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Home All issues Volume 196 (2025) BSGF - Earth Sci. Bull., 196 (2025) 17 Full HTML
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BSGF - Earth Sci. Bull.
Volume 196, 2025
Article Number 17
Number of page(s) 25
DOI https://doi.org/10.1051/bsgf/2025015
Published online 17 October 2025

© M.-B. Forel et al., Published by EDP Sciences 2025

Licence Creative CommonsThis is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

1 Introduction

Cold seeps are sites where low temperature fluids enriched in hydrocarbons and hydrogen sulphide are emitted onto the seafloor after they migrated through sediments (e.g., Paull et al., 1984; Dando et al., 1991; Baco et al., 2010). They occur on both active and passive margins and host communities relying on chemosynthetic production at the base of their food chain (e.g., Van Dover, 2000; Levin, 2005). Important members of these chemosynthetic communities today include vestimentiferan tube worms (polychaete Siboglinidae Caullery, 1914), vesicomyid clams and bathymodiolin mussels, accompanied by diverse bivalves (lucinids, solemyids, thyasirids), gastropods (buccinids, trochids), crustaceans (amphipods, bresiliid shrimps, galathaeoids), polychaetes and sponges (e.g., Sibuet and Olu, 1998; Tunnicliffe et al., 2003). The origin of most of them can be traced into the Eocene or Late Cretaceous (e.g., Amano and Kiel, 2007; Kaim et al., 2009).

It has long been considered that the evolutionary history of chemosynthetic communities during the Paleozoic was dominated by rhynchonellid brachiopods (e.g., Campbell and Bottjer, 1995a, b), while they are virtually absent from modern ones. This view was challenged by the discovery of the oldest known cold seep community dominated by bivalves of the extinct family Modiomorphidae Miller, 1877 from the Ludlow, Silurian (Ludfordian) of Morocco (Jakubowicz et al., 2017). Conversely, the Early Cretaceous history of cold seeps was marked by mass occurrences of large shells of the brachiopod Peregrinella (Peregrinella) multicarinata (Lamarck, 1819) that have intrigued geologists and palaeontologists since their first observation on the field (e.g., Paquier, 1900; Ascher, 1906; Trümpy, 1956). Representatives of Peregrinella (Peregrinella) multicarinata are the largest known rhynchonellids during the Mesozoic (e.g., Trümpy, 1956; Biernat, 1957) and have thus puzzled the community in regard of the evolution of brachiopod shell-size through the Permian-Triassic mass extinction when newly appearing orders were smaller than their ancestors (e.g., He et al., 2017). The discovery of present-day cold seeps and their communities in 1983 (Paull et al., 1984) allowed palaeontologists to understand that these unique Lower Cretaceous fossiliferous deposits represent ancient seeps (Campbell and Bottjer, 1995b), likely with diffusive seepage (Kiel et al., 2014). Such Peregrinella-rich deposits have been reported from several sites within the Vocontian Basin in south-eastern France since 1835, including Curnier in the Drôme department, which was long unlocated since its first mention in Paquier (1900). Here we describe the largest known collection of Peregrinella brachiopods from Curnier, stored in the Muséum national d’Histoire naturelle in Paris. For the first time, we describe the associated ostracods and scolecodonts and discuss their implications in terms of palaeocology and palaeoenvironments.

2 Geological setting

2.1 The Vocontian Basin

During the Early Cretaceous, most of south-eastern France was occupied by the Vocontian Basin, an east-west oriented sub-basin of the Tethys Ocean located at a palaeolatitude of 25 to 30°N (Fig. 1A; e.g., Dercourt et al., 1993; Cecca, 1997; Ferry, 2017). It was bounded by the partly emerging massifs of the Maures-Esterel on the south and Massif Central on the west, and was linked to the European continental margin by the Jura margin (Fig. 1B). During the Valanginian-lower Aptian interval, south-eastern France was characterized by the great development of carbonate platforms (northern subalpine massifs, Ardèche, Gard, Provence) around the Vocontian Basin, within which marl-limestone alternations were deposited in a pelagic to hemipelagic environment at several hundred of meters water depth (e.g., Lemoine et al., 1982, 1986; Cotillon, 1984; Reboulet et al., 1992; Wilpshaar et al., 1997; Adatte et al., 2005; Vermeulen et al., 2013; Ferry, 2017). During this interval, active synsedimentary faults or fault-bounded diapirs (e.g., Arnaud, 1981; Lemoine et al., 1982; Perthuisot and Guilhaumou, 1983; Mascle et al., 1988) were associated with unusual fossil-rich limestone lenses successively called “lentilles à Rhynchonelles” and “lentilles à Pérégrinelles” because of the abundance of large brachiopods today attributed to Peregrinella Oehlert, 1887 (e.g., Paquier, 1900; Thieuloy, 1972; Lemoine et al., 1982).

thumbnail Fig. 1

A. Palaeogeographical map of western Tethys during the latest Hauterivian (modified from Dercourt et al., 1993) with position of the south-eastern France Basin. B. Palaeogeography of south-eastern France Basin and nearby areas during the Hauterivian (after Arnaud-Vanneau et al., 1982; Ferry, 1984, Ferry, 2017) showing the three main sites yielding Peregrinella-lenses discussed in the text. F1: Menée fault, F2: Trente-Pas-Condorcet fault.

2.2 Peregrinella-lenses

The “lentilles à Pérégrinelles” or Peregrinella-lenses in the Vocontian area are known since the discovery of the Ravin de Quintel site (Drôme), later called Châtillon-en-Diois, by the geologist Scipion Gras in 1835, close to the Menée fault (Fig. 1B). The site of Curnier, close to the Trente-Pas-Condorcet fault, was discovered in 1894 by Victor Paquier who mentioned gastropods, bivalves and ammonites accompanying abundant brachiopod specimens of Rhynchonella peregrina von Buch, 1834, since re-identified as Peregrinella (Peregrinella) multicarinata (Fig. 1B; Paquier, 1900). The fossils collected by Paquier at Curnier were re-analyzed by Thieuloy (1972) who updated their identifications and concluded that this fauna was probably of early Hauterivian age based on ammonoids. Thieuloy (1972) also considered that the Curnier exposure was from a detached block that was a lateral equivalent of the upper part of the succession exposed at the Rottier site, close to the Menée fault (Hébert, 1871; Fig. 1B). Thieuloy (1972) considered that the Rottier sequence, including Peregrinella-lenses, was deposited on a relatively narrow shoal under shallow water, surrounded by pelagic area and possibly influenced by intertidal mechanisms. The discovery of hydrothermal ecosystems along present-day ocean ridges (Lonsdale, 1977; Corliss et al., 1979) coupled with fossil content, isotopic analyses, palaeogeographic and geological contexts led Macsotay (1980) and Lemoine et al. (1982) to propose that Peregrinella-lenses were rather related to hydrothermal activity in the deepest part of the Vocontian Basin. All Early Cretaceous sites displaying Peregrinella-lenses within the Vocontian Basin were finally linked to circulations of cold seep fluids (e.g., Campbell and Bottjer, 1995b; Kiel et al., 2014).

Peregrinella-lenses are today known worldwide, including Alaska (Sandy and Blodgett, 1996), Italy (Posenato and Morsilli, 1999), Switzerland (Trümpy, 1956), Crimean Peninsula (e.g., Kiel, 2008; Kiel and Peckmann, 2008), Poland (Biernat, 1957), California (e.g., Campbell et al., 2002), Romania (Sandy et al., 2012) and Tibet (Sandy and Peckmann, 2016). Strontium isotope stratigraphy suggests that they range from the late Berriasian to the early Hauterivian (Kiel et al., 2014). Peregrinella brachiopods may have been adapted to diffusive rather than advective seepage and lived at temperatures ranging from 10 to 19°C (Kiel et al., 2014).

2.3 Curnier site

Paquier (1900) did not provide precise geographical and stratigraphical indications for the Curnier site, which thus remained unlocated for decades. A float limestone block with abundant Peregrinella (Peregrinella) multicarinata within a dry creek near Curnier was considered as equivalent to the historical Curnier site. It was resampled, its macrofauna revised and petrographic, stable isotope, and biomarker analyses performed (Kiel, 2013; Kiel et al., 2014). Molluscs accompanying Peregrinella at Curnier, lucinid bivalves Tehamatea vocontiana Kiel 2013 and abyssochrysoid gastropods Humptulipsia macsotayi Kiel et al., 2010, are endemic of methane-seep deposits. Isotopic analyses report a slight δ13C depletion which, coupled with biomarkers, confirm methane seepage, probably through slow and diffuse flow. Lipid biomarkers indicate that seepage fluids may have included crude oil, though it may have intruded the limestone at a later stage. The temperatures reconstructed at Curnier range from 11 to 18°C (Kiel et al., 2014).

