shallow (littoral), warm marine environment (Rosenfeld and Raab, 1984). Meanwhile, Majungaella sp. K9A3 of Dingle (1996) has been recognized in AlbianeSantonian continental slope (and presumably cool-water) sediments from off Namibia (McMillan, 1990). Majungaella pavta from the Valanginian of the Neuquén Basin in northern Patagonia, Argentina is frequently recorded, although with few specimens; its association with diverse lagenid foraminifers and nannofossils indicates a shelf and nearshore environment

with clear, warm and well-oxygenated waters (Ballent et al., 2006). In the eastern part of the basin, its association with diverse cytherellids typify a marginal marine environment, with warmtemperate waters (Ballent and Ronchi, 1999). Majungaella santacruziana dominates the Albian fauna of the Argentinian part of the Austral Basin in southern Patagonia and represents 45% of the total individuals recorded (Ballent, 1998). The species is associated with abundant radiolarians and benthonic and planktonic foraminifers. In this case, the depositional environment corresponds to cooltemperate, well-oxygenated, outer shelf and upper slope (Ballent, 1998). This is also supported by oxygen isotope data of dimitobelid belemnites referred to by Pirrie et al. (2004), which suggest relatively cool temperatures (mean 9.5  C) for the early Albian of Argentina at a palaeolatitude of approximately 58 S. Malumián et al. (2000) referred Majungaella sp. of Ballent et al. (1998) to an infralittoral shallow environment, associated with diverse arenaceous foraminifers. Bertels (1975a) concluded that the mid-Maastrichtian ostracod species from northern Patagonia (including M. australis) lived in relatively warm (minimum of 17  C), normal salinity, clear waters in outer shelf (minimum of 150 m) conditions. Diverse benthonic (mainly species of Bolivina, Bulimina, Neobulimina) and planktonic (some carinate but the majority heterohelicids) associated foraminifers corroborated that environment (Bertels, 1975a, b). On the other hand, the rich association of calcareous nannofossils and abundant marine reptiles suggest shallow inner to middle shelf environments (Gasparini et al., 2007). By SantonianeCampanian times, at Gingin in the northern part of the Perth Basin in Western Australia, Majungaella is relatively rare and accounts for less than 1% of the fauna. It is suggested that the Gingin Chalk may be regarded as a warm-water deposit laid down in a shallow-shelf sea (around 100 m deep) with a minimum temperature of not less than 10  C. This agrees with the associated well-developed trachyleberid species, which suggests a relatively shallow-shelf sea area; in addition the Platycopina confirm warm, clear, shallow seas (Neale, 1975). The influence of warmer water currents in Western Australia is suggested by the moderate abundance of keeled planktonic foraminifera throughout the CampanianeMaastrichtian (see Huber, 1992). Considering the fauna of the Santos Basin on the Brazilian continental margin, the Majungaella species occur associated mostly with trachyleberidids, cytherurids, cytherellids and bairdiids (Piovesan et al., 2010; Bergue et al., 2011), which indicates shallow, warm environments. In all Brazilian basins where the genus has been recorded, it is always relatively rare in the samples. In relation to the late Cretaceous species of Majungaella in New Zealand, Dingle (2009, p. 187) concluded that the contrasts between the faunas at Waipara and Pukehou probably reflect different depositional environments. While M. wilsoni and M. waiparaensis preferred shallow and warm habitats (Waipara locality, in a proximal position within the Canterbury Basin), Majungaella sp. 4978 preferred a deep and cool outer shelf habitat (Pukehou locality, more exposed coast swept by cold currents and in an outer shelf “bathyal slope” environment relative to the East Coast Basin). The depositional environment during the middleelate Campanian in southern James Ross Island, Antarctica corresponded to a shallow platform of normal marine salinity and relatively warm water temperatures (Fauth et al., 2002, 2003). The occurrence of two species of Majungaella (nearly 14% of the total fauna) is outnumbered by Mandelstamia antarctica Fauth and Seeling (which represents 48% of the total individuals recorded) and Rostrocytheridea hamiltonensis Fauth and Seeling (nearly 22%). A relatively warm and humid climate with terrestrial floras of a high rainfall forest, “ever wet”, have been also corroborated by

