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Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright Author's personal copy Cretaceous Research 29 (2008) 925–929 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes A new amber deposit from the Cretaceous (uppermost Albian-lowermost Cenomanian) of southwestern France Didier Néraudeau a, *, Vincent Perrichot a, Jean-Paul Colin a, Vincent Girard a, Bernard Gomez a, François Guillocheau a, Edwige Masure b, Daniel Peyrot a, Florent Tostain a, Blaise Videt a, Romain Vullo a a b UMR CNRS 6118, Université Rennes 1, Campus de Beaulieu, 263 avenue du Général Leclerc, 35042 Rennes, France UMR CNRS 5143, Université P. & M. Curie, case 104, Tour 46-56, 5 ème étage, pièce 22D, 4, place Jussieu, 75252 Paris cedex 05, France a r t i c l e i n f o a b s t r a c t Article history: Received 14 August 2006 Accepted in revised form 4 May 2008 Available online 13 June 2008 A Cretaceous amber deposit has recently been discovered in a quarry of Charente-Maritime (southwestern France), at Cadeuil. This paper presents the sedimentary and palaeoenvironmental settings of the uppermost Albian-lowermost Cenomanian series including the amber deposit. A preliminary analysis of the amber samples reveals diverse fossil arthropods (a few mites and at least 20 insect families within 9 orders), as well as numerous micro-organisms, mainly algae and mycelia. A myceloid colony of bacteria, a flagellate algae and four especially well preserved insects are illustrated (Diptera Dolichopodidae, Diptera Chironomidae, Hymenoptera Parasitica, and Heteroptera Tingidae). The abundance of the limnic micro-organisms is discussed in terms of bloom events. Their relative scarcity in almost all the amber pieces containing fossil arthropods is attributed to differences in the origin of resin: production along trunk and branches for amber with arthropods; production by aquatic roots for amber rich in algae. The absence of pollen and spores in amber is attributed to differences in the respective periods of resin and palynomorph production, which may be related to a seasonal climate during the Albian-Cenomanian transition in Western Europe. Ó 2008 Elsevier Ltd. All rights reserved. Keywords: Insects Microorganisms Amber Uppermost Albian-lower Cenomanian Charente-Maritime SW France 1. Introduction 2. Geological setting During the last eight years, several amber localities were discovered in the department of Charente-Maritime, in southwestern France. These deposits appear to be amongst the most fossiliferous amber deposits globally known from the Cretaceous with those from Canada, Lebanon, Myanmar, New Jersey, Siberia, and Spain (Perrichot, 2005). The first fossiliferous amber deposit was discovered in 1999 in a sand quarry at Archingeay-Les Nouillers, and is dated as latest Albian (Néraudeau et al., 2002). In 2000, a second deposit from the mid lower Cenomanian was found on the tidal flat at Fouras (Néraudeau et al., 2003). The amber deposit described in the present paper was discovered at Cadeuil in 2001 in a new sand quarry, and provides additional data for analysing the geological and palaeoecological frameworks of the Cretaceous amber deposits from France. The Cadeuil area is located at about ten kilometres from the Seudre estuary (Fig. 1), in the western part of the Saintes synclinal (Néraudeau & Moreau, 1989). In the quarry under study, the uppermost Albian to lower Cenomanian deposits occur directly on top of the Upper Jurassic substratum (Kimmeridgian to Tithonian), and represent transgressive deposits which have eroded the underlying strata. These deposits are mainly composed of fluviatile and paralic sand, but contain several clayey intercalations with local concentrations of fossil plant cuticles (Gomez et al., 2004). At the regional scale, Arnaud (1877), Moreau (1976, 1993a) and Néraudeau et al. (1997, 2002, 2005) considered the alternating sand and clay beds as belonging to the lithological unit A, which they subdivided into two subunits: A1, sand of various grain sizes, arranged in large cross beddings, bearing abundant lignite and amber accumulations, and latestmost Albian in age according to the palynology (Néraudeau et al., 2002); A2, fine sand, mainly arranged in horizontal beds, bearing rare wood remains, and earlymost Cenomanian in age (Moreau, 1993a,b). The unit A is overlain by a shelly sand (‘‘falun’’ sensu Vullo et al., 2003) rich in orbitolines and oysters which marks the base of the lithological unit B (subunit B1). This unit includes the oldest Cenomanian rudist-bearing facies * Corresponding author. E-mail address: [email protected] (D. Néraudeau). 