Verónica V Vennari
email: vvennari@mendoza-conicet.gob.ar

Beatriz Aguirre-Urreta

Luciana S Marin
email: lmarin@gl.fcen.uba.ar

Pierre Pellenard
email: pierre.pellenard@u-bourgogne.fr

Mathieu Martínez
email: mathieu.martinez@univ-rennes1.fr

Maisa Tunik
email: mtunik@unrn.edu.ar

Mathieu Martínez

Journal Pre-proof Upper Jurassic (TITHONIAN) belemnites from the Neuquén basin, Argentina

Keywords: Fossil coleoidea, Belemnopsis, Hibolithes, Vaca Muerta Formation, Trans-Gondwana seaway

Belemnites were a group of Mesozoic coleoids with a global distribution from the Early Jurassic to the Late Cretaceous. During the Late Jurassic-Early Cretaceous, Gondwana's marine shelves were dominated by Belemnopsis and Hibolithes populations. In contrast to the Austral Basin, where both genera are quite common, their record in the Neuquén Basin (Argentina) is rather scarce and poor, probably due to a combination of locally unfavorable environmental conditions and predation pressure. Here we describe a new belemnite association from the Tithonian of the Vaca Muerta Formation from southern Mendoza and northern Neuquén provinces. Belemnite's findings are assignable to the Tithonian Windhauseniceras internispinosum and Corongoceras alternans Andean ammonoid Biozones. The studied rostra are identified as Belemnopsis cf. B. patagoniensis, Belemnopsis cf. B. launceloti, and Hibolithes argentinus? Affinities of the Neuquén Basin belemnite fauna are referred to the Madagascan-Eastern Africa associations (Ethiopian Subprovince, Tethyan Province), adding evidence to an established Trans-Gondwana seaway enabling the connection between those areas from the Tithonian onwards.

Introduction

Belemnites are an extinct group of Mesozoic nektonic cephalopods considered in the most recent contributions as stem-dechabrachian coleoids [START_REF] J O U R N A L P R E -P R O O F Fuchs | Longibelus gen. nov., a new Cretaceous coleoid genus linking Belemnoidea and early Decabrachia[END_REF][START_REF] Hoffmann | The palaeobiology of belemnites -foundation for the interpretation of rostrum geochemistry[END_REF]. Usually reconstructed as externally morphologically close to loginid squids [START_REF] Naef | Die fossilen Tintenfische[END_REF][START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF], belemnites had a mantle muscle structure similar to that of sepiids [START_REF] Monks | The function of the belemnite guard[END_REF]. Their mineralized inner shell was generally divided into three parts: proostracum, phragmocone, and rostrum. Of these, the low-Mg calcitic rostrum is the most commonly preserved part due to its resistant structure of radially arranged calcite fibers that, combined with variable amounts of organic matter, form concentric rings in cross-section [START_REF] Stolley | Beiträge zur Kenntnis der Cephalopoden der Norddeutschen Unteren Kreide 1: Die Belemnitiden der Norddeutschen Unteren Kreide[END_REF]Jeletzki, 1966;[START_REF] Riegraf | Revision der Belemniten der Schwäbischen Jura. Part 7[END_REF][START_REF] Hoffmann | Evidence for a composite organicinorganic fabric of belemnite rostra: implications for palaeoceanography and palaeoecology[END_REF][START_REF] Hoffmann | The palaeobiology of belemnites -foundation for the interpretation of rostrum geochemistry[END_REF]. Belemnite taxonomy mostly relies on morphological characters of the rostrum: general morphology, both in outline and profile view, shape of the transverse section, form and orientation of the apex, and presence and extension of grooves (apical, ventral, dorsal, and/or lateral). The orientation and form of the apical line and alveolus-related characters (i.e., where the phragmocone lies inside the rostrum) are usually accessorily described [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Jeletzky | Macroinvertebrate paleontology, biochronology, and paleoenvironments of Lower Cretaceous and Upper Jurassic rocks, Deep Sea Drilling Project Hole 511, Eastern Falkland Plateau[END_REF][START_REF] Doyle | The Jurassic and Cretaceous belemnites of Kong Karls land, Svalbard[END_REF][START_REF] Challinor | Belemnite biostratigraphy of the New Zealand Late Jurassic Mangaoran (Early Puaroan) Substage and the Puaroan Stage revisited[END_REF].

Belemnites are recorded from the Late Triassic [START_REF] Iba | The early evolutionary history of belemnites: new data from Japan[END_REF], but it is from the Toarcian onwards that they attain a worldwide distribution [START_REF] Doyle | Lower Jurassic-Lower Cretaceous belemnite biogeography and the origin of the Mesozoic Boreal Realm[END_REF]. By the Late Jurassic-Early Cretaceous Gondwana shelf-areas were dominated by Belemnopsis and Hibolithes [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Doyle | Belemnite distribution patterns[END_REF], both included in the "Belemnopseina," a group characterized by alveolar furrows and whose taxonomic validity has been recently challenged (Stevens, in press). Hibolithes was a cosmopolitan and ecologically wide-ranged genus whose species are mostly differentiated based on the ventral groove characters [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] J O U R N A L P R E -P R O O F Mutterlose | Migration and evolution patterns in Upper Jurassic and Lower Cretaceous belemnites[END_REF][START_REF] Doyle | The Jurassic and Cretaceous belemnites of Kong Karls land, Svalbard[END_REF] J o u r n a l P r e -p r o o f [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF]. Belemnopsis is a more variable genus [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Combémorel | Les bélemnites de Madagascar[END_REF][START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF][START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Challinor | Stratigraphy of Tithonian (Ohauan-Puaroan) marine beds near Port Waikato, New Zealand, and a redescription of Belemnopsis aucklandica (Hochstetter)[END_REF][START_REF] Challinor | Synonymy and stratigraphic ranges of Belemnopsis in the Heterian and Ohauan Stages (Callovian-Tithonian), southwest Auckland, New Zealand[END_REF][START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF] mostly represented in Gondwana by robust, cylindroconical to slightly depressed rostra with a variably strong ventral groove [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF][START_REF] Doyle | Jurassic belemnite distribution patterns: implications of new data from Antarctica and Argentina[END_REF][START_REF] Doyle | Belemnite distribution patterns[END_REF]. In Argentina, both Hibolithes and Belemnopsis had been long recognized from the Upper Jurassic and, mostly, from the Lower Cretaceous of the Austral Basin in south Patagonia (e.g., [START_REF] Dana | Geology[END_REF][START_REF] Favre | Die Ammoniten der unteres Kreide Patagonien[END_REF][START_REF] Feruglio | Paleontographica Patagónica[END_REF][START_REF] Feruglio | Geología de la Patagonia[END_REF] and have been revised or cited in the literature from there on (e.g., [START_REF] Aguirre-Urreta | Belemnites de una secuencia turbidítica volcanoclástica de la Formación Yaghán, Titoniano-Cretácico Inferior del extremo sur de Chile. 4° Congreso Geológico Chileno[END_REF][START_REF] Riccardi | Paleontología y edad de la Formación Springhill. 1° Congreso Geológico Chileno[END_REF][START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF][START_REF] Riccardi | The Cretaceous System of southern South America[END_REF]. In contrast, Neuquén Basin belemnite records of the same age are significantly less frequent, and taxonomic-focused contributions are lacking [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF]1997).

Therefore, the objective of this work is to fully document new belemnite rostra from the Upper Jurassic (Tithonian) of the Neuquén Basin to start filling the gap in the knowledge of this group in west-central Argentina. We also aim to refer the findings to the Andean ammonoid-based biostratigraphic framework, and to discuss the paleobiogeographic, biostratigraphic, and broad paleoecological implications of their occurrences.