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Fig. 5. Records of valid species of Majungaella in stratigraphical order, including the new records for Argentina and Brazil.

geochemical evidence (Dingle and Lavelle, 1998). This is in broad agreement with the results of Hayes et al. (2006) whose palaeoclimatic analysis based on angiosperm leaves indicated that the mean annual temperature for the Santa Marta Formation was 15e23  C (mean 19  C). The floras are indicative of warm climate

without extended periods of winter temperatures below freezing and with adequate moisture for growth. Majungaella adapted to the cool (pre-glacial) environments in the James Ross Basin by the end of the Eocene (Szczechura, 2001) and full-blown glacial waters by Oligocene in the Victoria Land

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Basin (Dingle and Majoran, 2001). In this case, the presence of Austrocythere reticulotuberculata Hartmann, which today is found only north of 70 S (Hartmann, 1997), suggests water temperatures somewhat higher than the present Ross Sea, but possibly lower than those of modern southern South America. The fauna from the Pliocene interglacial strata on Cockburn Island suggests an environment similar to that presently existing in Antartica (Szczechura and Blaszyk, 1996). Majungaella sp. co-existed with Copytus caligula Skogsberg, Pseudocythereis spinifera Skogsberg, Australicythere polylyca (G.W. Müller) and Antarctiloxoncha frigida (Neale), which live on the modern Antarctic shelf at depths between 60 and 200 m and at a temperature lower than 2  C (Hartmann, 1990). Concheyro et al. (2007) reached similar conclusions based on the association of Majungaella sp. with Ammoelphidiella antarctica, an exclusive Pliocene Antarctic foraminifer from the James Ross Basin.

6. Discussion 6.1. Origin and migration Majungaella was a medium to very large marine Mesozoic and Cenozoic podocopid ostracod that was incapable of free swimming and also lacked pelagic larvae. Therefore, their active migration is confined to walking distance or transport by marine agents during their lifetime. Ostracods can migrate latitudinally when temperature and other ecological parameters, such as the bathymetry, remain stable within certain limits and continental margins can function as migration pathways (Babinot and Colin, 1992). From all records of Majungaella, from the Middle Jurassic to the Pliocene (Fig. 5), and their plots on palaeogeographical maps (Fig. 6), interesting features are revealed.

Fig. 6. Palaeogeographical reconstructions with occurrences and distribution of Majungaella. Maps adapted from Smith et al. (1994), illustrating the ages: A, Callovian, 160 Ma; B, Tithonian, 148 Ma; C, HauterivianeBarremian, 130 Ma; D, Albian, 105 Ma; E, Santonian, 85 Ma; F, Maastrichtian, 70 Ma. Ar, Argentina; An, Antartica; Au, Australia; Br, Brazil; I, India; Is, Israel; M, Madagascar; NZ, New Zealand; Saf, South Africa; Som, Somalia; Tz, Tanzania.

