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A Review of Amber and Copal Occurrences in Africa and Their Paleontological Significance Valentine Bouju, Vincent Perrichot

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A Review of Amber and Copal Occurrences in Africa and Their Paleontological Significance Valentine Bouju, Vincent Perrichot A review of amber and copal occurrences in Africa and their paleontological significance Valentine Bouju, Vincent Perrichot To cite this version: Valentine Bouju, Vincent Perrichot. A review of amber and copal occurrences in Africa and their paleontological significance. Bulletin de la Société Géologique de France, Société géologique de France, 2020, 191 (1), pp.17. 10.1051/bsgf/2020018. insu-02881938 HAL Id: insu-02881938 https://hal-insu.archives-ouvertes.fr/insu-02881938 Submitted on 26 Jun 2020 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. Distributed under a Creative Commons Attribution| 4.0 International License See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/342466062 A review of amber and copal occurrences in Africa and their paleontological significance Article in Bulletin de la Societe Geologique de France · June 2020 DOI: 10.1051/bsgf/2020018 CITATIONS 0 2 authors: Valentine Bouju Vincent Perrichot Université de Rennes 1 Université de Rennes 1 1 PUBLICATION 0 CITATIONS 184 PUBLICATIONS 3,371 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Taxonomy, phylogeny, and biogeography of myrmicine ants: insight from Cenozoic fossils View project Systematics and taxonomy of Rhopalosomatidae (Hymenoptera) View project All content following this page was uploaded by Vincent Perrichot on 26 June 2020. The user has requested enhancement of the downloaded file. BSGF - Earth Sciences Bulletin 2020, 191, 17 © V. Bouju and V. Perrichot, Published by EDP Sciences 2020 https://doi.org/10.1051/bsgf/2020018 Available online at: Special Issue L’Ambre www.bsgf.fr A review of amber and copal occurrences in Africa and their paleontological significance Valentine Bouju and Vincent Perrichot* Univ. Rennes, CNRS, Géosciences Rennes, UMR 6118, F35000 Rennes, France Received: 9 April 2020 / Accepted: 19 May 2020 Abstract – The paleontological interest for fossil plant resins (amber and copal) has greatly increased in the last decades, as field studies have resulted in the discovery of various new deposits worldwide. Yet, amber- rich deposits remain particularly scarce on continents from former Gondwana. Here we review the known occurrences of copal and amber from Africa, with a state-of-the-art regarding the age dating, the putative plant sources, the fossil content, as well as the paleoenvironmental settings. The first African ambers known to yield arthropods and other organismal inclusions, found recently from the early Cretaceous of Congo and the Miocene of Ethiopia, are briefly overviewed. Keywords: fossil resins / amber / Africa / Cretaceous / Cenozoic Résumé – Une révision des gisements d’ambre et de copal en Afrique et leur signification paléontologique. L’intérêt paléontologique pour les résines végétales fossiles (ambre et copal) s’est fortement accru ces dernières décennies, les études de terrain ayant permis la découverte de divers nouveaux gisements à travers le monde. Les gisements riches en ambre demeurent néanmoins particulièrement rares sur les continents autrefois gondwaniens. Nous révisons ici les occurrences connues de copal et d’ambre en Afrique, avec un bilan concernant la datation, les sources botaniques probables, le contenu fossile, ainsi que le contexte paléoenvironnemental. Les premiers ambres africains livrant des arthropodes et autres inclusions organiques fossiles, découverts récemment dans le Crétacé inférieur du Congo et le Miocène d’Ethiopie, sont brièvement présentés. Mots clés : résines fossiles / ambre / Afrique / Crétacé / Cénozoïque 1 Introduction Crelling, 1995; Langenheim, 2003; Vávra, 2009; Lambert et al., 2012; Penney, 2016; Delclòs et al., 2020). The time Fossilized resins represent important sources of paleonto- needed for full polymerization greatly varies depending on the logical data for reconstructing ancient terrestrial ecosystems. A resin composition and the burial conditions, so that there is no single fossiliferous deposit can provide abundant, various, and temporal feature to characterize copal and amber. Commonly, often exquisitely preserved organic inclusions (mostly however, copals are considered as young, semifossilized resins arthropods, plants, protists, and fungi; more rarely microalgae, ranging