0195-6671/$ – see front matter Ó 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2008.05.009 Author's personal copy 926 D. Néraudeau et al. / Cretaceous Research 29 (2008) 925–929 Fig. 1. Geographical and geological settings of the uppermost Albian to lower Cenomanian amber deposits from Charente-Maritime (southwestern France): Archingeay/LesNouillers (upper Albian); Fouras (mid lower Cenomanian); Cadeuil (uppermost Albian to lowermost Cenomanian). in this region (Néraudeau et al., 1997). In the Cadeuil area, the main amber deposit is located in the top of the subunit A1. 3. Lithostratigraphy and palaeoenvironments The basal sediments of the geological section of Cadeuil consist of 2 m of coarse sand (A1), which is of brackish to coastal marine origin. The sand includes three remarkable fossiliferous horizons, which follow on top of 0.5 m of sand barren in fossils (Fig. 2). The first horizon (A1a) consists of a few clayey lenses bearing plant cuticle compressions (sensu Schopf, 1975 and Martin-Closas and Gomez, 2004); the second one (A1b) provides a large amount of lignite, i.e. an accumulation of fossil wood, together with numerous pieces of amber; the third horizon (A1c) consists in a hard and coarse sandstone with metric cross beddings, which correspond to a succession of channels. This horizon locally includes lignite and amber, sometimes with marine mollusk accumulations. A few selachian (e.g., ‘‘Carcharias’’ (Cappetta and Case, 1975)) and pycnodontiform teeth and echinoid fragments (undetermined spatangoids and cidaroids) are locally associated with the mollusks. The section continues with 10 to 15 m of estuarine sand and clay alternations, which are arranged in horizontal beds and which constitute the subunit A2. The lower part of A2 mainly consists of white fine sand devoid of fossils, while the middle and upper parts include clay beds, and display a few layers bearing fossil plant cuticles. In this quarry, the top of the coarse fossiliferous sandstones is considered as the limit between subunits A1 and A2. The sandstones from the level A1c are dated as early Cenomanian by ostracods, such as the species Neocythere inornata Colin, 1974 or the genera Curfsina, Mauritsina, Dordoniella, and Spinoleberis (Colin, 1974; Damotte et al., 1981). However, the lignite and clay from levels A1a and A1b are not clearly dated because they are devoid of any Cenomanian markers but contain dynocysts which apppeared in the late Albian and survived in the Cenomanian, such as Palaeohystrichophora infusorioides Deflandre, 1935 and Oligosphaeridia poculum Jain, 1977. According to the sequence stratigraphy analysis of all the Albian-Cenomanian sections from the same region (Videt, 2004) the subunit A1, as observed in Cadeuil, can be correlated with the A1 partly dated by palynology as latest Albian at Archingeay-Les Nouillers (level A1sl2 in Néraudeau et al., 2002). Finally, the amber from Cadeuil can be dated as latest Albian-earliest Cenomanian. The top of the Cadeuil section corresponds to the lower part of the subunit B1 (« faluns » sensu Vullo et al., 2003). It begins by 0.5 m of carbonate-rich sand (B1a), especially rich in the large benthic foraminifer Orbitolina concava (Lamarck, 1816), in small oysters (Rhynchostreon, Ceratostreon, Rastellum), in echinoids (Goniopygus menardi (Desmarest, 1825), Tetragramma variolare (Brongniart, 1822), Nucleopygus aff. similis (d’Orbigny, 1856)), and in selachian teeth (‘‘Carcharias’’ amonensis, Tribodus morlati (Landemaine, 1991), Hybodus sp.). Finally, this sand is overlain by limestones bearing large rudists (Ichthyosarcolithes triangularis Desmarest, 1817), crustaceans (Protocallianassa), oysters (Rhynchostreon, Ceratostreon), and echinoids (Periaster undulatus (Agassiz, 1847)). 