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All reliable Middle Jurassic records of Majungaella are from India, Madagascar and East Africa (Fig. 6A). The earliest, almost simultaneous, records seem to be M. biswasi Neale and Singh (probably ¼ M. perforata) from the BathonianeTithonian in India, and M. mundula (Grekoff) from the lower Callovian in southwest Madagascar. Fifteen species of Majungaella have been recognized in the Middle and Late Jurassic, M. mundula and M. perforata being the most abundant in the three regions. The Callovian faunas of Madagascar have several species in common with those of India (see Fig. 5), and therefore indicate intensive faunal exchange of ostracods between these two regions. This affinity suggests that the Tethyan Gulf, which covered the Kutch-Rajasthan region, extended to Madagascar after skirting the east coast of Africa (Bhalla, 1983). The distribution pattern is in accordance with current palaeogeographic models, which suggest that Madagascar and India were connected by a shallow marine shelf during Middle and Late Jurassic times (Mette, 2004b). In Australia, its first arrival (as suggested by Guzel, in press) could have been by the Oxfordian (Fig. 6B). An effective exchange between Madagascar and Australia was already known for the Middle Jurassic, as demonstrated by the contemporaneous occurrence of Paradoxorhyncha Chapman in the lower Bajocian of Western Australia and the Bajocian of southwest Madagascar (Mette and Geiger, 2004a). Considering the absence of Paradoxorhyncha in the North and East Africa, Arabia and the Near East, Mette (2004a, b) postulated an early Bajocian shallow marine route from Western Australia to Madagascar (via the incipient formation of the “Gulf Madagascar”). The coeval occurrence of Procytherura aerodynamica Bate, 1975 in the Kimmeridgian of Tanzania and in the Kimmeridgian of DSDP site 263 off Western Australia (as Indet. sp. E of Oertli, 1974, p. 964, pl. 7, fig. 5), also support this marine migration route between the Indian Ocean and the southeast Pacific. Suitable neritic habitats along the margins of Gondwana have been demonstrated by Damborenea and Manceñido (1992) by direct comparison of Jurassic marine invertebrates from equivalent facies settings in New Zealand and the Argentinian-Chilean basins. In South Africa, Majungaella is both abundant and diverse in “Neocomian” deposits (Fig. 6C). The genus remains an abundant taxon in Madagascar through the Neocomian with M. nematis (ValanginianeHauterivian), but this species does not survive into Aptian or post-Aptian strata. It migrated northwards to western Israel (Fig. 6C). Guzel (in press) has suggested that the cool climate of the late Valanginianeearly Hauterivian with a substantial amount of polar ice (as postulated by McArthur et al., 2007), resulted in the Gondwanan genera Majungaella and Arculicythere extending their range into the South Tethys Province. Leanza (1996) used the strong similarities of the ammonoid successional pattern between Argentina and Iraq to propose a migration route via the Mozambique Channel, Somaliland, eastern India (Spiti), Madagascar and eastern Antarctica to southern Gondwana for faunal interchange during the JurassiceCretaceous transition. This seaway would have facilitated the migration of Majungaella to western Israel, which is consistent with Rosenfeld and Raab’s (1984) ValanginianeHauterivian record there of M. cf. perforata. The last appearance is in the lowermost upper Albian (M. pyriformis) in northern Madagascar (Grekoff in Collignon et al., 1979; Babinot et al., 2009; Fig. 6D). Sandstones and shales of freshwater origin with a rich Ptilophyllum flora are widely distributed in “Neocomian” strata of Peninsular India (Bhalla, 1983) and therefore records of Majungaella have not been recognized. The last appearance of a representative of the genus is Majungaella manheratibbaensis Singh in coincidence with the widespread Cenomanian transgression (Fig. 6E). In southwestern Africa, Majungaella survived into the Santonian (Majungaella sp. K9A3 of Dingle, 1996), before becoming extinct there also (Fig. 6E).