from Recent up to the Pliocene age (5.3 Ma) (Poinar, molluscs, and vertebrate remains). Fresh liquid resins solidify 1992; Grimaldi, 1996; Delclòs et al., 2020). But there are older and progressively transform into fossil resins during burial resins that are only partially polymerized and can readily through the chemical process of polymerization, during which dissolve, or that remain sticky and aromatic when cut or their volatile compounds are lost (Anderson and Crelling, polished, and should therefore be copals. Nonetheless, these fi 1995; Clifford and Hatcher, 1995; Seyfullah et al., 2018). are generally classi ed as ambers because of their antiquity (e. “ ” “ ” Amber is a fully polymerized, cross-linked resin that can at g., Eocene Cambay amber from India, or Miocene amber most softened using organic solvents. A resin that is from Indonesia; see Rust et al., 2010; Mazur et al., 2014; incompletely cross-linked and not fully polymerized can Narudeesombat et al., 2014). In the present review of all “ ” “ ” dissolve in various solvents and is sometime called copal, known fossil resins from Africa, copal and amber are used although the use of this term is ambiguous (Anderson and in this temporal sense rather than chemical sense, i.e. regardless of their level of polymerization. The vast majority of amber deposits known today occurs in *Corresponding author: [email protected] the Northern Hemisphere, particularly from the former This 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. V. Bouju and V. Perrichot: BSGF 2020, 191, 17 Fig. 1. Present-day map with known distribution of African deposits of fossil resins. Laurasian continent. The paucity of Gondwanan amber 2 Copal deposits may results from a long lack of prospection for this geological material in the Southern Hemisphere, combined Copal had once a major export value for its industrial use in with typically limited outcrops due to the dense plant covering varnishes and lacquers. A rich literature exists on African copals, in intertropical zones. Amber deposits of Gondwanan origin their botany and their trade that developed during colonialism in comprise Indomalayan occurrences in India (early Eocene; the nineteenthand twentieth centuries.Abundant copal resources Rust et al., 2010) and possibly Myanmar (early Cenomanian; have been exploited mostly from two regions of the continent: at the time probably an island in the Tethys Ocean but Zanzibar and the German East Africa, that became a British originating from Austral Gondwana, see Rasnitsyn and Öhm- territory after WWI, and are now corresponding to the modern Kühnle, 2018, Poinar, 2019 or Westerweel et al., 2019 for Tanzania (Kirk, 1871; Schlüter and von Gnielinski, 1987; arguments); Australasian occurrences in Australia (Triassic to Sunseri, 2007); and the Democratic Republic of the Congo Neogene; Hand et al., 2010; Sonibare et al., 2014; Stilwell (DRC, formerly Congo Free State, then Belgian Congo) et al., 2020) and New Zealand (Cretaceous, Oligocene and (Pynaert, 1924; De Wildeman, 1927; Vervloet, 1932; Hellinckx, Miocene; Schmidt et al., 2018; Mays et al., 2019; Stilwell 1935; Anonymous, 1942; Billing,1944).Copal is moregenerally et al., 2020); South American occurrences in Brazil (early known from several countries in East Africa and West Africa Cretaceous; Pereira et al., 2011), Ecuador (Aptian–Albian; (Fig. 1). All African copals derive from legume trees in the Cadena et al., 2018), Venezuela (Miocene, Pérez et al., 2016), family Fabaceae, mostly Hymenaea or Guibourtia in the and Peru (Miocene; Antoine et al., 2006); and rare African Caesalpinioideae group, but also Colophospermum, Copaifera, occurrences that are detailed and reviewed hereafter along with Daniellia or Tessmannia in the Detarioideae group (Howes, 1949; the known copal deposits. Langenheim, 2003; McCoy et al., 2017; Delclòs et al., 2020). Page 2 of 11 V. Bouju and V. Perrichot: BSGF 2020, 191, 17 2.1 East African copals (Lambert et al., 1995, 2012; McCoy et al., 2017; Delclòs et al., 2020); and more specifically Hymenaea verrucosa, the sole Pleistocene or even younger copal has been reported from species found in East Africa and Madagascar, based on various countries along the African East coast: Kenya, botanical considerations (Kirk, 1871; Gilg, 1898; Zimmerman, Tanzania, Mozambique, Madagascar (Fig. 1). Copal was 1908; Howes, 1949; Langenheim, 1990, 2003; Delclòs et al., traded into Indian Ocean networks, including India, by Swahili 2020). communities as soon as in the
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