Previous Neocomian belemnite records in Argentina and nearby regions

Upper Jurassic-Lower Cretaceous belemnite records from Argentina are currently concentrated in the Austral Basin (Fig. 1). B. patagoniensis (Favre) was originally described from the Hauterivian-Barremian of Lago Belgrano area and later restricted to the Hauterivian [START_REF] Favre | Die Ammoniten der unteres Kreide Patagonien[END_REF][START_REF] Riccardi | The Cretaceous System of southern South America[END_REF]. Findings around Lago San Martín, at the top of the Springhill Formation, were alternatively ascribed to the Tithonian (Bonarelli and Nágera, 1921;[START_REF] Feruglio | Geología de la Patagonia[END_REF], the Hauterivian-Barremian [START_REF] Riccardi | Estratigrafía en el Oriente de la Bahía de La Lancha, Lago San Martín, Santa Cruz, Argentina[END_REF], or the Berriasian [START_REF] Riccardi | Paleontología y edad de la Formación Springhill. 1° Congreso Geológico Chileno[END_REF][START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF]. Finally, they were assigned to a Berriasian-

J o u r n a l P r e -p r o o f

Valanginian age [START_REF] Medina | Desmoceratidae, Silesitidae and Kossmaticeratidae (Ammonitida) from the Upper Aptian-Albian of Patagonia (Argentina)[END_REF][START_REF] Riccardi | The Cretaceous System of southern South America[END_REF]. From the same levels, [START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF] reported B. cf. B. madagascariensis (Besairie) and Hibolites aff. H. jaculum (Phillips). Also from the Lago San Martín area, at the base of the Río Mayer Formation, [START_REF] Medina | Desmoceratidae, Silesitidae and Kossmaticeratidae (Ammonitida) from the Upper Aptian-Albian of Patagonia (Argentina)[END_REF] reported B. patagoniensis, B. cf. gladiatoris Willey, and Belemnopsis sp. together with the ammonoids Favrella americana (Favre) and F. wilckensi (Favre), thus indicating a Hauterivian age for the association. To the south, in the Lago Argentino area, Estancia La Cristina, [START_REF] Feruglio | Paleontographica Patagónica[END_REF] Stevens (1965, p. 159) and alternatively as Parahibolites by Challinor (1991, p. 316). The specimen assigned to B. patagoniensis was reassigned to B. gladiatoris Willey due to morphological differences with the type material [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Willey | Belemnites from South-Eastern Alexander Island. II, The occurrence of the Family Belemnopseidae in the Upper Jurassic and Lower Cretaceous[END_REF] and subsequently to B. launceloti Howlett [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF]. Feruglio (1949, p. 169) also mentioned, but not illustrated, B. patagoniensis from another locality (Cerro de Los Fósiles) within the Lago Argentino area, there associated with Tithonian to Berriasian ammonoid taxa (cf. [START_REF] Leanza | Anotaciones sobre los fósiles Jurásico-Cretácicos de Patagonia Austral (Colección Feruglio) conservados en la Universidad de Bologna[END_REF]. A reassessment of the Indo-Pacific affinity of the whole assemblage [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] as well as a revision of the ammonoid-related fauna, pointed to an upper Tithonian age for Feruglio's original belemnite collection [START_REF] Leanza | Anotaciones sobre los fósiles Jurásico-Cretácicos de Patagonia Austral (Colección Feruglio) conservados en la Universidad de Bologna[END_REF]. The same seems valid for related specimens from the Springhill Formation near Lago Argentino, where Hibolithes ranges into the Berriasian and younger beds of the Río Mayer Formation sensu [START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF].

Other illustrated records of B. patagoniensis or B. cf. patagoniensis come from the southernmost end of South America, from Berriasian-Hauterivian beds of the Yaghán Formation, Bahía Nassau, Tierra del Fuego [START_REF] Dana | Geology[END_REF][START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] J o u r n a l P r e -p r o o f [START_REF] Stolley | Helicerus Dana als Erhaltungszundstand von Belemnopsis[END_REF] and from the Valanginian?-Hauterivian Beauvoir Formation in the western region of the Isla de los Estados [START_REF] Harrington | Observaciones geológicas en la Isla de los Estados[END_REF][START_REF] Caminos | Descripción geológica de la Hoja 67e, Isla de los Estados[END_REF]. Also, [START_REF] Aguirre-Urreta | Belemnites de una secuencia turbidítica volcanoclástica de la Formación Yaghán, Titoniano-Cretácico Inferior del extremo sur de Chile. 4° Congreso Geológico Chileno[END_REF] presented new Berriasian-Hauterivian records of B. madagascariensis from several sections within the Magallanes Archipelago.

Recently, [START_REF] Ippolitov | First find of Parabelemnopsis, the alien belemnite from the Southern Hemisphere, in the Upper Berriasian of Central Crimea[END_REF] described a single specimen from the Berriasian of Crimea as Parabelemnopsis cf. patagoniensis, indicating that its presence was probably the result of a rare immigration episode to the Northern Hemisphere. Aside from the later record, Belemnopsis patagoniensis appears to be a well-distributed species of the Austral Basin during the Lower Cretaceous.

Other records of Belemnopsis in nearby areas are known from the Valanginian-Hauterivian of the Rocas Verdes Basin, Chile, where robust forms were ascribed to the Antarctic species B. cf. B. launceloti Howlett [START_REF] Stinnesbeck | A Lower Cretaceous ichthyosaur graveyard in deep marine slope channel deposits at Torres del Paine National Park, southern Chile[END_REF]. Additional material retrieved from a drill hole at the Falkland/Malvinas Plateau was identified as Belemnopsis spp. and assigned a New Zealand to Himalayan affinity [START_REF] Jeletzky | Macroinvertebrate paleontology, biochronology, and paleoenvironments of Lower Cretaceous and Upper Jurassic rocks, Deep Sea Drilling Project Hole 511, Eastern Falkland Plateau[END_REF].

Within the Neuquén Basin (Fig. 1), Upper Jurassic-Lower Cretaceous belemnite records are sparser and poorly known. Oxfordian Belemnopsis with European affinities have been mentioned from La Manga Formation in southern Neuquén [START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF][START_REF] Riccardi | Megainvertebrados jurásicos y su importancia geobiológica[END_REF] and from the Callovian-Oxfordian of Vega de la Veranada, in northern Neuquén [START_REF] Alberti | Stable isotopes (δ13C, δ18O) and element ratios (Mg/Ca, Sr/Ca) of Jurassic belemnites, bivalves and brachiopods from the Neuquén Basin (Argentina): challenges and opportunities for palaeoenvironmental reconstructions[END_REF]. Younger not-illustrated records from Mendoza Province include an indeterminate rostrum fragment from the Tithonian of Arroyo Serrucho [START_REF] Krantz | La Fauna del Tithono superior y medio en la parte meridional de la provincia de Mendoza[END_REF] and some thin incomplete rostra from the upper Tithonian of the Vaca Muerta Formation in Bardas Blancas [START_REF] Gerth | Contribuciones a la estratigrafía y la paleontología de los Andes Argentinos II. La Fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza[END_REF]; here referred as Cañada de Leiva). Additional fragmentary belemnites were reported from a Berriasian interval in the Arroyo La Manga area [START_REF] Gerth | Contribuciones a la estratigrafía y la paleontología de los Andes Argentinos II. La Fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza[END_REF]. Other Tithonian records from Neuquén Province were either identified as Hibolithes sp. (e.g., Alberti et al., J o u r n a l P r e -p r o o f 2020, from Pampa Tril) or not specified (e.g., Kietzmann and Vennari, 2013, from Cerro Domuyo, here revised). Besides, Stevens (1965, p. 158)

