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Hidden Termite Coprolites Revealed by Synchrotron Microtomography Inside Eocene–Oligocene Filled Wood-Borings from the Malzieu

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Hidden Termite Coprolites Revealed by Synchrotron Microtomography Inside Eocene–Oligocene Filled Wood-Borings from the Malzieu Hidden termite coprolites revealed by Synchrotron microtomography inside Eocene–Oligocene filled wood-borings from the Malzieu Basin, Lozère, southern France Jean-David Moreau, Vincent Trincal, Andre Nel, Regine Simon-Coinçon, Vincent Sallé, Matthieu Le Couls, Didier Néraudeau, Vincent Fernandez To cite this version: Jean-David Moreau, Vincent Trincal, Andre Nel, Regine Simon-Coinçon, Vincent Sallé, et al.. Hidden termite coprolites revealed by Synchrotron microtomography inside Eocene–Oligocene filled wood- borings from the Malzieu Basin, Lozère, southern France. Lethaia, Wiley, 2020, 53 (1), pp.106-117. 10.1111/let.12344. hal-02280933 HAL Id: hal-02280933 https://hal.archives-ouvertes.fr/hal-02280933 Submitted on 7 Sep 2019 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Hidden termite coprolites revealed by Synchrotron microtomography inside Eocene–Oligocene filled wood‐ borings from the Malzieu Basin, Lozère, southern France JEAN‐DAVID MOREAU , VINCENT TRINCAL, ANDRE NEL, RÉGINE SIMON‐COINÇON, VINCENT SALLÉ, MATTHIEU LE COULS, DIDIER NÉRAUDEAU AND VINCENT FERNANDEZ Moreau, J.‐D., Trincal, V., Nel, A., Simon‐Coinçon, R., Sallé, V., Le Couls, M., Néraudeau, D. & Fernandez, V.2019: Hidden termite coprolites revealed by Synchrotron microtomography inside Eocene–Oligocene filled wood‐borings from the Malzieu Basin, Lozère, southern France Lethaia, https://doi.org/10.1111/let.12344. Fragments of filled wood‐borings were recently discovered from the Eocene–Oligocene Conglomerate and Sandstone Formation of the Malzieu Basin (Lozère, southern France). Propagation phase‐contrast X‐ray synchrotron microtomography (PPC‐ SRμCT) was used to characterize their hidden inner structures. Virtual 2D sections and 3D reconstructions show that the inner microstructures of borings display a mud- stone–wackestone siliceous matrix including tiny coprolites. The hexagonal cross‐sec- tion of these coprolites is characteristic of termite faecal pellets. Morphology and dimensions of the coprolites share strong similarities with those of extant Kalotermiti- dae (drywood termites). The depositional environment is interpreted as a fluvial valley in which the lenticular sandstone and conglomerate lenses correspond to channels. This discovery complements the record of Palaeogene termites which remain poor in France and throughout the world. □ Faecal pellets, Insecta, Kalotermitidae, Lozère, Palaeogene, PPC‐SRμCT. Jean‐David Moreau [[email protected]], CNRS UMR 6538 Laboratoire Géosciences Océan, Centre de Recherche Yves Coppens, Université Bretagne Sud, Cam- pus de Tohannic, rue André Lwoff, Vannes 56017, France; Vincent Trincal [vincenttrin- [email protected]], LMDC ‐ Laboratoire Matériaux & Durabilité des Constructions, Université de Toulouse, INSA/UPS ‐ 135 av. de Rangueil Toulouse31077, France; Andre Nel [[email protected]], Institut de Systématique, Évolution, Biodiversité, ISYEB‐UMR 7205‐CNRS, MNHN, UPMC, EPHE, Muséum national d'Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, CP 50, Entomologie ParisF‐75005, France; Régine Simon‐ Coinçon [[email protected]], 10, allée du Parchemin Buxerolles86180, France; Vincent Sallé [[email protected]], Association Paléontologique des Hauts Plateaux du Languedoc, 14 chemin des Ecureuils Mende48000, France; Matthieu Le Couls [mat- [email protected]], Didier Néraudeau [[email protected]], CNRS UMR 6118 Géosciences, Université Rennes 1, Campus de Beaulieu, 263 avenue du général Leclerc Rennes Cedex35042, France; Vincent Fernandez [vinfernand@gmail. com], Imaging and Analysis Centre, Natural History Museum, Cromwell Road LondonSW7 5BD, UK; Vincent Fernandez [[email protected]], European Syn- chrotron Radiation Facility, 7 avenue des Martyrs Grenoble38043, France; manuscript received on 17/01/2019; manuscript accepted on 5/04/2019. Although the diversity of extant termites (Isoptera) is evidences such as underground nests (Genise 2016 relatively modest compared with that of other insect and references therein; Roberts et al. 2016), borings major clades, in recent ecosystems they are probably in wood (Rohr et al. 1986; Genise 1995; Francis & the most ecologically important group of insects Harland 2006) and coprolites in wood, sediment and besides the bees and ants (Grimaldi & Engel 2005). amber (Colin et al. 2011). Although termite bodies They constitute recyclers of organic matter essential