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During the latest JurassiceBerriasian the opening of a shallow intermittent epicontinental seaway between southern South Africa and southern Argentinian Patagonia favoured faunal interchange. This seaway corresponds to the commencement of the continental separation between Africa and South America (124 Ma), which created the South Atlantic and a small ocean basin off south-east Africa (Natal Valley, see Dingle, 1988) producing a Valanginiane Hauterivian influx of new species and an increase in population diversity. The presence of Majungaella (M. pavta) and coeval species of Rostrocytheridea and Sondagella in the Neuquén Basin and South Africa (cf. Ballent, 2009) as well as the coeval records of M. hemigymnae, M. praehemymnae and M. nematis in South Africa and the Austral Basin corroborate close links between these two regions, thus supporting the existence of a southern Gondwana seaway (Ballent and Whatley, 2007; Ballent, 2009). The same route is known to have been adopted by foraminifers (Kielbowicz et al., 1983; Caramés, 2011). The migration of species, as shown by Dingle (1999), occurred along the southern flanks of the Walvis Ridge, an eastewest orientated barrier that separated in the newly opening South Atlantic, marine (and temperate) conditions in the south from a non-marine environment to the north. During the AptianeAlbian, these NeocomianeBarremian non-marine sediments were abruptly overlain by paralic/marine strata with marine sediments (Dingle, 1999). On both sides of the proto-Atlantic, uppermost AptianeAlbian deposits consist of alternating nearshore marine/ lagoonal facies that pass upwards into more open-water marine sediments (lateral equivalents of the Madiela Formation in Gabon and the Riachuelo Formation in Sergipe; see Fig. 6D). Similar facies/ ostracod associations were suggested by Krömmelbein (1967), Grosdidier (1979) and most recently by Do Carmo et al. (in press) with the description of the same species of the paralic ostracod Sergipella on both sides of the Atlantic. Consistently with the progressive opening of the South Atlantic Ocean from south to north, its two margins developed under different tectonic processes. On the eastern side, the early break-up phase of the South Atlantic began around 130 Ma when the West African rift system developed, resulting in differential movement between western and southern Africa (Fairhead and Binks, 1991). On the western side, along the eastern margin of South America, geodynamic evolution accompanied Andean deformation and the Late Cretaceous was a time of relative tectonic quiescence, recording the maximum flooding of the continent by Atlantic waters (Uliana and Biddle, 1988). Majungaella, already installed in southern Argentina, seems to have migrated to Brazil through the suitable shallow continental shelves (Fig. 6E, F). During the Aptian, sedimentation in Brazilian marginal basins was predominantly continental and the faunas are mainly non-marine (e.g., Swain, 1946; Wicher, 1959; Krömmelbein, 1961, 1962; Krömmelbein and Weber, 1985, Moura, 1988; Do Carmo et al., 1999; Dias, 2005). In the Albian, with the establishment of marine conditions, the marine fauna of ostracods became more abundant and diverse. The presence of Majungaella is characteristic of the marine intervals studied, but despite their wide distribution through the CampanianeMaastrichtian interval, numbers of its species are relatively low. The distribution of the genus along the Brazilian margin is uneven. In the Pelotas Basin, it is recorded from Maastrichtian beds (Majungaella sp. of Ceolin et al., 2011). From south to north the other recorded species are, Majungaella sp. 1, Majungaella sp. 2 and Majungaella. sp. 1 of Miller et al. (2002) from the Santonian and early Campanian of the Santos Basin; M. santosensis sp. nov. from the CampanianeMaastrichtian of the Santos and Espírito Santo basins; M. alta sp. nov., from the CampanianeMaastrichtian of the Santos Basin; Majungaella sp. 5 from the CampanianeMaastrichtian of the Sergipe Basin. Rostrocytheridea, other gondwanan ostracod, seems have followed the