Geological setting

The Neuquén Basin (Fig. 1) is a retro-arc basin situated over the western margin of Gondwana, between 32° to 40° South Latitude, extending over central-western Argentina and central Chile [START_REF] Legarreta | Jurassic-Cretaceous marine oscillations and geometry of back-arc basin fill, central Argentine Andes[END_REF][START_REF] Howell | The Neuquén Basin: A case study in sequence stratigraphy and basin dynamics[END_REF]. Its nearly continuous Mesozoic to lower Cenozoic sedimentary infill, intercalated with volcanic episodes, and the high quality and richness of its fossil record arouse the interest of the international scientific community over the basin. Tithonian successions bearing the belemnite fauna here presented are included in the Vaca Muerta Formation [START_REF] Weaver | Paleontology of the Jurassic and Cretaceous of West Central Argentina[END_REF], a marine geographically extended unit mainly composed by an alternation of shale/marls with a variety of limestone beds deposited over a carbonate ramp to basin setting from the lower Tithonian to the lower Valanginian [START_REF] Kietzmann | Sedimentology and sequence stratigraphy of a Tithonian-Valanginian carbonate ramp (Vaca Muerta Formation): a misunderstood exceptional source J o u r n a l P r e -p r o o f rock in the southern Mendoza area of the Neuquén basin, Argentina[END_REF][START_REF] Legarreta | Jurassic-Cretaceous marine oscillations and geometry of back-arc basin fill, central Argentine Andes[END_REF]. Ammonoids are the most common megascopic fossils of the Vaca Muerta Formation, including both endemic and more extended distributed taxa. Their correlation with Tethyan faunas and with calcareous nanofossils and microfossils bioevents provide the current Upper Jurassic-Lower Cretaceous biostratigraphic scheme for the Neuquén Basin (Leanza, H. et al., 2020 and references therein;[START_REF] Lescano | New Tithonian biostratigraphic data from the J o u r n a l P r e -p r o o f Neuquén Basin, Argentine Andes: a contribution from the Southern Hemisphere[END_REF].

Material and methods

Localities

J o u r n a l P r e -p r o o f

The twenty-one belemnites herein studied come from Tithonian beds of the Vaca Muerta Formation in four localities situated over the northern sector of the Neuquén Basin (Fig. 2). From the northeast to the southwest, they are Cañada de Leiva (fourteen specimens), Las Loicas (one specimen), Las Alcantarillas (one specimen), and Cerro Domuyo (five specimens).

Cañada de Leiva

It is a classic locality encompassing the Upper Jurassic-Lower Cretaceous transition in the Neuquén Basin (35º 48' S, 69º 48' W), 6 km northeast of the Bardas Blancas settlement [START_REF] Dessanti | Descripción geológica de la Hoja 29b, Bardas Blancas, provincia de Mendoza[END_REF][START_REF] Gerth | Contribuciones a la estratigrafía y la paleontología de los Andes Argentinos II. La Fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza[END_REF][START_REF] Gulisano | The Jurassic of the Neuquén Basin. Field Guide, Mendoza Province. Asociación Geológica Argentina, Serie E, 3[END_REF]. Here

the Vaca Muerta Formation spans nearly 240 m and is dominated by tempestite-facies deposited over a middle to proximal outer carbonate ramp setting [START_REF] Kietzmann | Las tempestitas peloidales de la Formación Vaca Muerta (Tithoniano-Valanginiano) en el sector surmendocino de la Cuenca Neuquina, Argentina[END_REF][START_REF] Kietzmann | Astronomical calibration of the Tithonian-Berriasian in the Neuquén Basin, Argentina: A contribution from the Southern Hemisphere to the Geologic Time Scale[END_REF]. Belemnites from this section come from up to 40 cm thick, very consolidated hummocky cross-bedding bioclastic packstone limestones (CLBe) intercalated with laminated marls/siltstone intervals, and a decimetric massive to laterally concretional wackestone limestone with ammonoids, and located about 24.5 m from the base of the logged section in Figure 2. The CLBe sample is a densely packed, poorly sorted, and oriented (Fig. 3) packstone with micrite and organic matter.

The grains are 70% echinoderm fragments ), 10% of bone fragments (Figs. 3.1,3.4), 10% of assorted fragmented bioclasts, probably bivalve shells (Fig. 3.2), bryozoan fragments and small gastropods (Fig. 3.4), and 10% of siliciclastic material such as volcanic lithic fragments, detrital mono and polycristallyne quartz, feldspar and abundant pyrite. Among the echinoderm fragments were differentiated: star-shaped crinoid columnals (Fig. 3.1), cross and longitudinal sections of stems, crinoid arm elements, some with lunate patterns (Figs. 3.1,3.3), and scarce echinoid spines. Most echinoid fragments are highly reworked, replaced and recrystallized, and some of them J o u r n a l P r e -p r o o f have microborings (Fig. 3.4). Most of the bone fragments have evidence of microbial boring activity as well. The matrix is micritic and some pore spaces are filled with authigenic quartz. The belemnite-bearing interval is the same identified by Gerth in 1911and 1913(Gerth, 1925, fig. 10).

FIGURE 2 and FIGURE 3 AROUND HERE

Las Loicas

It is located close to the 145 National Road heading towards the Pehuenche International Pass to Chile, approximately 1 km to the west of the homonymous settlement (35º 47' S, 70º 09' W). The marine sediments of the Vaca Muerta Formation are punctuated by several volcanic layers, some of which have provided precise U/Pb absolute ages [START_REF] Lena | High-precision U-Pb ages in the early Tithonian to early Berriasian and implications for the numerical age of the Jurassic-Cretaceous boundary[END_REF][START_REF] Vennari | New constraints in the Jurassic/Cretaceous boundary in the High Andes using high precision U-Pb data[END_REF]. Belemnites from this section were retrieved from decimetric to metric laminated to massive dark wackestones bearing ammonoid and brachiopod impressions, intercalated with dark laminated marls.

Based on the facies associations interpreted by [START_REF] Kietzmann | Facies analysis and stratigraphy across the Jurassic-Cretaceous boundary in a new basinal Tithonian-Berriasian section of the Vaca Muerta Formation, Las Tapaderas, Southern Mendoza Andes, Argentina[END_REF] in the nearby Las Tapaderas section, this interval is assignable to the transition between basinal to distal outer ramp settings.

Las Alcantarillas

Located about 20 km west of the Las Loicas section, it is well exposed over the right margin of the Arroyo Pehuenche that crosses 145 National Road (35º 57' S, 70º 14' W).

The single belemnite rostrum from this locality comes from a 12 cm thick, dark laminated wackestone limestone showing a thin pyritized band, which is intercalated within an interval dominated by calcareous-nodule bearing dark laminated marls.

Likewise, the previous locality, the facies association indicates a distal outer ramp setting [START_REF] Kietzmann | Facies analysis and stratigraphy across the Jurassic-Cretaceous boundary in a new basinal Tithonian-Berriasian section of the Vaca Muerta Formation, Las Tapaderas, Southern Mendoza Andes, Argentina[END_REF].

Cerro Domuyo

J o u r n a l P r e -p r o o f

This locality is at the foothills of the Cerro Domuyo, at the southern tip of the Cordillera del Viento (36º 41' S, 70º 26' W). Here the Vaca Muerta Formation is exposed over the left margin of the Arroyo Covunco, and all belemnites were recorded from centimetric to decimetric grey to dark laminated wackestones, intercalated between large intervals of laminated shales and marls. Facies association analyses indicate a basinal to distal outer ramp setting for this interval, occurring just below slumped calcareous sandstone deposits assigned to the Huncal Member of the Vaca Muerta Formation by [START_REF] Kietzmann | Sedimentología y estratigrafía de la Formación Vaca Muerta (Tithoniano-Berriasiano) en el área del cerro Domuyo, norte de Neuquén, Argentina[END_REF] and reinterpreted by [START_REF] Naipauer | The Huncal Member of the Vaca Muerta Formation, Neuquén Basin of Argentina: insight into biostratigraphy, structure, U-Pb detrital zircon, ages and provenance[END_REF].