Microfauna from Peregrinella-beds within the Vocontian Basin has never been studied in detail. Calpionellids (Stenosemellopsis hispanica (Colom, 1939)), as well as undetermined radiolaria, rare ostracods, benthic and planktonic foraminifera were mentioned in Rottier below and above Peregrinella-lenses and considered as pointing to deposition in an open sea with pelagic influences (Thieuloy, 1972).

3 Samples and methods

The Muséum national d’Histoire naturelle (MNHN) in Paris (France) stores invertebrate fossils sampled in 1976 and 1993 at Curnier. The specimens were gathered by Xavier Marie Philippe Rey-Jouvin, descendant of the glove-maker from Grenoble Xavier Jouvin. He was engineer at the Charbonnages de France (e.g., Rey-Jouvin 1963) and deposited numerous invertebrates in the MNHN collections from 1958 to 1993, mainly from France (e.g., Anonymous 1960, 1961). The Rey-Jouvin collection is composed of 44 Peregrinella brachiopods from Curnier, not including the additional four used for microfossils analysis, two internal moulds of bivalves and one gastropod. The lucinid bivalves (Tehamatea vocontiana; Fig. 2A–C) and abyssochrysoid gastropod (Humptulipsia macsotayi; Fig. 2D) are here pictured to illustrate the entire fauna. All brachiopods from Curnier were macroscopically imaged at the Centre de Recherche en Paléontologie-Paris (CR2P) and measured according to standard rules (Tab. 1).

The collection does not include bulk sediment so that four poorly preserved brachiopod specimens (two isolated valves and two broken shells) have been selected for extraction of microfossils from the enclosed sediment. They have been processed using the hot acetolysis technique (Bourdon, 1962; Lethiers and Crasquin-Soleau, 1988) and residue has been sieved through a 0.63 mm mesh and oven dried. They yielded ostracods, foraminifers, radiolarians, scolecodonts, microbivalves, microbrachiopods, microgastropods (the latter three not being identifiable). All specimens have been picked under stereomicroscope. Specimens of interest have been gold coated and photographed using the SEM JEOL JCM-600 at the CR2P. Stacked images of uncoated specimens were made using the ZEISS AXIO ZOOM V16 and Camera DELTAPIX USB3 20MP (20 stacks per specimen) using green and red UV light at the CR2P. Here we describe the first known ostracods and scolecodonts from Curnier, foraminifera and radiolaria will be considered in a forthcoming work. In the following, we consider the dorsal margin of ostracod carapaces/valves as subdivided into dorsal border (between the posterior and anterior cardinal angles), antero-dorsal border (in front of anterior cardinal angle) and postero-dorsal border (behind posterior cardinal angle). In the same manner, ventral margin is subdivided into ventral border (between the anterior and posterior slope breaks), antero-ventral border (in front of anterior slope break) and postero-ventral border (behind posterior slope break). The length convention of carapaces and valves is as follows: < 0.40 very small, 0.40–0.50 small, 0.50–0.70 medium, 0.70–1.00 large, > 1.0 very large.

All specimens are housed in the Palaeontology collections at the MNHN. All nomenclatural acts of this manuscript are registered in Zoobank at the following link: urn:lsid:zoobank.org:act:782A6BAC-F140-484D-8E81-1F9F37505628.

thumbnail Fig. 2

A–D. Molluscs from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–C. Tehamatea vocontiana Kiel, 2013 (A: MNHN.F.A97899, B, C: MNHN.F.A97898). D. Humptulipsia macsotayi Kiel et al., 2010 (MNHN.F.A97900). E, F. Palurites sp., scolecodonts from sedimentary infill within Peregrinella-shells, in dorsal view (Rey-Jouvin coll.). E. Left maxilla 1, MNHN.F.F73181. F. Right maxilla 1, MNHN.F.F73182. Scale bars: A–D, 1 cm; D, E, 200 µm.
Table 1

Measurements of all specimens of Peregrinella (Peregrinella) multicarinata from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). All are articulated shells except MNHN.F.97893 (ventral valve).

4 Giant brachiopods from Curnier

The present brachiopods are, to the best of our knowledge, the most abundant and well-preserved known from Curnier and entirely composed of specimens of Peregrinella (Peregrinella) multicarinata (Figs. 3 and 4). The 44 specimens range in length from 30.1 to at least 75.5 mm, in width from 29.1 to 73.9 mm and in thickness from 13.8 to 42.5 mm (Tab. 1; Fig. 5). The shells are large, mainly circular and display a variable costate ornamentation. Costae are fine in small specimens with a near planoconvex profile and an anterior margin often with a faint broad sulcation (e.g., Fig. 3A–F), to coarse in the largest specimens with ventribiconvex (e.g., Fig. 4A, B) approaching equibiconvex profile (e.g., Figs. 3Q, 4L, M). The ventral umbo (beak) is massive and incurved (e.g., Figs. 3S, 4C), cut dorsally by a round foramen, which is small for the size of the shell (e.g., Figs. 3E, 4I, N). The lateral margins are rather straight (e.g., Figs. 3B, F, I, 4C, P) and the anterior margin is rectimarginate (e.g., Figs. 3D, P, Q, 4B, M). The shells, often partly worn, show signs of alteration revealing a low dorsal septum (e.g., Figs. 3A, H, P, 4A, G), which supports long curved mergiform crura (Fig. 4R; cf., Manceñido et al., 2002, after Ager, 1968). Several specimens display major growth lines with intervening intermediary or elementary ones (Fig. 4O).

Though juveniles are uncommon (Fig. 5), they illustrate the trend from thin to coarse costae through ontogeny. It is nonetheless interesting that shape and size of costae is variable among pre-adults and adults, some of them also displaying relatively thin costae (e.g., Fig. 4C–E). Conversely, some of the smallest specimens (not necessary juveniles) display relatively coarse costae in regard to their size (e.g., Fig. 3D–F). The material at hand is however not sufficient to further discuss and interpret this pattern. The link between the ontogeny of Peregrinella specimens and their distance from the active seepage zone during their lifetime should be tested in the future. It is also not possible to exclude that the specimens at hand correspond to successive generations and/or populations that may illustrate different palaeoenvironmental conditions or seepage stages.

In spite of their reputed unpalatability (e.g., Thayer and Allmon, 1991), brachiopods are often targets for predators (gastropods for example) and may be important prey in habitats where molluscs are rare or absent (e.g., Harper, 2011; Tyler et al., 2013). At least 55% of the shells of the Curnier collection bear traces relatable to healed shell injuries, not considering marks that could not be undoubtedly identified (Figs. 3 and 4). They are seen on both juveniles and adults (Fig. 5), some shells displaying up to three distinct traces. Most are long, wide and deep marks occurring symmetrically on both valves (e.g., Figs. 3M, N, 4D–J, L–N). They seem to be preferentially located in the posterior (39%) and central (39%) portions of the valves, anterior marks being rarer (22%). The rhynchonellids survived the attacks that can be considered sublethal as the secretion of the shell went on after the attack. As the thickness of each valve was not important (Fig. 4R), it is likely that the predators themselves were not particularly large.

Injuries on shells of Peregrinella (Peregrinella) multicarinata have previously been reported from various sites. Biernat (1957) observed symmetrical damages on adults and juveniles from France and Poland. Straight cuts, convex cuts on the lateral shell margins, and distinct holes on shells from the Hauterivian of Crimean Peninsula in southern Ukraine have also been interpreted as healed shell injuries (Kiel and Peckmann, 2008). Rectilinear punctures composed of four small depressions and deep straight cuts, both symmetrical on both valves, have been reported from the Upper Sinaia Formation (late Hauterivian-early Barremian) exposed in Eastern Carpathian Mountains, Romania (Sandy et al., 2012). It is considered that predation rates on brachiopods from chemosynthetic environments are much higher than in normal marine settings, decapod crustaceans being the most probable predators although their remains rarely occur in the assemblages (Sandy, 2010). Alternatively, scars could possibly result from the action of cephalopod jaws (Biernat, 1957; Sandy, 2010).