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same way, as demonstrated by its record from the early Maastrichtian of the Pará-Maranhão Basin (Piovesan et al., 2009, pl. 3, fig. 12). On the eastern side of the South Atlantic, in southwestern Africa, Majungaella survives into the Santonian where it is represented by Majungaella sp. K9A3 of Dingle (1996) (Fig. 6E) in sequences that accumulated in a deep-marine and presumably cool-water environment (McMillan, 1990). In the “mid” and Late Cretaceous the affinities between the ostracod faunas of the West African and Brazilian margins are discussed by Viviers et al. (2000), who report the genera Amphicytherura, “Conchoecia” and Veenia from the Albian and Brachycythere, Nigeroloxoconcha and Haugthonileberis from the CenomanianeTuronian as elements common to both regions. In the ConiacianeMaastrichtian interval this similarity is represented by the genera Brachycythere, Cophinia, Ovocytheridea and Protobuntonia. Despite these similarities, Majungaella has not been recorded from Late CretaceousePaleogene deposits along the western continental margin of Africa (see Apostolescu, 1961, 1963; Neufville, 1973; Tambareau, 1982; Sarr, 1999; Elewa, 2002; Fauth, 2002 and references therein). An explanation for this absence requires additional research but it is possibly an artefact of insufficient sampling in West Africa. During mid-Cretaceous times there would have been no impediment to communication between Western Australia and Patagonia, as demonstrated by the presence of Majungaella santacruziana in the Albian of both regions. This connection was favoured by an eastward flow of water in a belt of westerly winds owing to the presence of a West Wind Drift type cool current comparable to that coming today from Australasia (Gordon, 1973). In Western Australia, Majungaella continued to be present in the Late Cretaceous with M. annula in the SantonianeCampanian of the Carnarvon Basin and Santonian in the Gingin Basin (Fig. 6E, F). Dingle (2009) demonstrated comparisons and new correlations between the Late Cretaceous faunas of Australasia, Antarctica and Patagonia. He described three new species of Majungaella from the uppermost Maastrichtian of New Zealand. The records of Majungaella in Antarctica (Majungaella sp. of Ballent et al., 1998 and Majungaella pseudonymos of Rossi de García and Proserpio, 1980) from the middleeupper Campanian of southern James Ross Island (Fig. 6F) and in the Upper Cretaceous of Patagonia show strong faunal connections with both southern South America and the Antarctic Peninsula. Also, the occurrence of Rostrocytheridea hamiltonensis Fauth and Seeling (close to R. canaliculata Bate in Western Australia) in Antarctica and of Rostrocytheridea? sp. of Ballent and Whatley (2007), which is close to males of R. westraliensis (Chapman) from the Santonian of Western Australia, in southern Patagonia, reinforces such links. These links support the view that the Ross Sea area was connected faunally to southeastern Australia, presumably via the Tasman shelf/rise. They are also in agreement with the assumption that the Drake Passage (now ca. 1000 km wide) started to open at about 36 Ma (or slightly earlier, as suggested by Ghiglione et al., 2008), so that, prior to that time, the distance between Antarctica and Patagonia would have been shorter (Lawver et al., 1992; Lawver and Gahagan, 2003). Despite the fact that an Oligocene palaeobiogeographic seaway between New Zealand and Patagonia along the West Antarctic Rift system based on similar invertebrate faunas has been postulated by Casadío et al. (2010), there are no Palaeogene records of Majungaella in the continental part of southern South America (Argentina and Chile). The genus apparently became extinct in this area during the Maastrichtian. The separation of South America from Antarctica and the subsequent formation of the Drake Passage influenced Cenozoic global cooling because these events enabled the development of the Antarctic Circumpolar Current (Lawver and Gahagan, 2003). The widening of the Drake Passage allowed a suite of newly

originated cryophilic genera to evolve there on the continental shelf during the Neogene (Wood et al., 1999). Frigid (glacial) temperatures have been indicated for Oligocene and younger ages in northern Antarctica (Dingle and Lavelle, 1998). The distribution of the flora (Gandolfo et al., 1998) and benthonic foraminifers (Malumián and Náñez, 1991) reinforce the existence of a latitudinal gradient between both continents and a progressive biotic migration towards the north as a consequence of climatic deterioration during the EoceneeOligocene. Based on marine invertebrates from Seymour Island, Zinsmeister and Feldman (1994) stated that highlatitude regions serve as “holding tanks” for taxa that remain isolated until they disperse from the region. It is highly likely, therefore, that the Cenozoic Antarctic populations of Majungaella (Fig. 6F) recorded by Szczechura (2001) as M. antarctica, by Szczechura and Blaszyk (1996) as Majungaella sp. and by Dingle and Majoran (2001) as Majungaella sp. 4471, derive from this already cryophyllically adapted root stock. The genus survived for a further 30 Ma under glacial/interglacial conditions and constitutes an example of a taxon for which high-latitude cold climates became a “refugia”, but for which the Pleistocene climate of Antarctica eventually proved to be too severe (Whatley et al., 2005). This propensity of migration from Cretaceous warm, shallow water to Cenozoic deeper or cold waters observed in certain genera has been considered a retrothermal adaptation (Dingle, 2009). 6.2. Control on migration patterns Investigations on the control of global ostracod distribution, suggest that, apart from physical barriers, such as tectonic plate motions (Smith, 1986; Benson, 1988; Whatley, 1988), the main abiotic criteria that affect shallow marine benthonic ostracod migrations are temperature, light penetration, pressure (which are water depth dependent) and ocean currents (Titterton and Whatley, 1988; Whatley, 1988; Dingle, 2002, among others). The thermospheric nature of the Jurassic oceans favoured the migration of benthonic ostracods, which could cross oceanic basins such as the Tethys because the marine bathymetric temperature gradient was not a barrier for shore to shore migration (Whatley, 1988; Whatley and Ballent, 1994). Considering that during Jurassic times the vertical oceanic circulation of water masses was of low intensity because of thermospheric conditions, there must have been a significant vertical gradient in dissolved oxygen. Low oxygen concentrations in areas of deeper shelf and basin would have been an effective migration barrier for shallow marine ostracods, with the exceptions of Metacopina and Platycopina (Whatley, 1991; Horne et al., 2011). Donze (1977) has previously demonstrated how the Berriasian shelf-dwelling ostracod faunas of France and Spain differ importantly from those of North Africa because of the presence of the oceanic Tethys, the depth of which was then an insuperable barrier for migration of benthonic ostracods. A similar faunal differentiation has been documented for the Late Cretaceous faunas of the northern and southern Tethys margin (Babinot, 1985), which suggests that migration was at least impeded by the depth central Tethys basin. During Middle and Late Jurassic times, almost the entire KutchRajasthan region of western India was covered by a gulf that was evidently in direct communication with the Tethys in the north; these Jurassic rocks form the first significant record of fossiliferous marine deposits on the Indian Peninsula (Bhalla, 1983). The peripheral geographic position of this shallow marine arm of the southern Tethys was probably the main prerequisite for the origination of several endemic ostracod taxa (Mette, 2004b). At this time the opening of the “Gulf of Madagascar” began and surely led to the influx of warm equatorial Tethyan water masses into this