date back to the Jurassic/Cretaceous boundary (Engel for carbon mineralization, humification and building et al. 2011; Vršanský & Aristov 2014), the record of of soils (Eggleton 2011). Worldwide, evidence of ter- Palaeogene termites remains sparse, except in Baltic mites in fossil ecosystems was demonstrated by the amber. This is particularly true for the Eocene–Oli- discovery of bodies preserved as inclusions in amber gocene interval which is weakly documented in Eur- (Krishna & Emerson 1983; Krishna & Grimaldi 1991; ope (Nel & Paicheler 1993; Engel et al. 2008). Poinar 1993; Nel & Bourguet 2006; Krishna et al. Recently, fragments of filled borings (inner casts) 2013; Coty et al. 2014) and compressions on slabs of containing coprolites were discovered in the Eocene– sedimentary rocks (e.g. Jarzembowski 1981; Nel & Oligocene deposits from the Malzieu Basin, in Lozère Paicheler 1993). Fossil record also yielded indirect (southern France). Although coprolites of termites DOI 10.1111/let.12344 © 2019 Lethaia Foundation. Published by John Wiley & Sons Ltd 2 Moreau et al. LETHAIA 10.1111/let.12344 are known since the Early Cretaceous to the Holo- movements, the CSF is also visible 2 km further cene (Rogers 1938; Corbett & Burrell 2001; Suther- north, at the top of the Verdezun hill as well as 8 km land 2003; Nuorteva & Kinnunen 2008; Pires & to the southwest, near La Chaumette (St‐Chély‐ Sommer 2009; Colin et al. 2011; Boucher 2012; Jud et d'Apcher; Colin 1963; Couturié & Geffroy 1974). The al. 2017), this report consists of the first evidence of CSF yields abundant plant macroremains, including Eocene–Oligocene termite borings containing copro- large trunks preserved as impressions and silicified lites in Europe. The fragments of pellet‐filled borings permineralizations. Their identification and dating were analysed using a non‐destructive imaging tech- were reported in an ancient study (Boule 1888). nique useful for the observation of inner and hidden Based on scanty and fragmented specimens, and by structures: propagation phase‐contrast X‐ray syn- comparison with the regional Cenozoic flora, the chrotron microtomography (PPC‐SRμCT). The author announced an Aquitanian age (Lower Mio- paper aims to compare specimens from Malzieu with cene). However, the comparisons were limited and faecal pellets of extinct and extant termites and dis- Boule (1888) indicated that this age is only an cuss probable authors of the coprolites as well as approximation, even suggesting that the deposit may taphonomical and palaeoecological implications. be more than 10 million years older and dated to the Fragments of borings were discovered in the Con- Priabonian (Upper Eocene). Simon‐Coinçon et al. glomerate and Sandstone Formation whose age is (1981) indicated that the plant species identified by uncertain and debated since the 19th century (Boule Boule (1888) are not characteristic of a particular 1888). stage and may be dated from the Oligocene to the end of the Miocene. Geological setting Age of the Conglomerate and Sandstone The Malzieu basin is located in the south of Massif Formation Central, in the Lozère department, between Mende The age of the CSF has historically been debated and Clermont‐Ferrand (Fig. 1). Deposits of the basin because of the lack of biostratigraphical evidence and lie on the ‘Granite de la Margeride’ basement, a Her- relationships with the other formations of the Mal- cynian porphyritic batholith dated to the Carbonifer- zieu basin (Coinçon et al. 1976). Furthermore, in the ous (Talbot et al. 2005). Numerous studies current topography, the CSF is located more than (Glangeaud 1922; Rey 1949; Bout 1961; Coinçon 100 m above the sediments that have succeeded 1972) tentatively divided the sedimentary series of them, suggesting a tectonic shift (Boule 1888; Bout the Malzieu basin into the seven following forma- 1961; Coinçon et al. 1976). The detachment border tions (Fig. 2), from the oldest to the most recent: the faults pushed blocks, creating thresholds and col- Red Sandstone Formation (RSF), the Conglomerate lapses (more than 600 m according to the unpub- and Sandstone Formation (CSF), the Red Clay For- lished CFMU drillings). Thus, it is possible to find at mation (RCF), the Green Clay Formation (GCF), the the same topographic level formations of different Silcrete Formation (SF), the Variegated Sandy Clays ages. This has misled many authors who thought that Formation (VSCF) and the Chert‐bearing Sand For- the CSF may be at the top of the sedimentary series, mation (Fy) (or Grey Sand Formation). and therefore posterior to the VSCF (Glangeaud 1922; Colin 1963; Couturié 1972; Coque‐Delhuille & The Conglomerate
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