Non-lethal shell damages such as those observed on Peregrinella (Peregrinella) multicarinata specimens are often attributed to failed durophagous attacks (e.g., Vermeij et al., 1981; Alexander and Dietl, 2003). Klompmaker et al. (2019) summarized the features distinguishing predation, non-predatory biotic causes from abiotic causes of non-lethal damage, including on invertebrate shells from past and modern cold seeps. Among these causes, compaction related to the abundance of specimens forming Peregrinella-lenses may have caused generalized morphological disruptions of the shells (Harper, 2005), as well as fractures or breakage that were neither observed in previous works (e.g., Vörös, 2009; Sandy et al., 2012) nor in the present material. The symmetry of the traces observed on Curnier specimens are rather evocative of crustacean claws, similar to marks observed on shells of the extant terebratulid Gryphus vitreus (Born, 1778) in the Mediterranean Sea related to lobsters cracking the fine anterior part of the three-layered shells (e.g., Boullier et al., 1986). Similar traces of unsuccessful crushing attacks were also for instance observed on extant brachiopods of Antarctic Peninsula, Falkland Islands, Chile (e.g., Harper et al., 2009; Harper and Peck, 2016) and fossils through the Mesozoic and Cenozoic (e.g., Harper, 2005; Vörös, 2009). However, remains of macro-crustaceans have never been observed at Curnier and we did not observe microcoprolites, in spite of their importance to evaluate the role of crustaceans in chemosynthetic ecosystems (Senowbari-Daryan et al., 2007). It is also interesting that brachiopods of all sizes were attacked in Curnier, including the largest ones (Fig. 5), contrasting with the idea that large size could be a refuge from predation (Harper et al., 2009). Occurrence of traces in all size classes of modern brachiopods was considered as indicative of abiotic factors or human intervention (Harper et al., 2009) but the observed morphology of the traces precludes such interpretation at Curnier.

thumbnail Fig. 3

Peregrinella (Peregrinella) multicarinata (Lamarck, 1819) from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–C. Dorsal, lateral and anterior views of a juvenile specimen with septum visible on the worn surface (MNHN.F.A97897). D–F. Anterior, dorsal and lateral views of a juvenile with coarse costae (MNHN.F.A97877). G. Dorsal view of specimen MNHN.F.A97871. H–J. Dorsal, lateral and anterior views of MNHN.F.A97881. K, L. Anterior and dorsal views of MNHN.F.A97856 with suspicion of foramen. M, N. Dorsal and ventral views of MNHN.F.A97866. O, P. Dorsal and anterior views of MNHN.F.A97857. Q–S. Anterior, dorsal, lateral views of MNHN.F.A33863. Black arrows: septum. Red arrows: predation marks. Scale bars: 1 cm.
thumbnail Fig. 4

Peregrinella (Peregrinella) multicarinata (Lamarck, 1819) from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A, B. Dorsal and anterior views of MNHN.F.A97862 with coarse costae and septum visible on a worn surface. C–E. Lateral, dorsal and ventral views of the relatively gibbous MNHN.F.A97869 specimen with marks of predation on both valves. F–H. Anterior, dorsal and lateral views of an equibiconvex specimen with relatively thin costae MNHN.F.A97865. I–L. Dorsal, ventral, lateral and anterior views of MNHN.F.A97864 with remaining parts of calcite shell and coarse costae. M–P. Anterior, dorsal, close up on major growth lines near the antero-lateral margin and lateral views of MNHN.F.A97863. Q. Large isolated ventral valve with coarse costae (MNHN.F.A97893). R. Transverse section trough a large specimen revealing the septum and beginning of crura (MNHN.F.A98232). Black arrows: septum. Red arrows: predation marks. Yellow arrows: growth lines. Scale bars: 1 cm.
thumbnail Fig. 5

Scatter plots of the size of Peregrinella (Peregrinella) multicarinata at Curnier (Rey-Jouvin coll.) showing the width (top) and thickness (bottom) as a function of length of the shells. Drawings from d’Orbigny (1847).

5 Ostracods from Curnier

5.1 Systematic palaeontology

Phylum Arthropoda Gravenhorst, 1843

Class Ostracoda Latreille, 1802

Suborder Cypridocopina Jones, 1901

Superfamily Pontocypridoidea Müller, 1894

Family Pontocyprididae Müller, 1894

Genus Nekrocypris Forel gen. nov.

Zoobank link. urn:lsid:zoobank.org:act:782A6BAC-F140-484D-8E81-1F9F37505628.

Type species. Nekrocypris sepultinconcha Forel gen. nov. sp. nov., by monotypy.

Diagnosis. A small Pontocyprididae genus characterized by Pontocypris-like subtriangular valves with produced posterior end pointing downward and left valve narrowly overlapping right one with major antero-ventral offset. Internally, calcified inner lamella wide anteriorly and subcentral muscle scars composed of at least four scars in a packed circular pattern.

Etymology. From the Greek νεκρóς (nekrós), for “dead”, referring to the specimens being preserved within the shells of Peregrinella brachiopods. Gender of the genus: feminine.

Remarks. The subtriangular elongate cyprid outline of Nekrocypris carapace relates it to Cypridoidea, and more precisely to Pontocyprididae or Paracyprididae Sars, 1923. All specimens being articulated carapaces, their hinge could not be observed but the use of green UV fluorescence reveals a subcircular central muscle scar pattern indeed resembling that of Pontocyprididae or Paracyprididae (see Fig. 7 in Maddocks, 1969 for a summary). In spite of the recrystallization of carapaces, at least four wedge-shape individual scars are observed (Fig. 7D, E). Because of their compact arrangement and absence of a cap scar (sensu Maddocks, 1969), we favour Pontocyprididae as they are characterized by a simple arrangement of five adductor scars, though six scars have been reported in the extant Peripontocypris Wouters, 1997 and Tabukicypris Chiu et al., 2015 (Maddocks, 1969; Wouters, 1997; Chiu et al., 2015). The UV fluorescence also highlights the calcified inner lamella which is wide anteriorly but imperceptible ventrally and posteriorly, as for instance in Argilloecia Sars, 1866 and Australoecia McKenzie, 1967. Pontocyprididae are also characterized by right valve overlapping left one. Nekrocypris has a reversed overlap and thus belongs to the few genera of the family with such a character, including the extant Maddocksella McKenzie, 1982, some species of Australoecia McKenzie, 1967 and the fossil Pontocyprella Mandelstam in Ljubimova, 1955.

Nekrocypris differs from all Pontocyprididae by the combination of its produced posterior end and reversed overlap with antero-ventral offset. Among fossil Pontocyprididae, Nekrocypris differs from the Jurassic Liasina Gramann, 1963 (senior synonym of the recent Iliffeoecia Maddocks, 1991 according to Wouters, 1997) by its Pontocypris-like outline, while Liasina has an Argilloecia-outline with thicker overlap. Nekrocypris furthermore lacks the sack-like anterior vestibule of Liasina. Nekrocypris differs from the overall Mesozoic Pontocyprella by its smaller size and general outline of cyprid morphology with produced posterior end pointing downward. It also differs from Pseudomacrocypris Michelsen, 1975, known from the Late Triassic to the Early Cretaceous (Maddocks, 1991; Forel and Grădinaru, 2020), by left valve overlapping right one and produced posterior end pointing downward.

Nekrocypris sepultinconcha Forel gen. nov. sp. nov.

Figs. 6C–, 7A–E, 8A

Zoobank link. urn:lsid:zoobank.org:act:782A6BAC-F140-484D-8E81-1F9F37505628.

Type material. Holotype: MNHN.F.F73159, an articulated carapace (Figs. 6H–K, 8A). Paratype: MNHN.F.F73160, an articulated carapace (Fig. 6L–P).

Additional material. 15 articulated carapaces.

Type locality and horizon. Peregrinella-lenses, Curnier, Drôme department, France; early Hauterivian, Early Cretaceous.

Etymology. From the Latin sepulta in concha, for “buried in a shell”, referring to the specimens being preserved within shells of Peregrinella brachiopods. By apposition.

Diagnosis. A small, elongate, fusiform Nekrocypris species with thin ridges radiating from posterior end. The upper ridges do not extend past the posterior cardinal angle while lower ones extend to the posterior part of the antero-ventral offset of overlap. A short, thick ridge develops perpendicular to the anterior margin, at maximum of curvature.

Description. In lateral view, carapace small, elongate, fusiform with produced posterior end, maximum of height at anterior cardinal angle and maximum of length close to ventral margin. In dorsal view, carapace biconvex with maximum of width very slightly posterior to mid-length and narrow constriction in front of it, likely corresponding to the central muscle scar area. Left valve narrowly overlaps right one along dorsal and ventral margins (Fig. 6C, H, Q, S, U). An antero-ventral offset of the overlap develops below anterior cardinal angle, clearly visible in ventral (Figs. 6F, J, N, R, 8A) and lateral views (Figs. 6C, H, L, Q, S, U, 8A), even in poorly preserved specimens (Fig. 7A, C).