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incipient way (Mette, 2004a, b), contributing to the migration of Majungaella during the Middle Jurassic and the coeval records of several species in Madagascar and India (Fig. 5). Only two common species (M. mundula and M. perforata) are recorded from the CallovianeUpper Jurassic of south-western Madagascar and Tanzania, showing an increasing endemism in the Indo-East Africa region. The successive widening and deepening of the “Gulf of Madagascar” because of sea floor spreading between East Africa and India/Madagascar, and the subsequent reduced oxygen concentration within the deeper shelf and basin areas, acted as an effective barrier to migration of the genus (Mette, 2004a). This is in accordance with the assumption of Bate (1975), who attributed the reduced diversity of the Kimmeridgian ostracod fauna of Tanzania to a probable lowering of the water temperature. The initial history of all Cretaceous sedimentary basins of southern South America is related to the initial break-up of Gondwana. TriassiceLate Jurassic intracontinental rifting spreading and subsidence culminated in the earliest Cretaceous with the opening of the South Atlantic (Riccardi, 1987). During the Jurassic the geodynamic evolution of the Neuquén Basin in central western Argentina was intimately linked to the development of the central Andes. The basin originated in the Late Triassic as a result of continental intraplate extension. Jurassice Early Cretaceous sedimentation was coeval with progressively more widespread marine invasions from the northwest and west, leading to an elongate marine seaway which connected central Patagonia with the Pacific domain (Legarreta and Uliana, 1996; Howell et al., 2005). This geodynamic scenario favoured the development of shallow marine-margin environments that facilitated the colonization and migration of benthonic ostracods. Palaeomagnetic data (Iglesia Llanos, 2008) show that, by the Late Jurassic, the position of the Neuquén Basin was similar to that of the present day (ca. 30 S). Abundant remains of marine crocodiles in mid-Late Jurassic deposits indicate warm water temperatures, probably in excess of 20  C (Volkheimer et al., 2008). This is in agreement with temperatures of nearly 25  C during the Oxfordian (Matheos, 1992) and between 22 and 27  C as suggested by Gasparini et al. (2002) from the d18O values for oysters and ammonite septa. Warm temperatures are also in accordance with the development of epifaunal small oyster and serpulid “boundstones” in the Valanginian (Schwarz, 1999) and with the data of Lazo et al. (2008) who concluded that a temperature of 25  C was supported by records of coral assemblages, coral patch-reef facies, oolithic carbonates, and thick-shelled bivalves in the ValanginianeHauterivian Agrio Formation. However, Majungaella is represented by only one species (the late TithonianeBerriasain and Valanginian M. pavta) despite the favourable warm temperatures mentioned. Coinciding with the commencement of the continental separation between Africa and South America (124 Ma), which created the South Atlantic and a small ocean basin off southeast Africa (Natal Valley; see Dingle, 1988) a ValanginianeHauterivian influx of new species and an increase in population diversity occurred. This is demostrated by the coeval records of Majungaella during this time in South Africa and the Austral Basin. A more widespread entrance of surface waters in the South Atlantic was facilitated in the late Aptian by deepening of the Malvinas Plateau, where an outermost neritic to bathyal environment developed (Riccardi, 1991). Also, some palaeoceanographical and palaeogeographical alterations, such as the opening pole change at 105 Ma (early Albian; see Dingle, 1988, table 3), probably led to deep-water passages forming along the line of the Falkland Plateau-southern Africa fracture zones, allowing the dysaerobic temperate South Atlantic to be flushed by oxygenated waters, which favoured the appearance of new taxa. This was relatively short-lived and coincided with the