Measurements and illustrations

Belemnite rostra's study and description follow the outlines of [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] and [START_REF] Challinor | Belemnite biostratigraphy of the New Zealand Late Jurassic Mangaoran (Early Puaroan) Substage and the Puaroan Stage revisited[END_REF][START_REF] Challinor | Synonymy and stratigraphic ranges of Belemnopsis in the Heterian and Ohauan Stages (Callovian-Tithonian), southwest Auckland, New Zealand[END_REF]. [START_REF] Challinor | Belemnite biostratigraphy of the New Zealand Late Jurassic Mangaoran (Early Puaroan) Substage and the Puaroan Stage revisited[END_REF] redefined the flattening index (A) of [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] as a percentage ratio: A= rostrum transverse diameter/rostrum sagittal diameter x 100 Rostra linear measurements were acquired using a vernier digital caliper (± 1 mm accuracy). Angular measurements were obtained with a protractor (± 1°) or by digital means. Pictures of the belemnite rostra covered with sublimated ammonium chloride were taken with a Canon SX60 semi-reflex camera. Linear illustrations and plates were prepared with CorelDraw 2020 Graphics Suite and Adobe Photoshop 2020.

Abbreviations

Institutional abbreviations. BAS KG,

Systematic paleontology

Order BELEMNOPSEINA Jeletzky, 1965 Family BELEMNOPSEIDAE Naef, 1922, amend Jeletzky, 1946 Genus Belemnopsis [START_REF] Bayle | Atlas. Première partie. Fossiles principaux des terrains. -Explication de la Carte géologique de la France[END_REF] Type species. Belemnopsis apiciconus Blainville, 1827 (pl. 2, figs. 2, 2a). ?Upper Aalenian-Bajocian, England, France, and Southern Germany (cf. [START_REF] Riegraf | Taxonomic status of the belemnite genus Belemnopsis Bayle 1878[END_REF].

Lectotype subsequently designated by Riegraf (1999, p. 66). A reassessment of the validity of the genus was recently undertaken by [START_REF] Mitchell | A reassessment of the validity and affinities of Belemnites sulcatus Miller, 1826, Belemnopsis Edwards in Gray, 1849, and Belemnopsis Bayle[END_REF] and followed here. Cross-section elliptical and depressed throughout (A c. 115 over the stem region), although this character might be accentuated by sedimentary flattening. A narrow, shallow ventral groove with angular margins runs over the mid-region of the stem and extends posteriorly at least up to 18 mm before the apex. The groove is 1 mm deep over the apical region, where it shows a "v" profile in cross-section and appears to deepen anteriorly. The apical line is slightly ventral situated, and no lateral lines were observed. [START_REF] Bayle | Atlas. Première partie. Fossiles principaux des terrains. -Explication de la Carte géologique de la France[END_REF]) groups more than one hundred European and Indo-Pacific belemnite species ranging from the Middle Jurassic up to the Lower Cretaceous [START_REF] Riegraf | Taxonomic status of the belemnite genus Belemnopsis Bayle 1878[END_REF]. Although we agree with [START_REF] Riegraf | Revision der Belemniten der Schwäbischen Jura. Part 7[END_REF], [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF][START_REF] Ippolitov | First find of Parabelemnopsis, the alien belemnite from the Southern Hemisphere, in the Upper Berriasian of Central Crimea[END_REF] that such a broad taxonomic concept is problematic, there is still no agreement among belemnite workers on the validity of the proposed subgeneric categories (e.g., [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF] and the poor preservation state of our material does not enable further insight on the matter, so we follow the classic taxonomic assignation to the genus Belemnopsis.

Remarks. The Tethyan genus Belemnopsis

The outline and profile general forms, as well as the length and distinctiveness of the ventral groove, are important taxonomic characters [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] that justify the assignation of the studied material to Belemnopsis. The rostrum cross-section appears to be a more variable trait since different authors have described Belemnopsis species with either circular to elliptical cross-sections (e.g., [START_REF] Doyle | The Jurassic and Cretaceous belemnites of Kong Karls land, Svalbard[END_REF][START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF] J o u r n a l P r e -p r o o f [START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF]. More specifically, the elongated rostrum, with an acute apex and a narrow ventral groove with angular margins, enables comparing the Neuquén Basin material with Belemnopsis patagoniensis [START_REF] Favre | Die Ammoniten der unteres Kreide Patagonien[END_REF], a well-known Lower Cretaceous species frequently recorded in the Austral Basin from the Berriasian to the Hauterivian (see section 1.2). Upper Tithonian (cf. [START_REF] Leanza | Anotaciones sobre los fósiles Jurásico-Cretácicos de Patagonia Austral (Colección Feruglio) conservados en la Universidad de Bologna[END_REF] B. patagoniensis described by [START_REF] Feruglio | Paleontographica Patagónica[END_REF] from Santa Cruz differ from the type material of the species [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] and has been repeatedly reassigned (see section 1.2). Independently of their systematic position, Feruglio's Belemnopsis and Hibolites argentinus show morphological affinities with Indonesian and New Zealand assemblages cf. [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] and to elements from Madagascar, cf. [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF].

Nonetheless, Patagonian specimens are typically more robust, almost twice wider and longer than the material described here (e.g., Favre, 1908, pl. 37, figs. 6-7;Riccardi, 1977, figs. 6a-c, g-i). A similar slender Berriasian specimen was described from Crimea by [START_REF] Ippolitov | First find of Parabelemnopsis, the alien belemnite from the Southern Hemisphere, in the Upper Berriasian of Central Crimea[END_REF], who assigned it to Parabelemnopsis cf. patagoniensis and interpreted its record in the Northern Hemisphere as a singular short invasion episode. The Crimean specimen differs from the Neuquén Basin material in having a much shorter ventral groove. Antarctic Belemnopsis, on the other hand, seem to be closely related to those from New Zealand-Indonesian of the "uhligi Complex" [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF], though more recently [START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF] indicated some affinities with Madagascan forms.

Belemnopsis patagoniensis, B. madagascariensis [START_REF] Besairie | Recherches geologiques a Madagascar[END_REF], and B. casterasi [START_REF] J O U R N A L P R E -P R O O F Besairie | Recherches geologiques a Madagascar. Premiere suite. La geologie du Nord-Ouest[END_REF] are recognized as a cluster of closely related Madagascan-South American Late Jurassic-Early Cretaceous large-sized, narrow-grooved belemnites [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF]. [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF], [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF], and [START_REF] Ippolitov | First find of Parabelemnopsis, the alien belemnite from the Southern Hemisphere, in the Upper Berriasian of Central Crimea[END_REF] have pointed out the strong similitude of some specimens J o u r n a l P r e -p r o o f assigned to B. patagoniensis, B. aff. madagascariensis [START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF][START_REF] Riccardi | The Cretaceous System of southern South America[END_REF] and B. madagascariensis (Aguirre-Urreta and Suarez, 1985). Even more, [START_REF] Combémorel | Les bélemnites de Madagascar[END_REF] suggested that B. madagascariensis and B. casterasi might conform to a dimorphic pair.

Phylogenetic relationships between them await a detailed revision to assess the extent of ontogenetic and population variability [START_REF] Challinor | Synonymy and stratigraphic ranges of Belemnopsis in the Heterian and Ohauan Stages (Callovian-Tithonian), southwest Auckland, New Zealand[END_REF][START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF]. Moderately preserved, 119 mm long, slender (nearly 22 times longer than thicker), symmetrical, and markedly hastate rostrum both in outline and profile. The maximum transverse diameter (dtM= 9.84 mm) is attained over the apical region, at about 25 mm from the apex, while the minimum transverse diameter (dtm= 5.4 mm) is placed over the broken anterior portion where no traces of the alveolus is observed. Apex moderately obtuse. Cross-section circular to slightly depressed over the apical and middle-stem region, where the venter is almost flattened, and more compressed to the anterior-stem region where the venter is more excavated. A medium-width, shallow ventral groove with rounded and weakly pronounced edges extends up to two-thirds J o u r n a l P r e -p r o o f over the stem. In a probable adult specimen (MCNAM-PI 24647.2; Figs. 4.7-4.9), the groove still has rounded margins, but it is broader (3 mm wide) and runs into the apical region, at least up to 30 mm from the apex. Specimen MCNAM-PI 24647.2 also shows a slightly asymmetrical profile due to inflation of the venter, and the outline appears to be cylindroconical (dtM= 12.2 mm) with a circular cross-section (A= 100). No dorsal or lateral grooves are observed in any of the specimens. The rest of the rostra recorded in the same bed, 15.4 mm long on average (dtM= 1.4), are too poorly preserved to provide any further insights into ontogenetic traits.