Dorsal margin largely convex with right valve slightly more tripartite than left one, with poorly defined cardinal angles. Anterior cardinal angle around anterior 1/3 of length, delimiting straight to slightly convex antero-dorsal border steeply sloping anteriorly (30–35°) and short, straight dorsal border gently sloping posteriorly (10–15°). Posterior cardinal angle weakly expressed at most specimens, around posterior 1/4 of length, delimiting straight postero-dorsal border steeply sloping downward (35–40°). Ventral margin long and sinuous with concavity behind mid-length. Antero-ventral margin short, only moderately raised upward and postero-ventral margin sloping downward. Anterior margin laterally compressed with gentle angulation below mid-height. A thick, short ridge extends perpendicular to anterior border, starting from angulation, with beak-like morphology when thinner surrounding parts are weathered (Figs. 6H, L, Q, S, T, 7A, B, 8A). Posterior border narrow, at or slightly below ventral margin and pointing downward. Laterally, thin ridges radiate from posterior end: ventralmost ridges extend across mid-length to ventral offset of overlap, dorsalmost ones do not extend past posterior cardinal angle.

Adductor muscle scars composed of at least four individual scars organized in a packed circular pattern located slightly posterior to mid-length and around mid-height (Fig. 7D). Individual scars wedge-shape, elongated along an antero-dorsal and postero-ventral axis (Fig. 7E). Calcified inner lamella wide along anterior margin, less so posteriorly (Fig. 7D). Sexual dimorphism not observed.

Dimensions (Fig. 8A). Holotype: length = 434 µm; height = 182 µm; width = 157 µm. Paratype: length = 421 µm; height = 187 µm; width = 159 µm.

Remarks. Nekrocypris sepultinconcha is the second report of an ornamented Pontocyprididae, with the Holocene Tabukicypris decoris Chiu et al., 2015 (type species and only member of Tabukicypris Chiu et al., 2015. Nekrocypris nonetheless lacks the flat ventral surface of Tabukicypris and has a downward pointing posterior end. The height/length scatter plot of all measurable specimens of Nekrocypris sepultinconcha documents at least two ontogenetic stages (Fig. 8A).

Occurrences. Peregrinella-lenses, Curnier, Drôme department, France; early Hauterivian, Early Cretaceous (this work).

Suborder Cytherocopina Baird, 1850

Superfamily Cytheroidea Baird, 1850

Family Cytheruridae Müller, 1894

Subfamily Eucytherurinae Müller, 1894 emend. Maddocks and Steineck, 1987

Genus Eucytherura Müller, 1894 emend. Horne and Lord, 2024

Subgenus Vesticytherura Gründel, 1964

Type species. Eucytherura neocomiana Kaye, 1964, subsequently designated by Gründel (1964).

Remarks. Vesticytherura Gründel, 1964 was described as a subgenus of Eucytherura Müller, 1894 to accommodate species with a distinct anterior vestibule, seven to 10 straight anterior and three posterior marginal pore canals, dorsal ridge replaced by nodes and median ridge developed only in the anterior region. It was later raised to genus level by Gründel (1981) and ascribed to Eucytherurinae, as was Eucytherura, by Maddocks and Steineck (1987). Opinions on the status of Vesticytherura have varied, from justified genus (e.g., Whatley, 1970; Malz et al., 1985; Maddocks and Steineck, 1987; Tesakova et al., 2008; Piovesan et al., 2014) to unnecessary (Whatley and Boomer, 2000), while others went on considering it as a subgenus of Eucytherura (Slipper, 2021). Morphological transitions between Eucytherura, Renicytherura Gründel, 1981 and Vesticytherura led Franz et al. (2018) to subsume all three genera into Eucytherura. It is beyond the scope and intention of this paper and its material to attempt going further in this debate. However, we consider that the morphological differences distinguishing Eucytherura from Vesticytherura are sufficient to consider that they should not be synonymized. We thus follow the most recent taxonomic largescale investigation of Cretaceous taxa by Slipper (2021) and consider Vesticytherura as a subgenus of Eucytherura.

Eucytherura (Vesticytherura) captinconcha Forel sp. nov.

Figs. 7I–S, 8B

1979 Eucytherura sp. B 377; Donze in Busnardo et al., p. 78, pl. 4, Fig. 2–4.

Zoobank link. urn:lsid:zoobank.org:act:782A6BAC-F140-484D-8E81-1F9F37505628.

Type material. Holotype: MNHN.F.F73174, an articulated carapace (Figs. 7O, 8B). Paratype: MNHN.F.F73170, an articulated carapace (Fig. 7J).

Additional material. 24 articulated carapaces.

Type locality and horizon. Peregrinella-lenses, Curnier, Drôme department, France; early Hauterivian, Early Cretaceous.

Etymology. From the Latin capti in concha, for “trapped in a shell”, referring to the specimens preserved within shells of Peregrinella brachiopods. By apposition.

Diagnosis. A reticulate and ridged Eucytherura (Vesticytherura) with long, deep, sinuous sulcus and three dorsal nodes.

Description. Carapace very small, elongate, ear-shaped subtriangular in lateral view, strongly tapered postero-ventrally with maximum of height at the anterior cardinal angle and maximum of length around mid-height. In dorsal view, carapace weakly inflate, lanceolate with maximum of width behind mid-length at postero-ventral node. Dorsal margin straight to very gently sinuous, stepped posteriorly toward caudate posterior border located only slightly below dorsum. Anterior border largely convex with maximum at or below mid-height. Ventral margin mostly obscured by lateral inflation, straight and raised posteriorly towards posterior border.

Lateral surface bearing an oblique median sulcus and three circular subdorsal nodes, from front to back (N1–N3; Fig. 8B):

  • anterior node (N1) below dorsal margin directly behind anterior cardinal angle.

  • behind N1, a sinuate, narrow, long and deep sulcus extends from dorsal margin to lower 1/4 of height, just in front of mid-length.

  • median node N2 slightly raised above dorsum, directly behind sulcus.

  • posterior node N3 slightly raised above dorsum, in front of postero-dorsal angulation, posterior to postero-ventral node.

Ventrally, postero-ventral node located between N2 and N3. Juvenile carapace with well-expressed sulcus, subdorsal and postero-ventral nodes (Fig. 7L).

Lateral surface reticulated with large and shallow pentagonal fossae separated by thin muri, solum of each fossa bearing eight to 15 secondary pits. Except narrow area in front of marginal ridge R1, entire lateral surface reticulated, including subdorsal nodes. Reticulation weaker through muscular sulcus but still visible as ridges on adults and secondary pits on juveniles.

At least four prominent ridges recognizable through reticulation, variably expressed through ontogeny (R1–R4; Fig. 8B):

  • marginal ridge R1 delimiting the reticulated area, parallel to anterior margin, only slightly removed from it.

  • two subparallel anterior ridges R2 and R3, separated by two rows of fossae. R2 emerges from N1, runs downward and rapidly turns anteriorly to reach R1 above mid-height. R3 runs obliquely from pore conuli P2 and reaches R1 below mid-height.

  • curvate circum-sulcus ridge R4 emerges antero-ventrally in a loop below sulcus and rises upward as a single branch toward N1. It is fully expressed on small juveniles and only seen below sulcus and toward N1 on larger specimens, supposedly adults.

Five conjunctive pore conuli are scattered over the lateral surface (P1–P5; Figs. 7N, 8B): P1 on R2 below N1, P2 below and behind N1, P3 subdorsally behind N1 and in front of sulcus, P4 at postero-ventral tip of sulcus, P5 above postero-ventral nodule.

Dimensions (Fig. 8B). Holotype: length = 338 µm; height = 171 µm. Paratype: length = 317 µm; height = 168 µm.

Remarks. Donze in Busnardo et al. (1979) illustrated Eucytherura (Vesticytherura) captinconcha, identified as Eucytherura sp. B 377, from upper Valanginian deposits from Hautes-Alpes department in south-eastern France. They described it as “remarkable by its three tubercles on the dorsal border of each valve” (p. 78 in Busnardo et al., 1979). Noteworthy, the reticulation was not mentioned but is visible on the illustrated specimens. Even when badly preserved specimens no longer display reticulation, the morphology of Eucytherura (Vesticytherura) captinconcha is easily recognizable based on its three subdorsal nodes and well-expressed sulcus (e.g., Fig. 7Q–S).

The height/length scatter plot of all measurable specimens of Eucytherura (Vesticytherura) captinconcha from Curnier documents at least two ontogenetic stages (Fig. 8B). The size of the largest specimen (length = 363 µm; height = 175 µm; Fig. 7R) is consistent with that of fossil and recent Eucytherura species (e.g., Ayress et al., 1995; Ballent and Whatley, 2009). We thus consider the largest specimens, including holotype and paratype, as adults. Two morphologies are distinguished among specimens of both scatter plots:

  • Morphology 1 with subrectangular carapace, anterior maximum at mid-height, posterior end strongly stepped with caudate extremity below dorsum (Fig. 7Q, R). The smallest specimen at hand, though broken posteriorly, seems to illustrate this morphology (Fig. 7L).