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rapid distribution of M. santacruziana and Arculicythere tumida (Ballent, 1998; Ballent and Whatley, 2006), with Albian records connecting Australia and southern Patagonia. The latter species seems to have tolerated depths of the order of 200e600 m (external shelfeupper slope) and, therefore, cooler water in offshore Western Australia and southern Argentina during the Albian. This is also supported by data on the oxygen isotope geochemistry of dimitobelid belemnites referred to by Pirrie et al. (2004), which suggest relatively cool temperatures (mean 9.5  C) for the early Albian of Argentina at a palaeolatitude of approximately 58 S. This value was compared with previous data on dimitobelid belemnites (the group was particularly shelf-dwelling nektobenthonic) from Antarctica (James Ross Basin, mean 8.5  C) and Australia (Carnarvon Basin, mean 10.1  C). Together these data suggest that shelf temperatures around the Gondwanan margin during the early Albian were relatively cool, but probably not cold enough to indicate the presence of significant polar glaciation at this time. The preference of Arculicythere for cool waters during the Albian could have caused its extintion (Ballent and Whatley, 2006) as indicated by its absence in the Albian of the mid-South Atlantic where the waters were warmer than those of the Falkland Plateau (cf. Azevedo, 2004). In contrast, these “warmer waters” favoured the migration of Majungaella, as indicated by its records along the Brazilian margin. According to Maisey (2000) the communication between Equatorial and Temperate South Atlantic oceans was fully developed by the late Aptian, and the mid-late Albian dispersal of anoxic and hypersaline conditions north to the Walvis RidgeeRio Grande Rise barrier thus postdates the opening of the South Atlantic and may instead be related to its deepening. The existence of the Walvis Ridge barrier and the ubiquitous oxygen depletion in the South Atlantic during the mid-Cretaceous probably acted as a barrier and contributed to the isolation of its benthonic faunas (Delicio et al., 2000). During the Early Cretaceous, the Sergipe Basin in northeastern Brazil lay towards the northern end of the South Atlantic. It has provided crucial evidence for the timing of the establishment of the equatorial seaway linking the North and South Atlantic oceans. The influx of foraminifers in this basin documented by Koutsoukos and Hart (1990) signalled the establishment of the first permanent shallow-water equatorial seaway and was probably related to the alleviation of anoxic and hypersaline conditions in the northern part of the South Atlantic (Maisey, 2000). This situation also favoured the migration of Majungaella and its record from the CampanianeMaastrichtian of the Sergipe Basin constitutes the last known in Brazil. There are no Late Cretaceous records of Majungaella in the most northern basins (Pernambuco, Pará-Maranhão), which show strong affinities with faunas from northwest Africa, the Caribbean and North America (Fauth et al., 2005; Piovesan et al., 2009 and references therein). Probably the evolving retrothermal tendency (see below) together with some degree of competitive exclusion with respect to the “warmer water faunas” could explain the extinction the Majungaella along the Brazilian continental margin. 6.3. Retrothermal tendency of Majungaella The retrothermal propensity of Majungaella is well known because the genus survived and colonized much of the Antarctic seaboard by Eocene, Oligocene and Pliocene. According to Dingle (2009), the genus had developed a retrothermal disposition at least by the latest Maastrichtian as indicated by Majungaella sp. 4978, which occurs sparsely deposits representing the cool, outer shelf environment at Pukehou, New Zealand. It is possible that this tendency may have developed even by mid-Late Cretaceous times in Majungaella p. K9A3, which Dingle (1996) recorded in