Remarks. The studied specimens were retrieved from the same interval where Gerth (1925) collected some material which was then revised by Stolley and identified as Hibolithes sp. (cf. Gerth, 1925, p. 42, footnote). However, some rostra traits, like the slender and long stem region and the shallow, medium to broad ventral groove, make them akin to Belemnopsis launceloti Howlett (1989, p. 40, pl. 6, figs. 1-2, 6-7;pl. 8, figs. 5-6) from the Alexander Island. Still, the Antarctic material is less hastate, has a maximum transverse diameter over the middle stem region, and the ventral groove narrows anteriorly (pictures of the type material of B. launceloti are available at the BAS website: https://legacy.bas.ac.uk/bas_research/data/access/fossildatabase/filter.php?taxonKeywor d=BELEMNITIDA). Some of the mentioned differences might have an ontogenetic explanation. Stevens (1965, p. 40) indicated that Belemnopsis suffered changes in its degree of hastation in the early growth stages of the rostrum. Challinor (2001, p. 234) also pointed out that Belemnopsis aucklandica presents the point of maximum transverse diameter nearer the apex in juvenile specimens but higher up in adults, a trait that affects the degree of hastation of the rostrum. He also noted that in more hastated individuals, the ventral groove tends to weaken at about the mid-rostrum and extend J o u r n a l P r e -p r o o f apically as a wide shallow depression. Unfortunately, the poor preservation state of the material hinders further insight into its ontogeny, so its systematic assignation is left open. [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF] included in the B. launceloti synonymy list two specimens described by Feruglio (1936, pl. 10, figs. 3-4) from the Austral Basin, formerly reassigned to B. gladiatoris by [START_REF] Willey | Belemnites from South-Eastern Alexander Island. II, The occurrence of the Family Belemnopseidae in the Upper Jurassic and Lower Cretaceous[END_REF], both of which are more robust than the Neuquén Basin material. [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF], also renamed as B. launceloti a specimen from Madagascar figured as B. africanus Tate, 1867 by Besairie (1936, p. 23, figs. 6-7).

Genus Hibolithes [START_REF] Montfort | Conchyliologie systematique et classification methodique des Coquilles[END_REF] Type species. Hibolithes hastatus Montfort, 1808 (text-fig. 47), by monotypy.

Bajocian-Aptian of some Boreal and most Tethyan regions [START_REF] Doyle | The Jurassic and Cretaceous belemnites of Kong Karls land, Svalbard[END_REF][START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF].

Hibolithes argentinus? (Feruglio, 1936)

J o u r n a l P r e -p r o o f

Remarks. The circular form of the rostrum cross-section and the absence of a ventral groove in the preserved portion leads us to consider this fragment as part of the lower stem region of an Hibolithes representative. The most frequent Hibolithes species in Argentina is H. argentinus [START_REF] Feruglio | Paleontographica Patagónica[END_REF], which is characterized by a slender hastate rostrum with circular to slightly depressed transverse sections and a shallow ventral groove that extends no further than the upper stem region. Howlett (1989, p. 52) designated one of the specimens figured by Feruglio (1936, pl. 10, fig. 17) as lectotype and added new material from Alexander Island. The fragment here described agrees well both with the lower stem regions of the lectotype and the with the specimen BAS KG 2909.18 figured by Howlett (1989, pl. 6, figs. 4, 10), but as no other features are preserved, the assignation to H. argentinus is stated with doubts.

Discussion

Paleobiogeographic considerations

Several authors differentiated an Indo-Pacific Province within the Tethyan Realm from the Kimmeridgian onwards over the basis of belemnite geographic distribution, particularly of Belemnopsis and Hibolithes regional assemblages (e.g., [START_REF] Combémorel | Les bélemnites de Madagascar[END_REF][START_REF] Doyle | Antarctic belemnites biogeography and the breack-up of Gondwana[END_REF][START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF], 1992[START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF], 1973).

Alternatively, [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Grant-Mackie | Jurassic palaeobiogeography of Australasia[END_REF] proposed a different subdivision of the Tethyan Realm into two provinces, 1) Tethyan Province, including a Mediterranean, Ethiopian and Tethyan Subprovinces (the last partly equivalent to the Indo-Pacific Province), and 2) South Pacific Province, extending along the Pacific coasts of Gondwana, from southern South America (including the Neuquén Basin) through West Antarctica to New Zealand, and probably including New Caledonia. The South Pacific Province was not completely isolated from the Tethyan Province and at least from the latest Jurassic, some belemnite migrations took place J o u r n a l P r e -p r o o f from the Ethiopian Subprovince into Antarctica and South America [START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF][START_REF] Mutterlose | Early Cretaceous belemnites from the East Indian Ocean and their paleobiogeographic implications[END_REF]. From the Oxfordian to the Valanginian, a long-ranging lineage of narrowly-grooved Belemnopsis developed in Madagascar and Eastern Africa, forming the B. madagascariensis-B. casterasi group [START_REF] Besairie | Recherches geologiques a Madagascar[END_REF][START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF][START_REF] Combémorel | Les bélemnites de Madagascar[END_REF]. Some components of that cluster, as well as Hibolithes, might have reached the South Pacific coast from the Tithonian onwards through the Trans-Gondwana or Indo-Madagascan seaway (e.g., [START_REF] Cecca | Palaeobiogeography of Tethyan ammonites during the Tithonian (latest Jurassic)[END_REF][START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF]Fig. 6).

Even more, during the Berriasian, that marine connection is supposed to have held a "relict" population of Belemnopsis of the madagascariensis-casterasi group, since Belemnopsis is unknown or poorly represented from European and most Indo-Pacific regions at that time [START_REF] Doyle | Antarctic belemnites biogeography and the breack-up of Gondwana[END_REF][START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF]. That remnant population is thought to be related to the Belemnopsis patagoniensis assemblages of the Austral Basin [START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF][START_REF] Riccardi | The Cretaceous System of southern South America[END_REF] and now from the upper Tithonian in the Neuquén Basin.

The same is valid for B. launceloti, recorded in Madagascar, Alexander Island, and in the Austral Basin. Belemnopsis records here described add new elements connecting the Neuquén Basin assemblages with those of Madagascar and Eastern Africa, passing through Antarctica during the Upper Jurassic. These basins are known to share several ammonoid genera among other marine invertebrate and vertebrate taxa [START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF][START_REF] Riccardi | Jurassic and Cretaceous marine connections between the Southeast Pacific and Tethys[END_REF]Vennari and Aguirre-Urreta, 2019 and references therein;Campos et al., 2021).