  • Morphology 2 with subtriangular carapace, anterior maximum below mid-height and posterior end strongly tapered ventrally with apex at dorsum (e.g., Fig. 7I, J, O). This morphology is also expressed among the smallest specimens (Fig. 7S).

Males of extant cytherurids are generally more elongate and inflated than females, although this is not universal to all species (e.g., Weingeist, 1949; Whatley et al., 1988; Witte, 1993; Brouwers, 1994; Ayress et al., 1995; Ramos et al., 1999; Jöst et al., 2022). This morphological pattern occurs for both extant and fossil Eucytherura, males generally being more subrectangular and elongate laterally (e.g., Ayress et al., 1995; Ballent and Whatley, 2009). We thus consider that the two morphologies of Eucytherura (Vesticytherura) captinconcha illustrate sexual dimorphism, morphology 1 likely corresponding to males while morphology 2 may correspond to females.

Among Eucytherura species with subdorsal nodes, here considered as members of the subgenus Vesticytherura, the new species differs from Eucytherura ansata Weingeist, 1949 from the Albian of Texas (Weingeist, 1949) by its circular rather than elongated nodes, long and well-developed sulcus, lack of a second node in front of sulcus, of antero-ventral tubercles and of keel-like ventral ridge. Eucytherura (Vesticytherura) captinconcha differs from Eucytherura dorsotuberculata van Veen, 1938 from the Maastrichtian of Netherlands (van Veen, 1938) by its sulcus, reticulation and lack of node anterior to sulcus. The new species also differs from Eucytherura multituberculata Gründel, 1964 from the Early Cretaceous of Germany (Gründel, 1964) and Eucytherura trinodosa Pokorný, 1973 from the Late Jurassic of Czech Republic (Pokorný, 1973) by its reticulate surface, sulcus and lack of a subdorsal node directly in front of sulcus. We take this opportunity to note that Eucytherura multituberculata Ayress et al., 1995 described from the Early Pliocene of Lord Howe Rise (Ayress et al., 1995) is a junior homonym of Eucytherura multituberculata Gründel, 1964, and a replacement name should be introduced following ICZN rules.

Occurrences. Route d’Angles and Barret-le-Bas sections, Hautes-Alpes department, France; upper Valanginian, Early Cretaceous (Donze in Busnardo et al., 1979). Peregrinella-lenses, Curnier, Drôme department, France; early Hauterivian, Early Cretaceous (this work).

Family Pleurocytheridae Mandelstam, 1960

Genus Vocontiana Donze, 1968

Type species. Vocontiana longicostata Donze, 1968 by original designation.

Remarks. Vocontiana Donze, 1968 was described from the Berriasian-Valanginian transition in Berrias (Ardèche, France; Donze, 1968). It was questionably placed within the Paradoxostomatidae Brady and Norman, 1889 and was more recently attributed to Pleurocytheridae (e.g., Savelieva, 2014), a position we follow. Colin (1974) proposed, without discussion, that Vocontiana might be a junior synonym of Annosacythere Kuznetsova, 1957. Until a detailed analysis of the type material of both genera can be performed, we consider Vocontiana as a valid genus, in line with the current consensus.

?Vocontiana sp.

Fig. 7T

Studied material. MNHN.F.F73179, an articulated carapace. Right valve: length = 319 µm; height = 154 µm. Left valve: length = 319 µm; height = 158 µm.

Remarks. ?Vocontiana sp. is rare in the present material and the unique carapace at hand precludes in-depth discussion. Its outline and ornamentation are reminiscent of Vocontiana but the specimen lacks the eye node of Vocontiana and has a maximum of height at mid-length, while it is generally at the antero-dorsal angulation on Vocontiana. Vocontiana is furthermore characterized by more or less sinuous longitudinal ridges (Donze, 1968): they are here seen ventrally and posteriorly, and replaced by large reticulation in front of mid-length. ?Vocontiana sp. is undeniably new to science but more specimens are needed to describe its characters and fully understand its generic placement.

Occurrences. Peregrinella-lenses, Curnier, Drôme department, France; early Hauterivian, Early Cretaceous (this work).

thumbnail Fig. 7

Ostracods from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–E. Nekrocypris sepultinconcha Forel gen. nov. sp. nov. A, B. Articulated carapace broken posteriorly (MNHN.F.F73164) in right view (A) and left view (B). C–E. Articulated carapace broken posteriorly (MNHN.F.F73165) in right view (C), in left view seen in green UV light with white arrows showing the limits of the anterior calcified inner lamella and dashed square showing the adductor muscle scars (D), enlarged in (E). F. Pontocyprella sp. 1, right view of an articulated carapace (MNHN.F.F73166). G. Pontocyprella sp. 2, right view of an articulated carapace (MNHN.F.F73167). H. ?Pseudomacrocypris sp., right view of an articulated carapace (MNHN.F.F73168). I–S. Eucytherura (Vesticytherura) caspinconcha Forel sp. nov. I. Articulated carapace MNHN.F.F73169, right view. J. Paratype, articulated carapace MNHN.F.F73170, right view. K. Articulated carapace (MNHN.F.F73171) in dorsal view, with anterior to the left. L. Articulated juvenile carapace broken anteriorly (MNHN.F.F73172) in right view. M, N. Articulated carapace broken posteriorly (MNHN.F.F73173) in left view (M), with close-up on surface ornamentation and pore conuli P2, P4, P5 (N). O. Holotype, articulated carapace (MNHN.F.F73174) in left view. P. Articulated carapace broken posteriorly (MNHN.F.F73175) in right view. Q. Articulated carapace (MNHN.F.F73176) in left view. R. Articulated carapace (MNHN.F.F73177) in right view. S. Articulated carapace (MNHN.F.F73178) in right view. T. ?Vocontiana sp., right view of an articulated carapace (MNHN.F.F73179). U. Polycope sp., right view of an articulated carapace (MNHN.F.F73180). Scale bars: 100 µm except E: 50 µm, N: 20 µm.
thumbnail Fig. 6

Ostracods from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A. ?Macrosarisa sp., articulated carapace, right view (MNHN.F.F73156). B. Paracypris sp., articulated carapace, right view (MNHN.F.F73157). C–U. Nekrocypris sepultinconcha Forel gen. nov. sp. nov. C–G. Articulated carapace (MNHN.F.F73158) in right view (C), detail of the posterior end (D), left view (E), ventral view with anterior to the left (F), dorsal view with anterior to the right (G). H–K. Holotype, articulated carapace (MNHN.F.F73159) in right view (H), left view (I), ventral view with anterior to the right (J), dorsal view with anterior to the right (K). L–P. Paratype, articulated carapace (MNHN.F.F73160) in right view (L), left view (M), ventral view with anterior to the left (N), detail of the posterior end (O), dorsal view with anterior to the left (P). Q, R. Articulated carapace broken posteriorly (MNHN.F.F73161) in right view (Q) and ventral view with anterior to the right (R). S, T. Articulated carapace (MNHN.F.F73162) in right view (S) and left view (T). U. Articulated carapace, right view (MNHN.F.F73163). Scale bars: 100 µm except D: 20 µm, O: 50 µm.
thumbnail Fig. 8

A. H/L scatter plot of Nekrocypris sepultinconcha Forel gen. nov. sp. nov. and line drawing in right view (top) and ventral view (bottom). B. H/L scatter plot of Eucytherura (Vesticytherura) caspinconcha Forel sp. nov. and line drawings of left views of holotype (top, Fig. 7O) and juvenile (bottom, Fig. 7L), including subdorsal nodes (N1–N3), ridges (R1–R4) and pore conuli (P1–P5). Scale bars: 100 µm.

5.2 Taphonomy and diversity

The sediment infilling Peregrinella shells from Curnier provided a low abundance ostracod assemblage of 57 identifiable specimens, together with several fragments and unidentifiable specimens. Their taxonomic diversity is relatively low with nine species distributed into eight genera and six families (Tab. 2). How much does this assemblage illustrate the original ostracod fauna? To answer this question, all specimens of each species were counted and individual rarefaction was calculated with PAST version 4.04 (Hammer et al., 2001; Hammer and Harper, 2005), showing the expected number of species as a function of the number of specimens in the community (Fig. 9A). The rarefaction curve indicates that the species count is not representative of the entire fauna and that a larger sample would have given better counts and a higher diversity level. Although the curve is far from flattening out, its slope indicates that the assemblage at hand gathers an important proportion of the original set of species.