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AlbianeSantonian continental slope (and presumably cool-water) sediments off southern Namibia (McMillan, 1990) and also in Majungaella santacruziana which dominates the Albian fauna of the Austral Basin in Argentina in a cool-temperate outer shelf and upper slope environment (Ballent, 1998). Speculating a bit more, in the lower Kimmeridgian of Tanzania, the abundance of Majungaella kimmeridgiana within a fauna of reduced diversity attributed by Bate (1975) to a probable cooling tendency may have predicted the retrothermal temdency later verified in the genus. A similar pattern has been recently proposed for Pelecocythere Athersuch from the Upper Cretaceous of the Santos Basin by Piovesan et al. (2010). 7. Summary and conclusions The record of new species of Majungaella in Argentina and Brazil have contributed to the understanding of the migration routes of the genus and the evolution of the South Atlantic during the Late Cretaceous. The first records of the genus are from India and Madagascar, in the Middle Jurassic. It spread to East Africa and South America during the Late Jurassic. In the Early Cretaceous, it was also present in Australia and Israel. In the Late Cretaceous it was in New Zealand and Antarctica and widely distributed on the Brazilian continental margin. In the Cenozoic, records of the genus are restricted to Antarctica, where it has been documented up to the Pliocene. With respect to palaeoenvironmental preferences, Jurassic and early Cretaceous species seemed to inhabit mainly warm shallowwater environments. From the mid-Late Cretaceous and especially through the Cenozoic, the genus apparently had a retrothermal propensity. Our paper comprises a further contribution to the taxonomy, palaeozoogeography and evolution of Majungaella along the southern margin of Gondwanaland. Acknowledgements EKP and GF thank Marta Claudia Viviers (Petrobras-Cenpes/ DIVEX) for access to some of the material examined and Rogério Martins (Petrobras-Cenpes/DIVEX) for the SEM work. We also thank Cristianini Trescastro Bergue, Daiane Ceolin and Gabriel Henneman Klaser of the Laboratório de Micropaleontologia of the Universidade do Vale do Rio dos Sinos (UNISINOS) for support during the preparation of the paper. SB (now sadly deceased) thanks the Universidad Nacional de La Plata (UNLP) of Argentina and the Universidad do Vale do Rio dos Sinos (Brazil) for an international cooperation grant which enabled her to visit São Leopoldo in order to carry out this and other research. Funding for this study was provided by Conicet (PIP 0819) and UNLP (N/589). References Andreu, B., Colin, J.-P., Singh, J., 2007. Cretaceous (Albian to Coniacian) ostracodes from the subsurface of the Jaisalmer Basin, Rajasthan, India. Micropaleontology 53, 345e370. Apostolescu, V., 1961. Contribution a l’étude paléontologique (Ostracodes) et de stratigraphique des bassins crétacés et tertiaires de l’Afrique Occidentale. Revue l’Institut Français de Pétrole 16, 779e867. Apostolescu, V., 1963. Essai de zonation par les Ostracodes dans le Crétacé du Bassin du Sénégal. Révue de l’Institut Français de Pétrole 18, 1675e1694. Azevedo, R.L.M., 2004. Paleoceanografia e a evolução do Atlântico sul no Albiano. Boletim de Geociências da Petrobras 12, 231e249. Babinot, J.F., 1985. Paléobiogeographie des ostracodes du Crétacé supérieur des marges ouest-européennes et nord- africaines de la Tethys. Bulletin de la Societé Géologique de France 8, 739e745. Babinot, J.F., Colin, J.-P., 1992. Marine ostracode provincialism in the late Cretaceous of the Tethyan realm and the austral province. Palaeogeography, Palaeoclimatology, Palaeoecology 233, 63e95. Babinot, J.F., Colin, J.-P., Randrianasolo, A., 2009. Les ostracodes de l’Albiene Turonien moyen de la région d’Antsiranana (Nord Madagascar):

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