4. Amber, inclusions, and plant remains 4.1. Plant assemblage The wood assemblage preserved in the uppermost Albianlowermost Cenomanian lignite of Cadeuil quarry is entirely composed of conifers, mainly of the very common Mesozoic morphogenus Agathoxylon (Araucariaceae?), and this is valid both for the basal lignite accumulation (A1b) as well as for the overlying sandstone (A1c). In addition, A1c also bears a few wood fragments identified as Podocarpoxylon (Podocarpaceae), Brachyoxylon, and Protopodocarpoxylon (Cheirolepidiaceae?) (Perrichot, 2005). The palynological analysis of A1a by Peyrot et al. (2005) revealed lycophytes (Retitriletes), ferns (e.g. Cicatricocisporites, Deltoidospora, Reticulosporis), conifers (e.g. Araucariacites, Classopollis, Inaperturopollenites), cycads/gingkoes (Monosulcites/Cycadopites), and angiosperms (Afropollis, Clavatipollenites, Tricolpites). Though a few cuticles of the conifer Glenrosa have been identified, the main component of the fossil plant cuticles from A1a is by far the cheirolepidiaceous conifer Frenelopsis. The recurrent association of a large number of these conifer leaves together with the pollen grains Classopollis provides a challenge to the question of the identity of the main amber-producing trees for the AlbianCenomanian deposits of the Charentes (Cheirolepidiaceae or/and Araucariaceae?). Author's personal copy D. Néraudeau et al. / Cretaceous Research 29 (2008) 925–929 927 to honey-coloured and brown pieces (frequent, up to 80 mm). According to the analysis of the wood fragments embedded in amber, Agathoxylon would be the main plant source for the amber from Cadeuil, as is suggested for other amber deposits from Charente-Maritime by Perrichot (2005). Until now, we have identified about fifty fossil arthropods, mainly insects (Table 1), and numerous micro-organisms preserved in the amber samples. Hymenoptera is the largest group in terms of numbers of specimens as well as in terms of diversity, with at least five families recognized: Braconidae, Megalyridae (Fig. 3A), Platygastridae, Scolebythidae, Trigonalidae, and an undetermined family in Chalcidoidea. Diptera is the second largest group in terms of numbers of specimens, including the families Ceratopogonidae, Chironomidae (Fig. 3B), Dolichopodidae (Fig. 3C), and Psychodidae (Azar et al., 2007). Other arthropods include a few Acari, Blattodea, Homoptera, Lepidoptera, Psocoptera, Thysanoptera, and Heteroptera with the oldest known bug of the family Tingidae (Fig. 3D). Amongst the micro-organisms, colonies of bacteria (Fig. 3E), green algae of the genus Enallax (Girard, in press) (Fig. 3F), cyanophyceae, mycelia, and undetermined germinating fungal spores have been found. As in the other uppermost Albian-lower Cenomanian amber deposits from the Charente-Maritime region (Archingeay-Les Nouillers and Fouras, Fig. 1), almost all the amber samples from Cadeuil which contain algae are lacking arthropod inclusions, while the amber pieces which contain arthropods are lacking palynomorphs. 5. Discussion In previous sedimentological and palaeontological studies on the Charente-Maritime amber deposits, an estuarine environment partially influenced by marine incursions was assumed for the latest Albian epoch at Archingeay-Les Nouillers (Néraudeau et al., 2002), and a more strongly marine-influenced environment was assumed for the earliest Cenomanian period at Fouras (Néraudeau et al., 2003). This is highlighted by the invertebrate fauna, which at Archingeay-Les Nouillers is restricted to a few brackish to coastal oysters of one single species (Striostrea sp., Videt and Platel, 2005), whereas at Fouras oysters are more abundant and diversified Table 1 Arthropod inclusions in amber from Cadeuil Order Taxon N Acari Auchenorrhyncha Blattaria Coleoptera Heteroptera Lepidoptera Psocoptera Thysanoptera Diptera indet. Fulgoroidea indet. indet. Tingidae indet. indet. indet. Ceratopogonidae Chironomidae Dolichopodidae Psychodidae Nematocera indet. Brachycera indet. Braconidae Megalyridae Platygastridae Scolebythidae Trigonalidae Chalcidoidea indet. TOTAL, inclusions 2 1 2 1 1a 1 2 1 4 4 3b 1c 2 4 2 1 4 1 2d 2 5 46 Hymenoptera Fig. 2. Stratigraphical section of the Cadeuil quarry. 