FIGURE 6 AROUND HERE

Biostratigraphic considerations

Belemnites' biostratigraphic potential has long been acknowledged for Jurassic and Cretaceous successions, though belemnite-based correlations are more widely used for the Cretaceous of the Northern Hemisphere, as summarized by [START_REF] Christensen | Upper Cretaceous belemnite biostratigraphy of Europe[END_REF] and J o u r n a l P r e -p r o o f [START_REF] Mutterlose | A belemnite scale for the Lower Cretaceous[END_REF]. [START_REF] Doyle | Belemnites in biostratigraphy[END_REF] complemented those contributions by reviewing Jurassic belemnite occurrences of the Southern Hemisphere, and then [START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF][START_REF] Doyle | Jurassic belemnite distribution patterns: implications of new data from Antarctica and Argentina[END_REF] focused on the Antarctic and the Argentinean Jurassic records. The former includes the Kimmeridgian-Tithonian Belemnopsis fauna described from the Antarctic Peninsula and Alexander Island (Crame [START_REF] J O U R N A L P R E -P R O O F Crame | Late Jurassic and Early Cretaceous biostratigraphy of the Fossil Bluff Formation, Alexander Island[END_REF][START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF][START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF][START_REF] Riley | Late Jurassic (Kimmeridgian-Tithonian) macrofossil assemblage from Jason Peninsula, Graham Land: evidence for a significant northward extension of the Latady Formation[END_REF]. Upwards, the Tithonian/Berriasian transition is encompassed by the Hibolithes fauna (or Biozone according to [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF], which includes several species of Hibolithes, with H. argentinus, and Belemnopsis species of Indo-Pacific affinities as rare components. Tithonian Antarctic belemnite associations also include Duvalia/Produvalia [START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF][START_REF] J O U R N A L P R E -P R O O F Crame | Late Jurassic and Early Cretaceous biostratigraphy of the Fossil Bluff Formation, Alexander Island[END_REF], a strongly compressed genus with a dorsal median groove that is supposed to have immigrated into the Indo-Pacific region from the Tethys during the Tithonian [START_REF] Stevens | The Jurassic and Cretaceous belemnites of New Zealand, and a review of the Jurassic and Cretaceous belemnites of the Indo-Pacific region[END_REF] or as early as the Callovian [START_REF] Challinor | New Middle and Upper Jurassic Belemnite Assemblages from West Antarctica (Latady Group, Ellsworth Land): Taxonomy and Paleobiogeography[END_REF].

Concerning Argentinean Jurassic successions, [START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF] recognized two successive belemnite faunas for the Upper Jurassic-Lower Cretaceous namely early Belemnopsis (Oxfordian) and Hibolithes (Tithonian-Berriasian), which was named after H. argentinus. In that contribution, the Lower Cretaceous occurrences of B. patagoniensis were treated separately. Meanwhile, B. launceloti is an upper Tithonian-Hauterivian species that defines an assemblage biozone in Antarctica [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF] and has been recorded from the lower Valanginian of Madagascar [START_REF] J O U R N A L P R E -P R O O F Besairie | Recherches geologiques a Madagascar. Premiere suite. La geologie du Nord-Ouest[END_REF][START_REF] Combémorel | Les bélemnites de Madagascar[END_REF], the upper Tithonian of the Austral Basin in the Lago Argentino area (cf. [START_REF] Leanza | Anotaciones sobre los fósiles Jurásico-Cretácicos de Patagonia Austral (Colección Feruglio) conservados en la Universidad de Bologna[END_REF], and with reservations from the Valanginian-Hauterivian of southern Chile [START_REF] Stinnesbeck | A Lower Cretaceous ichthyosaur graveyard in deep marine slope channel deposits at Torres del Paine National Park, southern Chile[END_REF]. So, Belemnopsis records described in this contribution indicate that this genus should also be considered in the Neuquén Basin as J o u r n a l P r e -p r o o f a component of the well-established Upper Jurassic-Lower Cretaceous Hibolithes fauna of Gondwana [START_REF] Doyle | Jurassic belemnite biostratigraphy of the Southern Hemisphere: a comparative study from Antarctica and Argentina[END_REF][START_REF] Challinor | Belemnite successions and faunal provinces in the southwest Pacific, and the belemnites of Gondwana[END_REF].

Whenever possible, the correlation of the Neuquén Basin belemnite records in each locality with the current ammonoid Andean zonation [START_REF] Aguirre-Urreta | The Tithonian chronobiostratigraphy of the Neuquén Basin and related Andean areas: a review and update[END_REF][START_REF] Leanza | Relevant marine paleobiological markers of the Vaca Muerta Formation[END_REF][START_REF] Lescano | New Tithonian biostratigraphic data from the J o u r n a l P r e -p r o o f Neuquén Basin, Argentine Andes: a contribution from the Southern Hemisphere[END_REF] is summarized in Figure 7. Hibolithes sp. from the Arroyo La Manga area was found by [START_REF] Gerth | Contribuciones a la estratigrafía y la paleontología de los Andes Argentinos II. La Fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza[END_REF] koeneni Biozone [START_REF] Kietzmann | Sedimentology and sequence stratigraphy of a Tithonian-Valanginian carbonate ramp (Vaca Muerta Formation): a misunderstood exceptional source J o u r n a l P r e -p r o o f rock in the southern Mendoza area of the Neuquén basin, Argentina[END_REF][START_REF] Kietzmann | Facies analysis and stratigraphy across the Jurassic-Cretaceous boundary in a new basinal Tithonian-Berriasian section of the Vaca Muerta Formation, Las Tapaderas, Southern Mendoza Andes, Argentina[END_REF]. Meanwhile, belemnites from Cañada de Leiva come from a storm-dominated middle ramp setting, corresponding to a highstand system tract [START_REF] Kietzmann | Sedimentology and sequence stratigraphy of a Tithonian-Valanginian carbonate ramp (Vaca Muerta Formation): a misunderstood exceptional source J o u r n a l P r e -p r o o f rock in the southern Mendoza area of the Neuquén basin, Argentina[END_REF][START_REF] Kietzmann | Astronomical calibration of the Tithonian-Berriasian in the Neuquén Basin, Argentina: A contribution from the Southern Hemisphere to the Geologic Time Scale[END_REF]. However, other factors than a high local sea level alone would have influenced belemnite distribution.

Although some authors have modeled the existence of counter-oceanic currents at the latitude of the Neuquén Basin [START_REF] Parrish | Jurassic climate and oceanography of the Pacific region[END_REF], up to the authors' best knowledge, there were no physical geographic barriers between the Austral and the Neuquén Basin that prevented a free interchange of nektonic fauna as both were retro-arc basins open to J o u r n a l P r e -p r o o f the Paleo-Pacific. Indeed, both basins share several ammonoid genera [START_REF] Riccardi | Jurassic and Cretaceous marine connections between the Southeast Pacific and Tethys[END_REF], although comparatively poorly preserved and less abundant in the case of the former [START_REF] Riccardi | Jurassic and Cretaceous marine connections between the Southeast Pacific and Tethys[END_REF]. Nevertheless, two alternative paleogeographic models for the westernmargin configuration of the Neuquén Basin were proposed, either enabling a direct connection with the Paleopacific Ocean through a discontinuous arc of volcanic islands (e.g., [START_REF] Howell | The Neuquén Basin: A case study in sequence stratigraphy and basin dynamics[END_REF] or a more restricted seaway in the north-western sector: the Curepto Strait [START_REF] Vicente | Dynamic paleogeography of the Jurassic Andean Basin: patterns of transgression and localization of main straits through the magmatic arc[END_REF]. Nevertheless, the high diversity and Tethyan affinities of several Neuquén Basin ammonoids and herpetofauna militate against the possibility of a confined basin [START_REF] Gasparini | Jurassic marine reptiles of the Neuquén Basin: records, faunas and their palaeobiogeographic significance[END_REF][START_REF] Riccardi | Remarks on the Tithonian-Berriasian ammonite biostratigraphy of west-central Argentina[END_REF].

Besides, [START_REF] Alberti | Stable isotopes (δ13C, δ18O) and element ratios (Mg/Ca, Sr/Ca) of Jurassic belemnites, bivalves and brachiopods from the Neuquén Basin (Argentina): challenges and opportunities for palaeoenvironmental reconstructions[END_REF] recently demonstrated, over Sr/Ca ratios and their comparison with the global curve, the existence of water exchange with the open ocean in several localities of the Neuquén Province.