The proportion of articulated carapaces vs isolated valves and the demographic structure of populations are important indicators of the autochthonous or allochthonous nature of ostracod assemblages (e.g., Oertli, 1971; Boomer et al., 2003). At Curnier, all specimens are articulated carapaces, including unidentifiable and broken ones. Because of the low abundance of the assemblage, its demographic structure can only be cautiously discussed. Nonetheless, at least two submature to mature ontogenetic stages of Nekrocypris sepultinconcha (Fig. 8A) and Eucytherura (Vesticytherura) caspinconcha (Fig. 8B) are recovered, with lack of small juveniles. Altogether, these observations indicate that transportation occurred, though limited, and that the Curnier ostracods correspond to a truncate community that was living nearby to the brachiopod shells they were recovered in.

When considering the composition of the assemblage, the generic diversity (species per genus) is dominated by monospecific genera (11% each: Eucytherura, ?Macrosarisa, Nekrocypris, Paracypris, Polycope, Pseudomacrocypris, ?Vocontiana). Only Pontocyprella is represented by two species (22%). Though of low abundance, richness patterns can be distinguished among species and families, species and genus richness patterns being similar. In terms of specimens per species, the assemblage is dominated by Eucytherura (Vesticytherura) caspinconcha (46% of specimens) and Nekrocypris sepultinconcha (30%) (Fig. 9B, left panel). All other species are secondary (Pontocyprella sp. 1, Pontocyprella sp. 2: 7% each) and rare (Paracypris sp.: 4%; ?Vocontiana sp., ?Macrosarisa sp., Polycope sp., ?Pseudomacrocypris sp.: 2% each). When considering the overall familial abundance (i.e., number of specimens per family), Pontocyprididae (Nekrocypris, Pontocyprella, Pseudomacrocypris) and Cytheruridae (Eucytherura) co-dominate the assemblage, each being 46% of the specimens (Fig. 9B, right panel).

Table 2

Taxonomic list of ostracod species identified from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.).

thumbnail Fig. 9

A. Individual rarefaction curve for the ostracod assemblage from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). B. Circular diagrams summarizing the composition of the ostracod assemblage by number of specimens per species (left) and family (right).

5.3 Palaeoecology of ostracods

The ostracods here studied were extracted from sediment infilling the valves of four Peregrinella shells: were they scavenging on the decaying flesh or members of the Curnier chemosynthetic community and transported inside the shells post-mortem?

Scavenging is a common feeding strategy among modern nektobenthic Myodocopida, more specifically Cypridinidae, that form swarms around decaying carcasses, sometimes gathering thousands of specimens (e.g., Cohen, 1983, 1989; Collins et al., 1984; Stepien and Brusca, 1985; Vannier and Abe, 1993; Keable, 1995; Parker, 1997; Vannier et al., 1998). Swarms of scavenging Podocopida, Cyprididae, have also been documented from freshwater environments (e.g., Kiefer, 1936; Reichholf, 1983; Meisch, 2000) but scavenging by marine Podocopida has conversely rarely been reported. Among Cytheruridae, that are important components of the Curnier assemblage, species of Cytheropteron Sars, 1866 are carnivorous, scavengers or predators on polychaeta (Hartmann, 1975). Xylocythere Maddocks and Steineck, 1987 from deep-sea sunken wood may feed on chemoautotrophic bacteria, be necrophagous, coprophagous scavengers, symbiotic, commensal or parasitic (Van Harten, 1992). Pontocyprididae, the only known family of Pontocypridoidea, are generally detritus feeders, some species also being commensal and symbiotically associated with echinoderms (Maddocks, 1968, 1979, 1987). Species of Thomontocypris Maddocks, 1991 associated with tubeworms from hydrothermal vents in the Pacific Ocean are likely deposit feeders living from detritus, mucus secretions, or fecal pellets of sessile organisms (Maddocks, 2005; Tanaka and Yasuhara, 2016). Propontocypris Sylvester-Bradley, 1947 has been suggested to feed on decaying plant or animal tissues, mucus secretions, bacterial slime, or fecal pellets (Maddocks and Steineck, 1987). The morphology of Tabukicypris may be indicative of scavenging activities in a dark environment (Chiu et al., 2015).

In the fossil record, evidence of scavenging ostracods is rare and the most convincing reports are swarms of nektobenthic Myodocopida associated with carcasses during the Ordovician, Silurian, Carboniferous and Triassic (e.g., Wilby et al., 2005; Wilkinson et al., 2007; Perrier et al., 2011). The myodocopid Juraleberis jubata Vannier and Siveter, 1995 from the Tithonian of Russia proposed to have been scavenging on remains of a pliosaur (Dzik, 1978; Boucot, 1990) was finally demonstrated as filter feeding (Vannier and Siveter, 1995). Similarly, the proposed scavenging lifestyle of the Early Cretaceous cypridid Pattersoncypris micropapillosa Bate, 1972 from Brazil (Bate, 1971, 1972) was ruled out and detritus feeding was rather considered likely (Smith, 2000). No example of scavenging marine Podocopida has been formally reported from the fossil record.

At Curnier, the low abundance of ostracods is far from the high-density populations expected in relation to scavenging activities, but appears to be largely natural based on rarefaction curve, as discussed. Myodocopida are furthermore lacking and none of the taxa at hand can formally be related to scavenger groups in modern or past marine environments. All in all, the ostracod specimens recovered from within the Peregrinella shells were likely part of the Curnier chemosynthetic community and were washed into the shells post-mortem. Though feeding on living Preregrinella specimens can’t be ruled out, we rather hypothesize that Nekrocypris, so far only known from Curnier, had a soft diet similar to that of modern Propontocypris and Thomontocypris, possibly related to faecal pellets and other detritus of associated organisms.

The occurrence of secondary pits within the reticulation of Eucytherura (Vesticytherura) caspinconcha (e.g., Fig. 7N) is evocative of pore clusters, that have been proposed to be related to ectosymbiosis with chemosynthetic bacteria in the context of extreme environments (Van Harten, 1993; Maddocks, 2005; Karanovic and Brandão, 2015; Yasuhara et al., 2018; Tanaka et al., 2019, 2021). Though diagnostic of Eucytherurinae (see discussion in Horne and Lord, 2024), the formal occurrence of pore clusters in Early Cretaceous cold seeps would have crucial implications on the understanding of the evolution and adaptation of ostracods to such environments. Until these features can be further studied on well-preserved specimens and their link with ectosymbiosis verified, we rather consider that Eucytherura (Vesticytherura) captinconcha was a deposit-feeder, possibly linked to Peregrinella faeces. The palaeoecology of ostracods at Curnier would thus be characterized by deposit-feeding species likely relying on organic matter issued from the abundant Peregrinella specimens. Such a chemosynthetic community would thus differ from that of the older Sahune seep site (middle Oxfordian, Late Jurassic) where the cytherurid Procytherura? praecoquum Forel in Forel et al., 2024, likely endemic to the active centre of fluid emission, may display the oldest pore clusters known to date, possibly illustrating ectosymbiosis (Forel et al., 2024, 2025).

5.4 Palaeoenvironmental discussion

Pontocyprididae are marine ostracods today distributed from shallow littoral to deep-sea areas (e.g., van den Bold, 1974; Maddocks, 1969, 1977; Karanovic, 2019). Within the Vocontian Basin, they were largely represented by species of Pontocyprella that were members of oligotrophic deep-sea faunas during the Early Cretaceous (e.g., Donze, 1971; Donze and Lafarge, 1979; Donze in Busnardo et al., 1979; Scarenzi-Carboni, 1984; Babinot et al., 1985). Abundant specimens of Pontocyprella were also cornerstones of the Sahune seepage communities at bathyal depth during the Late Jurassic (Forel et al., 2024, 2025). Pontocyprididae are also key components of the Curnier community but Pontocyprella species are only secondary, and their dominance is replaced by that of Nekrocypris (Fig. 9B). Nekrocypris specimens however do not proliferate and the largely natural low abundance of this assemblage likely excludes taphonomic bias. These different abundances between two diachronic seep communities within the same basin may illustrate 1) different moments of the evolution of the basin, which may include different fluids such as the possible emission of crude oil in Curnier (Kiel et al., 2014), 2) different fluxes, reported diffuse in Curnier (Kiel et al., 2014), 3) different palaeoecological features, with abundant Peregrinella brachiopods structuring the ecosystem in Curnier. Nekrocypris is unknown from any other locality within the Vocontian Basin, which limits its palaeoenvironmental significance at that stage, but indicates that it may be specifically adapted to Peregrinella-dominated ecosystems. The high diversity of Pontocyprididae at Curnier (46% of species; Fig. 9B) is essentially natural and even higher than in Sahune (27%; Forel et al., 2024). Such a feature is unknown from other Hauterivian platform communities (e.g., Stchépinsky, 1955; Neale, 1960, 1962; Kaye, 1963a, b, 1964; Luppold, 2001; Schudack, 2004; Musacchio and Simeoni, 2008) but was only observed from bathyal environments during the Middle Jurassic (Monostori, 1995) and Early Cretaceous (Scarenzi-Carboni, 1984; Ayress and Gould, 2018). Overall, the observed high diversity of Pontocyprididae appears a typical feature of bathyal ostracod communities during this period. Conversely, the high abundance of ostracods and more specifically Pontocyprididae in Sahune may have been related to cold seep fluids and associated complex ecosystems (Forel et al., 2025). The comparatively poor population in Curnier may relate to palaeoenvironmental or palaeoecological features that remain to be determined.