4.2. Amber, arthropods and micro-organisms a b The preserved amber shows a wide variety of colours and sizes, ranging from yellow translucent pieces (rare, smaller than 30 mm) c d Ambarcader eugenie Perrichot et al., 2006. Microphorites deploegi Nel et al., 2004. Sycorax neli Azar et al., 2007. Guyotemaimetsha enigmatica Perrichot et al., 2004. Author's personal copy 928 D. Néraudeau et al. / Cretaceous Research 29 (2008) 925–929 Fig. 3. Various fossil inclusions in Albian-Cenomanian amber from Cadeuil (Charente-Maritime, SW France). A, Hymenoptera Megalyridae; B, Diptera Chironomidae; C, Diptera Dolichopodidae, Microphorinae: Microphorites deploegi Nel et al., 2004; D, Heteroptera Tingidae Ambarcader eugenei Perrichot et al., 2006; E, Myceloid colony of bacteria; F, Green algae. (Acutostrea lingularis (Lamarck, 1819), Gyrostrea delettrei (Coquand, 1865), Rhynchostreon suborbiculatum (Lamarck, 1801)) (Videt and Platel, 2005), and are associated with marine mollusks (Arca sp., Pterodonta, sp., Turbo sp.), selachians and ophidians (Néraudeau et al., 2003; Vullo et al., 2005). In comparison, the amber deposit from Cadeuil represents a depositional environment which is more strongly influenced by marine conditions. Indeed, the sandstones containing lignite and amber (A1c) bear locally marine invertebrates, including numerous bivalves (Arca sp., Neitheia sp.) and gastropods (Emarginula sp., Fusus sp., Turbo sp., Turritella sp.), and a few test fragments of irregular and regular echinoids (spatangoids and cidaroids), which are stenohaline organisms. This palaeoenvironmental interpretation fits well with the palaeogeographical location of the Cadeuil quarry, which is interpreted to have been more central in the Aquitain basin than those assumed for the Archingeay-Les Nouillers and Fouras localities (see Fig. 1). Finally, this new amber deposit is especially interesting because it contains both species previously identified in the contemporaneous Author's personal copy D. Néraudeau et al. / Cretaceous Research 29 (2008) 925–929 amber of Archingeay-Les Nouillers, as well as unpublished, possibly endemic (?) species. The species common to the two amber deposits, such as the dolichopodid fly Microphorites deploegi described for Archingeay by Nel et al. (2004), confirm their biostratigraphic correlation inferred from palynological data, whereas the newly identified species, such as the megalyrid wasp figured in this paper (Fig. 3A), increase our knowledge of the latest Albian to early Cenomanian entomofauna from the coastal ecosystems of Western Europe. Moreover, the abundance of algae in numerous large, nondrop-shaped pieces of amber in which arthropod inclusions are generally absent, indicates that a large part of the resin probably exuded from submerged conifer roots, and/or was produced during periods of algal blooms. Finally, the absence of pollen and spores of higher plants in fossil resin that has trapped terrestrial arthropods (insects, arachnids) lends further support to the hypothesis that the period of resin secretion did not coincide with the period of pollen production. These different observations could argue in favour of a season influenced climate pattern during the Albian-Cenomanian transition. Acknowledgements The authors are greatly indebted to Eugène Arnaud and Laurent Lacombe who collected some amber arthropods studied herein. They thank Peter Cobbold who revised the english language. This article is a contribution to to the ANR Project AMBRACE (n BLAN071-184190),to the IFB program ‘‘Interactions biodiversité végétale– changements globaux à la transition Crétacé inférieur–supérieur d’Europe occidentale,’’ and to the French-Spanish program Picasso ‘‘Evolution paléobiologique et paléoenvironnementale des gisements d’ambre du Crétacé d’Espagne et de France.’’ Our research was financially supported by UMR 6118 CNRS, and BG by projects BTE2001–0185-C02–01 and B052001–0173 of the Spanish government and project 2001SGR-75 of the Catalan government. References Arnaud, H., 1877. Mémoire sur le terrain crétacé du Sud-Ouest de la France. Mémoire de la Société Géologique de France 10, 1–110. Azar, D., Tahchy, A., Perrichot, V., 2007. The oldest Sycoracinae (Diptera: Psychodidae) from the French Cretaceous amber. Alavesia 1, 5–10. Cappetta, H., Case, G., 1975. Contribution à l’étude des sélaciens du groupe Monmouth (Campanien - Maestrichtian) du New Jersey. Palaeontographica Abteilung A 151, 1–46. Colin, J.P., 1974. Contribution à l’étude des ostracodes du Crétacé supérieur de Dordogne (France). Géobios 7, 19–42. Damotte, R., Babinot, J.F., Colin, J.P., 1981. Les ostracodes du Crétacé Moyen Européen. Cretaceous Research 2, 287–306. Girard, V. Evidence of Scenedesmaceae (Chlorophyta) from 100 million-year-old amber. Geodiversitas, in press. Gomez, B., Daviero-Gomez, V., Perrichot, V., Thévenard, F., Coiffard, C., Philippe, M., Néraudeau, D., 2004. Assemblages floristiques de l’Albien-Cénomanien de Charente-Maritime (SO France). Annales de Paléontologie 90, 147–159. 