Regarding average seawater temperature and salinity, [START_REF] Alberti | Stable isotopes (δ13C, δ18O) and element ratios (Mg/Ca, Sr/Ca) of Jurassic belemnites, bivalves and brachiopods from the Neuquén Basin (Argentina): challenges and opportunities for palaeoenvironmental reconstructions[END_REF] found some strong influence of rainfall and/or a continental freshwater influx in their northernmost sampled sites, and interpreted a water temperature of 20-25 °C from δ 18 O data from oyster and brachiopod shells. That temperature range agrees well with previous estimations by Gómez [START_REF] Dacal | Petrología, geoquímica y diagénesis de sedimentitas y fósiles carbonáticos del Jurásico Superior-Cretácico Inferior de las cuencas Neuquina y Austral, Argentina[END_REF] and Gómez [START_REF] Dacal | Evidence of Warm Seas in High Latitudes of Southern South America During the Early Cretaceous[END_REF] of a medium water temperature of 23 °C from oysters of the Neuquén Basin, and 24 °C from belemnites (Belemnopsis sp.) of the Austral Basin. Temperatures obtained in both cases are in agreement with the supposed range of 10-30 °C that belemnites inhabited by comparison with modern cephalopod preferences [START_REF] Hoffmann | The palaeobiology of belemnites -foundation for the interpretation of rostrum geochemistry[END_REF], though we do not know to which extent minor temperature changes affected belemnite distribution since it is considered the main parameter controlling their migration routes at species and genus levels (e.g., [START_REF] J O U R N A L P R E -P R O O F Mutterlose | Migration and evolution patterns in Upper Jurassic and Lower Cretaceous belemnites[END_REF][START_REF] Mutterlose | The first belemnite of boreal ancestry from the Early Cretaceous (Valanginian) of the western Tethys: implications for belemnite ecology[END_REF][START_REF] J O U R N A L P R E -P R O O F Stevens | Jurassic belemnites[END_REF]. Other paleoecologic requirements listed by [START_REF] Hoffmann | The palaeobiology of belemnites -foundation for the interpretation of rostrum geochemistry[END_REF], such J o u r n a l P r e -p r o o f as well-oxygenated waters, with a normal salinity of 27-37 psu and a depth range of no more than 200 m would characterize the Vaca Muerta Formation Upper Jurassic carbonate ramp (e.g., [START_REF] Kietzmann | Sedimentology and sequence stratigraphy of a Tithonian-Valanginian carbonate ramp (Vaca Muerta Formation): a misunderstood exceptional source J o u r n a l P r e -p r o o f rock in the southern Mendoza area of the Neuquén basin, Argentina[END_REF]. Even if, locally, some of the parameters would deviate from the optimum (e.g., by the influx of freshwater), belemnites could have just actively swum away from such conflicting areas.

In most recent revisions of the group, belemnites are considered medium-sized predators of Mesozoic food webs, most likely feeding upon small vertebrates, mollusks, and crustaceans (Hoffman and Stevens, 2020 and references therein). All these food sources were present at the Neuquén Basin, and possibly some v-shaped scars over ammonoid shells were caused by non-lethal belemnite attacks [START_REF] Vennari | Estudio preliminar del registro de heridas letales y subletales en amonoideos del Jurásico Tardío[END_REF]. In time, belemnites were occasionally preyed upon by marine reptiles such as ichthyosaurs and metriorhynchids [START_REF] Fernández | Marine reptiles from the Jurassic/Cretaceous transition at the High Andes, Mendoza, Argentina[END_REF], which might have gotten into the basin mostly for reproductive purposes [START_REF] Gasparini | Jurassic marine reptiles of the Neuquén Basin: records, faunas and their palaeobiogeographic significance[END_REF]. However, most probably, belemnite hatchlings were consumed by pachycormids, a stem-teleost clade of suspensivorous-feeding fishes that were quite successful from the Early Jurassic to the Late Cretaceous [START_REF] Friedman | 100-million year dynasty of giant planktivorous bony fishes in the Mesozoic seas[END_REF]. Pachycormids, which are well-known from the Neuquén Basin [START_REF] Gouiric-Cavalli | Large and mainly unnoticed: the first lower Tithonian record of a suspension-feeding pachycormid from Southern Gondwana[END_REF][START_REF] J O U R N A L P R E -P R O O F Gouiric-Cavalli | A new Pachycormiformes (Actinopterygii) from the Upper Jurassic of Gondwana sheds light on the evolutionary history of the group[END_REF], to date have not been recorded in the Austral Basin, though they have cited from Chile [START_REF] Arratia | Los peces osteíctios fósiles de Chile y su importancia en los contextos paleobiogeográfico y evolutivo[END_REF]. Although pachycormid gill rakers spacing was suitable to filter out both hatchlings and juvenile ammonites [START_REF] Tajika | The old and the new plankton: ecological replacement of associations of mollusc plankton and giant filter feeders after the Cretaceous?[END_REF], the different reproductive strategies of both cephalopod groups may have turned the odds in favor of ammonite survival. On the one hand, ammonites, either semelparous or iteroparous, are assumed to have been extreme r-strategists, with high fecundity rates and the possibility, for medium-sized macroconchs, to lay more than 10,000 eggs [START_REF] De Baets | Ammonoid embryonic development[END_REF][START_REF] Mironenko | First direct evidence of ammonoid ovoviviparity[END_REF][START_REF] Tajika | The old and the new plankton: ecological replacement of associations of mollusc plankton and giant filter feeders after the Cretaceous?[END_REF][START_REF] Greif | Extreme abundance of ammonoids in mass accumulations from the Late Devonian of the Moroccan Anti-Atlas[END_REF]. Even more, J o u r n a l P r e -p r o o f ovoviviparity has been proven for some ammonites [START_REF] Mironenko | First direct evidence of ammonoid ovoviviparity[END_REF], thus incrementing the chances of post-hatching survival. Nevertheless, direct evidence of this predation relation should be proven through bromalite analyses (i.e., oral or anal depositions or intestinal content) as conducted in other basins [START_REF] Zatón | Durophagus predation on middle Jurassic Molluscs, as evidenced from shell fragmentation[END_REF][START_REF] Ojeda | Las bromalitas de la Formación Ameghino (Jurásico Tardío) de la Península Antártica[END_REF][START_REF] Lukeneder | Bromalites from the Upper Triassic Polzberg section (Austria); insights into trophic interactions and food chains of the Polzberg palaeobiota[END_REF]. Belemnites, on the other hand, are assumed to have been r-strategists (e.g., [START_REF] Doguzhaeva | Embryonic shell structure of Early-Middle Jurassic belemnites, and its significance for belemnite expansion and diversification in the Jurassic[END_REF]) and to have depicted high fecundity rates as well, but of several magnitude order lower than that of ammonites due to a smaller size difference between adults and embryos [START_REF] Tajika | The old and the new plankton: ecological replacement of associations of mollusc plankton and giant filter feeders after the Cretaceous?[END_REF]. In consequence, ammonite surviving into adulthood was more likely than for belemnites.