Today, Cytheruridae are widespread in shelf and deep-sea environments as they were during the entire Cenozoic (e.g., Ayress, 1995; Ayress et al., 1995; Titterton and Whatley, 2006; Yasuhara et al., 2009; Jöst et al., 2022). It is thought that they were mainly confined to shelf and marginal areas during the Mesozoic (Ballent and Whatley, 2009), though they also occur in bathyal environments since the Late Triassic (e.g., Donze, 1975; Forel et al., 2019). At Curnier, they are as important as Pontocyprididae in terms of species per family (Fig. 9B), reminiscent of the ostracod community at Sahune seep (Forel et al., 2024, 2025). Eucytherura (Vesticytherura) captinconcha is the most abundant species at Curnier (Fig. 9B) and was previously reported in open nomenclature from upper Valanginian sub-bathyal to bathyal deposits of the route d’Angles and Barret-le-Bas sections in Hautes-Alpes (Donze in Busnardo et al., 1979). At both sections, it is associated with species of Orthonotacythere Alexander, 1933, Eucytherura, and Tethysia Donze, 1975, the latter being marker of bathyal to sub-bathyal water depth from the Tithonian to the Hauterivian (Donze, 1975). Higher in both sections, Donze in Busnardo et al. (1979) mentioned that species with ocular nodes, questionably attributed to Eucytherura, set the maximum depth at 600 m. In spite of the lack of Tethysia at Curnier, the presence of Eucytherura (Vesticytherura) captinconcha, dominance of Pontocyprididae and lack of otherwise typical platform taxa point to bathyal water depths. Of note, Eucytherura (Vesticytherura) captinconcha was first reported from Valanginian beds and is here for the first time reported from slightly younger Hauterivian deposits, leading to two hypotheses: 1) it was restricted to oligotrophic bathyal areas during the Valanginian and adapted to conditions of cold seepage later during the Hauterivian, 2) it was a member of both background deep-sea and cold-seep faunas through the Valanginian-Hauterivian interval but its distribution is so far only partly known. Future works should aim at clarifying the two hypotheses.

6 Scolecodonts from Curnier

Two scolecodonts have been retrieved from sediment infilling the brachiopods from Curnier, both corresponding to maxillae 1 (M1; Fig. 2E, F). They are strongly elongated, triangular in outline, with a distinct outer spur, a small yet clear fang and few well-separated denticles, corresponding to the emended diagnosis of the Hartmaniellidae Palurites Kozur, 1967 by Szaniawski and Imajima (1996). Both M1 display a gently curved fang bigger than the largest denticles (2nd and 3rd), anterior half of inner margin scarcely denticulate with a small 1st denticle visible only on right M1 (broken on left M1 or specimen tilted), a long 2nd denticle, slightly shorter 3rd one followed by a long and thin flange. The difference in size of the two specimens, identified as Palurites sp., likely indicate that they correspond to different developmental stages (Fig. 2E: length = 1276 µm; width = 379 µm; Fig. 2F: length = 1027 µm; width = 333 µm).

Palurites sp. is morphologically close to Palurites jurassicus Szaniawski and Imajima, 1996 from the Callovian and Oxfordian of Poland (Szaniawski and Imajima, 1996), both differing from other species by their large size. Palurites sp. however differs by its curved outer margin, fang not angulate, spur located more posteriorly, ramal arch concave and presence of a flange behind 3rd denticle. To the best of our knowledge, such morphological features do not correspond to any known Palurites species but specimens are too few to describe this new species. Of note, Courtinat et al. (1991) and Courtinat (1998) reported scolecodonts from Middle Jurassic and Cretaceous deposits from the south-eastern France basin, including Palurites sp. 1 from the Bathonian of Chabrières, none being conspecific with the present material.

Scolecodonts, i.e., fossil elements of polychaete jaw apparatuses, are here reported for the first time from Curnier and more generally from cold-seep deposits from south-eastern France. Among scolecodonts reported from the area (e.g., Courtinat and Howlett, 1990; Courtinat et al., 1990, 1991), members of Dorvilleidae Chamberlin, 1919 and Arabellidae Hartman, 1944 (junior synonym of Oenonidae Kinberg, 1865) have been considered as indicators of dysaerobic palaeoenvironments during the Mesozoic (Courtinat and Howlett, 1990). The two M1 at hand are distinct from all species from the area but it would be premature to consider that Palurites sp. was seep-endemic. It is nonetheless an important addition to the community developed around the Peregrinella shells of Curnier, polychaete worms being among the most abundant and diverse animals at seeps today (e.g., Levin, 2005; Decker et al., 2012; Alalykina, 2022).

Interestingly, traces on bivalves from Late Jurassic to Early Cretaceous seeps from California and Japan (Jenkins et al., 2013) and calcareous tubes on Peregrinella (Peregrinella) multicarinata shells from Early Cretaceous seeps from Crimean Peninsula (Kiel, 2008) have been interpreted as related to parasitic polychaetes. Today, parasitic polychaetes mainly belong to the families Nautiliniellidae Miura and Laubier, 1990 (junior synonym of Calamyzinae Hartmann-Schröder, 1971; e.g., Miura and Hashimoto, 1996; Quiroga and Sellanes, 2009) and Arabellidae (Dean, 1992). The diet and mode of life of extant Hartmaniellidae, which includes Palurites sp., are unclear but their morphology suggests that they are motile and predating on meiofauna (e.g., Jumars et al., 2015). It is thus likely that Palurites sp. illustrates free-living polychaetes around the Curnier seep, predating on microgastropods, microbrachiopods and microbivalves for instance.

It may be interesting to add that Hartmaniellidae are poorly understood polychaete worms that are today restricted to soft sediments from water depths ranging from 40 to 210 m (e.g., Jumars et al., 2015; Zanol et al., 2021). However, Hartmaniellidae are enigmatic as they are today represented by only a few species while they were common in ancient marine ecosystems, with fossils ranging from the Carboniferous to Triassic (e.g., Szaniawski and Imagima, 1996; Paxton, 2009).

7 Conclusions

Lower Cretaceous Peregrinella-lenses are known since the beginning of the 19th century and have only recently been linked with the diffusive seepage of hydrocarbons on the seafloor. The Muséum national d’Histoire naturelle in Paris stores invertebrates (brachiopods, bivalves and gastropods) from an Hauterivian Peregrinella-lens of the historical site of Curnier (Drôme, France), within the Vocontian Basin. Here we described this material encompassing 44 rhynchonellids Peregrinella (Peregrinella) multicarinata, two lucinids Tehamatea vocontiana and one abyssochrysoid Humptulipsia macsotayi, all corresponding to typical seep taxa from that site and area. Four additional badly preserved brachiopods were processed for the first micropalaeontological analysis of Peregrinella-lenses. They yielded a low abundance ostracod community, yet relatively natural based on rarefaction curve, indicative of a bathyal water depth. The community provided the new pontocypridid Nekrocypris (type species: Nekrocypris sepultinconcha Forel sp. nov.) and Eucytherura (Vesticytherura) captinconcha Forel sp. nov. that was previously known from Valanginian deep-sea deposits of the Hautes-Alpes. Pontocyprididae appear as cornerstones of cold seep ostracod communities within the south-eastern France basin from the Late Jurassic, as shown with the previously described Sahune site, to the Hauterivian, here illustrated from Curnier. However, the lower abundance, replacement of Pontocyprella by Nekrocypris and absence of taxa displaying possible pore clusters at Curnier may witness different contexts compared to the Sahune seepage, including the abundance of Peregrinella brachiopods. We also reported the first scolecodonts that confirm the presence of polychaete annelids around the seepage zone. The present analysis reported a variety of palaeoecological interactions and life modes at the Curnier seep, involving predatory attacks on numerous brachiopod specimens likely by decapod crustaceans, Hartmaniellidae polychaete annelids possibly predating on meiofauna and filter-feeding ostracods.

Acknowledgments

We thank Dr. Frank Sénégas (CNRS) for processing of the Peregrinella specimens for microfossils extraction. Thanks are due to Philippe Loubry (MNHN) for brachiopod photos. This work was granted by MNHN funds ATM 2022 MILEX Paléobiodiversité des milieux extrêmes : les suintements froids du Jurassique supérieur du Sud-Est de la France. Prof. Alan Lord (Senckenberg Forschungsinstitut Frankfurt) and an anonymous reviewer are warmly thanked for their careful reading of an earlier version of this manuscript.

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Cite this article as: Forel M-B, Charbonnier S, Gaspard D, Bergue CT. 2025. Ostracods, brachiopods (Peregrinella) and scolecodonts from the Early Cretaceous cold seeps of Curnier, France, BSGF - Earth Sciences Bulletin 196: 17. https://doi.org/10.1051/bsgf/2025015

All Tables

Table 1

Measurements of all specimens of Peregrinella (Peregrinella) multicarinata from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). All are articulated shells except MNHN.F.97893 (ventral valve).

In the text
Table 2

Taxonomic list of ostracod species identified from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.).

In the text

All Figures

thumbnail Fig. 1

A. Palaeogeographical map of western Tethys during the latest Hauterivian (modified from Dercourt et al., 1993) with position of the south-eastern France Basin. B. Palaeogeography of south-eastern France Basin and nearby areas during the Hauterivian (after Arnaud-Vanneau et al., 1982; Ferry, 1984, Ferry, 2017) showing the three main sites yielding Peregrinella-lenses discussed in the text. F1: Menée fault, F2: Trente-Pas-Condorcet fault.
In the text
thumbnail Fig. 2

A–D. Molluscs from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–C. Tehamatea vocontiana Kiel, 2013 (A: MNHN.F.A97899, B, C: MNHN.F.A97898). D. Humptulipsia macsotayi Kiel et al., 2010 (MNHN.F.A97900). E, F. Palurites sp., scolecodonts from sedimentary infill within Peregrinella-shells, in dorsal view (Rey-Jouvin coll.). E. Left maxilla 1, MNHN.F.F73181. F. Right maxilla 1, MNHN.F.F73182. Scale bars: A–D, 1 cm; D, E, 200 µm.
In the text
thumbnail Fig. 3

Peregrinella (Peregrinella) multicarinata (Lamarck, 1819) from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–C. Dorsal, lateral and anterior views of a juvenile specimen with septum visible on the worn surface (MNHN.F.A97897). D–F. Anterior, dorsal and lateral views of a juvenile with coarse costae (MNHN.F.A97877). G. Dorsal view of specimen MNHN.F.A97871. H–J. Dorsal, lateral and anterior views of MNHN.F.A97881. K, L. Anterior and dorsal views of MNHN.F.A97856 with suspicion of foramen. M, N. Dorsal and ventral views of MNHN.F.A97866. O, P. Dorsal and anterior views of MNHN.F.A97857. Q–S. Anterior, dorsal, lateral views of MNHN.F.A33863. Black arrows: septum. Red arrows: predation marks. Scale bars: 1 cm.
In the text
thumbnail Fig. 4

Peregrinella (Peregrinella) multicarinata (Lamarck, 1819) from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A, B. Dorsal and anterior views of MNHN.F.A97862 with coarse costae and septum visible on a worn surface. C–E. Lateral, dorsal and ventral views of the relatively gibbous MNHN.F.A97869 specimen with marks of predation on both valves. F–H. Anterior, dorsal and lateral views of an equibiconvex specimen with relatively thin costae MNHN.F.A97865. I–L. Dorsal, ventral, lateral and anterior views of MNHN.F.A97864 with remaining parts of calcite shell and coarse costae. M–P. Anterior, dorsal, close up on major growth lines near the antero-lateral margin and lateral views of MNHN.F.A97863. Q. Large isolated ventral valve with coarse costae (MNHN.F.A97893). R. Transverse section trough a large specimen revealing the septum and beginning of crura (MNHN.F.A98232). Black arrows: septum. Red arrows: predation marks. Yellow arrows: growth lines. Scale bars: 1 cm.
In the text
thumbnail Fig. 5

Scatter plots of the size of Peregrinella (Peregrinella) multicarinata at Curnier (Rey-Jouvin coll.) showing the width (top) and thickness (bottom) as a function of length of the shells. Drawings from d’Orbigny (1847).
In the text
thumbnail Fig. 7

Ostracods from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A–E. Nekrocypris sepultinconcha Forel gen. nov. sp. nov. A, B. Articulated carapace broken posteriorly (MNHN.F.F73164) in right view (A) and left view (B). C–E. Articulated carapace broken posteriorly (MNHN.F.F73165) in right view (C), in left view seen in green UV light with white arrows showing the limits of the anterior calcified inner lamella and dashed square showing the adductor muscle scars (D), enlarged in (E). F. Pontocyprella sp. 1, right view of an articulated carapace (MNHN.F.F73166). G. Pontocyprella sp. 2, right view of an articulated carapace (MNHN.F.F73167). H. ?Pseudomacrocypris sp., right view of an articulated carapace (MNHN.F.F73168). I–S. Eucytherura (Vesticytherura) caspinconcha Forel sp. nov. I. Articulated carapace MNHN.F.F73169, right view. J. Paratype, articulated carapace MNHN.F.F73170, right view. K. Articulated carapace (MNHN.F.F73171) in dorsal view, with anterior to the left. L. Articulated juvenile carapace broken anteriorly (MNHN.F.F73172) in right view. M, N. Articulated carapace broken posteriorly (MNHN.F.F73173) in left view (M), with close-up on surface ornamentation and pore conuli P2, P4, P5 (N). O. Holotype, articulated carapace (MNHN.F.F73174) in left view. P. Articulated carapace broken posteriorly (MNHN.F.F73175) in right view. Q. Articulated carapace (MNHN.F.F73176) in left view. R. Articulated carapace (MNHN.F.F73177) in right view. S. Articulated carapace (MNHN.F.F73178) in right view. T. ?Vocontiana sp., right view of an articulated carapace (MNHN.F.F73179). U. Polycope sp., right view of an articulated carapace (MNHN.F.F73180). Scale bars: 100 µm except E: 50 µm, N: 20 µm.
In the text
thumbnail Fig. 6

Ostracods from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). A. ?Macrosarisa sp., articulated carapace, right view (MNHN.F.F73156). B. Paracypris sp., articulated carapace, right view (MNHN.F.F73157). C–U. Nekrocypris sepultinconcha Forel gen. nov. sp. nov. C–G. Articulated carapace (MNHN.F.F73158) in right view (C), detail of the posterior end (D), left view (E), ventral view with anterior to the left (F), dorsal view with anterior to the right (G). H–K. Holotype, articulated carapace (MNHN.F.F73159) in right view (H), left view (I), ventral view with anterior to the right (J), dorsal view with anterior to the right (K). L–P. Paratype, articulated carapace (MNHN.F.F73160) in right view (L), left view (M), ventral view with anterior to the left (N), detail of the posterior end (O), dorsal view with anterior to the left (P). Q, R. Articulated carapace broken posteriorly (MNHN.F.F73161) in right view (Q) and ventral view with anterior to the right (R). S, T. Articulated carapace (MNHN.F.F73162) in right view (S) and left view (T). U. Articulated carapace, right view (MNHN.F.F73163). Scale bars: 100 µm except D: 20 µm, O: 50 µm.
In the text
thumbnail Fig. 8

A. H/L scatter plot of Nekrocypris sepultinconcha Forel gen. nov. sp. nov. and line drawing in right view (top) and ventral view (bottom). B. H/L scatter plot of Eucytherura (Vesticytherura) caspinconcha Forel sp. nov. and line drawings of left views of holotype (top, Fig. 7O) and juvenile (bottom, Fig. 7L), including subdorsal nodes (N1–N3), ridges (R1–R4) and pore conuli (P1–P5). Scale bars: 100 µm.
In the text
thumbnail Fig. 9

A. Individual rarefaction curve for the ostracod assemblage from sedimentary infill within Peregrinella-shells from Curnier, Drôme, south-eastern France Basin, Hauterivian, Early Cretaceous (Rey-Jouvin coll.). B. Circular diagrams summarizing the composition of the ostracod assemblage by number of specimens per species (left) and family (right).
In the text

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