929 Martin-Closas, C., Gomez, B., 2004. Taphonomie des plantes et interprétations paléoécologiques. Une synthèse. Geobios 37, 65–88. Moreau, P., 1976. Cadre stratigraphique et rythmes sédimentaires du Cénomanien nord-aquitain (région de Rochefort). Bulletin de la Société Géologique de France 18, 747–755. Moreau, P., 1993a. La transgression cénomanienne sur la marge septentrionale du bassin de l’Aquitaine (Charentes), flanc Nord du synclinal de Saintes et Angoumois. Modalités d’une invasion marine, aspects stratigraphiques, sédimentologiques et paléogéographiques. I. Analyse stratigraphique et identification des milieux. Unpublished thesis, Université Poitiers, 505 pp. Moreau, P., 1993b. La transgression cénomanienne sur la marge septentrionale du bassin de l’Aquitaine (Charentes), flanc Nord du synclinal de Saintes et Angoumois. Modalités d’une invasion marine, aspects stratigraphiques, sédimentologiques et paléogéographiques. III. Paléontologie stratigraphique et biochronologique. Paléogéographie et structure, Unpublished thesis, Université Poitiers, 322 pp. Nel, A., Perrichot, V., Daugeron, C., Néreaudeau, D., 2004. A new Microphorites in the Lower Cretaceous amber of the southwest of France (Diptera: Dolichopodidae, « Microphorinae »). Annales de la Société Entomologique de France. Nouvelle Série 40, 23–29. Néraudeau, D., Moreau, P., 1989. Paléoécologie et paléobiogéographie des faunes d’échinides du Cénomanien nord-aquitain (Charente-Maritime, France). Geobios 22, 293–324. Néraudeau, D., Thierry, J., Moreau, P., 1997. Variation in echinoid biodiversity during the Cenomanian-early Turonian transgressive episode in Charentes (France). Bulletin de la Société Géologique de France 168, 51–61. Néraudeau, D., Allain, A., Perrichot, V., Videt, B., De Lapparent De Broin, F., Guillocheau, F., Philippe, M., Rage, J.-C., Vullo, R., 2003. Découverte d’un dépôt paralique à bois fossiles, ambre insectifère et restes d’Iguanodontidae (Dinosauria, Ornithopoda) dans le Cénomanien inférieur de Fouras (CharenteMaritime, Sud-Ouest de la France). Comptes Rendus Palevol 2, 221–230. Néraudeau, D., Perrichot, V., Dejax, J., Masure, E., Nel, A., Philippe, M., Moreau, P., Guillocheau, F., Guyot, T., 2002. Un nouveau gisement à ambre insectifère et à végétaux (Albien terminal probable): Archingeay (Charente-Maritime, France). Geobios 35, 233–240. Néraudeau, D., Vullo, R., Gomez, B., Perrichot, V., Videt, B., 2005. Stratigraphie et paléontologie (plantes, vertébrés) de la série paralique Albien terminalCénomanien basal de Tonnay-Charente (Charente-Maritime, France). Comptes Rendus Palevol 4, 79–94. Perrichot, V., 2005. Environnements paraliques à ambre et à végétaux du Crétacé Nord-Aquitain (Charentes, Sud-Ouest de la France). Mémoires de Géosciences Rennes 118, 310 pp. Perrichot, V., Nel, A., Néraudeau, D., 2004. A new, enigmatic, evaniomorphan wasp in the Albian amber of France (Insecta, Hymenoptera). Journal of Systematic Palaeontology 2, 159–162. Perrichot, V., Nel, A., Guilbert, E., Néraudeau, D., 2006. Fossil Tingoidea (Heteroptera: Cimicomorpha) from French Cretaceous amber, including Tingidae and a new family, Ebboidae. Zootaxa 1203, 57–68. Peyrot, D., Jolly, D., Barron, E., 2005. Apports de données palynologiques à la reconstruction paléoenvironnementale de l’Albo-Cénomanien des Charentes (Sud-Ouest de la France). Comptes Rendus Palevol 4, 151–166. Schopf, J.M., 1975. Modes of fossil preservation. Review of Palaeobotany and Palynology 20, 27–53. Videt, B., 2004. Dynamique des paléoenvironnements à huı̂tres du Crétacé Supérieur nord-aquitain (SO France) et du Mio-Pliocène andalou (SE Espagne): biodiversité, analyse séquentielle, biogéochimie. Mémoires Géosciences Rennes 108, 1–261. Videt, B., Platel, J.-P., 2005. Les ostréidés des faciès lignitifères du Crétacé moyen du Sud-Ouest de la France (Charentes et Sarladais). Comptes Rendus Palevol 4, 167–176. Vullo, R., Néraudeau, D., Videt, B., 2003. Un faciès de type falun dans le Cénomanien basal de Charente-Maritime (France). Annales de Paléontologie 89, 171–189. Vullo, R., Néraudeau, D., Allain, R., Cappetta, H., 2005. Un nouveau gisement à microrestes de vertébrés continentaux et littoraux dans le Cénomanien inférieur de Fouras (Charente-Maritime, Sud-Ouest de la France). Comptes Rendus Palevol 4, 95–107.