To sum up, marine connections, seawater physical parameters, or food sources availability seem not to have constrained, on their own, the presence of belemnites in the Neuquén Basin in favor of ammonites. However, they were likely affected by the predation pressure exerted by filter-feeding organisms on hatchlings, combined with unsuitable paleoecological conditions over shallow marine settings relatively close to the coast where belemnites were assumed to inhabit [START_REF] Naef | Die fossilen Tintenfische[END_REF][START_REF] Doguzhaeva | Embryonic shell structure of Early-Middle Jurassic belemnites, and its significance for belemnite expansion and diversification in the Jurassic[END_REF][START_REF] Mutterlose | The first belemnite of boreal ancestry from the Early Cretaceous (Valanginian) of the western Tethys: implications for belemnite ecology[END_REF]. [START_REF] Favre | Die Ammoniten der unteres Kreide Patagonien[END_REF], [START_REF] Besairie | Recherches geologiques a Madagascar[END_REF][START_REF] J O U R N A L P R E -P R O O F Besairie | Recherches geologiques a Madagascar. Premiere suite. La geologie du Nord-Ouest[END_REF], [START_REF] Feruglio | Paleontographica Patagónica[END_REF]Feruglio ( -1937[START_REF] Feruglio | Geología de la Patagonia[END_REF], [START_REF] Harrington | Observaciones geológicas en la Isla de los Estados[END_REF], Stevens (1965, and references therein), [START_REF] Willey | Belemnites from South-Eastern Alexander Island. II, The occurrence of the Family Belemnopseidae in the Upper Jurassic and Lower Cretaceous[END_REF], [START_REF] Riccardi | Berriasian invertebrate faunas from the Springhill Formation in southern Patagonia[END_REF]), Jeletzki (1983), [START_REF] Combémorel | Les bélemnites de Madagascar[END_REF], [START_REF] J O U R N A L P R E -P R O O F Crame | Late Jurassic and Early Cretaceous biostratigraphy of the Fossil Bluff Formation, Alexander Island[END_REF], [START_REF] Howlett | Late Jurassic-Early Cretaceous cephalopods of eastern Alexander Island, Antarctica[END_REF], Doyle and Howlett (1989, and references therein), Aguirre-Urreta and Suárez (1985, and references therein), [START_REF] Mutterlose | Upper Jurassic Belemnites from the Orville Coast, Western Antarctica, and their palaeobiogeographical significance[END_REF] references therein), [START_REF] Olivero | Cefalópodos y bivalvos titonianos y hauterivianos de la Formación Lago La Plata, Chubut[END_REF], Challinor (1991, and references therein), Challinor andHikuroa (2007), Crame et al. (2000), and [START_REF] Stinnesbeck | A Lower Cretaceous ichthyosaur graveyard in deep marine slope channel deposits at Torres del Paine National Park, southern Chile[END_REF]. Map based on [START_REF] Smith | Atlas of Mesozoic and Cenozoic Coastlines[END_REF] et al., 2019;Leanza et al., 2020, Lescano et al., in press). White stars, Hibolithes sp. records [START_REF] Gerth | Contribuciones a la estratigrafía y la paleontología de los Andes Argentinos II. La Fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza[END_REF][START_REF] Alberti | Stable isotopes (δ13C, δ18O) and element ratios (Mg/Ca, Sr/Ca) of Jurassic belemnites, bivalves and brachiopods from the Neuquén Basin (Argentina): challenges and opportunities for palaeoenvironmental reconstructions[END_REF] J o u r n a l P r e -p r o o f

Concluding remarks

  British Antarctic Survey Collection, Cambridge, United Kingdom; CPBA, Paleontological Collection of the University of Buenos Aires, Buenos Aires, Argentina; MCNAM-PI, Colección de Paleontología de Invertebrados del Museo de Ciencias Naturales y Antropológicas "Juan Cornelio Moyano," Mendoza, Argentina. J o u r n a l P r e -p r o o f Anatomical abbreviations. A, index of flattening; dtM, maximum transverse diameter; dtm, minimum transverse diameter. Stratigraphical abbreviations. BB, Cañada de Leiva section; DO, Cerro Domuyo section; LA, Las Alcantarillas section; LL, Las Loicas section.

  Figures 4.1-4.6; 5.1-5.2

FIGURE 4

 4 FIGURE 4 AND FIGURE 5 AROUND HERE

Figure

  Figure 4.14-4.16

  FIGURE 7 AROUND HERE

  We presented in this contribution new records of Upper Jurassic belemnites from the Vaca Muerta Formation, Neuquén Basin, west-central Argentina. The studied rostra were assigned to Belemnopsis cf. B. patagoniensis, Belemnopsis cf. B. launceloti, and Hibolithes argentinus?. These are the first thoroughly described belemnites from the Neuquén Basin. By their association with ammonoid faunas, the new belemnite records can be referred to the Tithonian Windhauseniceras internispinosum to Corongoceras alternans Biozones. The biostratigraphic framework suggests that Belemnopsis should J o u r n a l P r e -p r o o f be contemplated as a component of the widely-known Hibolithes fauna recorded across the Late Jurassic-Early Cretaceous of Gondwana. Affinities of the described fauna seem close to Madasgascan-Eastern African belemnite associations of the Ethiopian Subprovince, thus reinforcing the hypothesis of a Trans-Gondwana seaway as an open marine connection at least since the Tithonian. The baffling scarcity of belemnites from the Late Jurassic in the Neuquén Basin and their absence from the Early Cretaceous upwards is in utter contrast with their abundant record in the Austral Basin. Though still unresolved, it is suggested that a combination of factors, such as local variations in seawater salinity/temperature in shallow environments and, particularly, the predation pressure of filter-feeding fishes on belemnite hatchlings, may have affected the presence of the group in the basin. Also, the possible existence of counter-currents at the latitude of the Neuquén Basin may have limited the extension to lower latitudes of the Belemnopsis/Hibolithes Gondwanan faunas.

Figure captions Figure 1 . 1 . 1 .

 captions111 Figure captions

Figure 2 .

 2 Figure 2. Log sections of the belemnite-bearing localities indicating the stratigraphic

Figure 3 . 1 ,

 31 Figure 3. Hummocky cross-bedding bioclastic packstone (CLBe) thin section, views of

Figure 4 .

 4 Figure 4. Belemnites from the Vaca Muerta Formation, Neuquén Basin. 1-6,

Figure 5 .Figure 6 .

 56 Figure 5. Belemnopsis rostra sketch diagrams. 1-2, Belemnopsis cf. B. patagoniensis

Figure 7 .

 7 Figure 7. Summary of occurrences of the Tithonian-Berriasian Neuquén Basin

  ; grey stars, Hibolithes argentinus and H. argentinus? records (this work); black stars, Belemnopsis cf. B. patagoniensis records (this work). DO, Cerro Domuyo; CL, Cañada de Leiva; LA, Las Alcantarillas; LL, Las Loicas; LM, Arroyo La Manga; PT, Pampa Tril. J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f Highlights • New Tithonian belemnite data from the Neuquén Basin, western Argentina.• 21 Belemnites are assigned to the genera Belemnopsis and Hibolithes.•The belemnite association is close to Madagascan-Eastern African faunas.• Generic composition and affinity of the belemnite fauna add new evidence supporting an open Trans-Gondwana seaway from the Tithonian onwards.

  introduced a supposedly Valanginian belemnite association composed of B. patagoniensis, Hibolithes argentinus Feruglio, H. aff. H. jaculum, and a specimen openly assigned as Belemnites sp. I, which was reinterpreted as a Belemnopsis sp. by

  . 1, Neuquén Basin; 2, Lago San Martín, Austral Basin; 3, Lago Argentino, Austral Basin; 4, Torres del Paine, Chile; 5, Tierra del Fuego Archipelago); 6, Alexander Island, Antarctic Peninsula; 7, Malvinas/Falkland Plateau; 8, Orville Coast and Ellsworth Land, Western Antarctica; 9, Eastern Wedell Sea; 10, Madagascar; 11, Kutchh; 12, Northwestern Pakistan; 13, Spiti Shales; 14, Australia; 15, New Guinea; 16, Moluccas and Sula Islands; 17, Misool, Indonesia; 18, New Zealand.

Acknowledgments

The authors are grateful to all field assistants, to Leonardo Pazo (Fundación Azara) for preparing the specimens, and to the editors and reviewers that greatly improved this contribution. This work was partially financed by the ANPCyT PICT 1016-2016 and PICT 2020-02067 (V.V.V); Sepkoski Grant 2019 (V.V.V.); PIP CONICET 11220170101117CO (B.A.U.) and 11220200102844CO (V.V.V.); CNRS Grants SYSTER-TELLUS "ACJC" in 2019 and "GC-JC" in 2021; and MITI AAP Temps "AstroCarb" (M.M. and P.P.). This is contribution R-445 of the Instituto de Estudios Andinos "Don Pablo Groeber".

Authors statement

Verónica V. Vennari: Conceptualization, methodology, data acquisition, investigation, writing, supervision, and funding acquisition. Beatriz Aguirre-Urreta: investigation, data acquisition, writing, and funding acquisition. Luciana S. Marin: data acquisition, methodology and writing. Pierre Pellenard: data acquisition, writing, and funding acquisition. Mathieu Martinez: data acquisition, writing, and funding acquisition. Maisa Tunik: data acquisition and writing.

J o u r n a l P r e -p r o o f

Declaration of interests

☒ The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

☐The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: