Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia

RESEARCH ARTICLE Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia Eduardo Barro´ nID1*, Daniel Peyrot2, Carlos A. Bueno-Cebollada1☯, Jiřı´ Kvaček3, Sergio A´ lvarez-ParraID4,5, Yul Altolaguirre6, Nieves Mele´ ndez7☯ 1 Museo Geominero, Centro Nacional Instituto Geolo´ gico y Minero de España CN-IGME CSIC, Madrid, Spain, 2 School of Earth and Environment, Centre for Energy Geoscience, University of Western Australia, Crawley, Western Australia, Australia, 3 National Museum Prague, Praha, Czechia, 4 Departament de Dinàmica de la Terra i de l’Oceà, Facultat de Ciències de la Terra, Universitat de Barcelona, Barcelona, Spain, 5 Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain, 6 OCEEH Research Center ‘The role of cultura in early expansions of humans’, Heidelberg Academy of Sciences, Senckenberg Research Institute, Frankfurt am Main, Germany, 7 Deparment of Geodina´ mica, Estratigrafı´a y Paleontologı´a, Facultad de Ciencias Geolo´gicas, Universidad Complutense de Madrid, Madrid, Spain ☯ These authors contributed equally to this work. * e.barron@igme.es Abstract Deserts are stressful environments where the living beings must acquire different strategies to survive due to the water stress conditions. From the late Albian to the early Cenomanian, the northern and eastern parts of Iberia were the location of the desert system represented by deposits assigned to the Utrillas Group, which bear abundant amber with numerous

bioinclusions, including diverse arthropods and vertebrate remains. In the Maestrazgo Basin (E Spain), the late Albian to early Cenomanian sedimentary succession represents the most distal part of the desert system (fore-erg) that was characterised by an alternation of aeolian and shallow marine sedimentary environments in the proximity of the Western Tethys palaeo-coast, with rare to frequent dinoflagellate cysts. The terrestrial ecosystems from this area were biodiverse, and comprised plant communities whose fossils are associ- ated with sedimentological indicators of aridity. The palynoflora dominated by wind-trans- ported conifer pollen is interpreted to reflect various types of xerophytic woodlands from the hinterlands and the coastal settings. Therefore, fern and angiosperm communities abun- dantly grew in wet interdunes and coastal wetlands (temporary to semi-permanent freshwa- ter/salt marshes and water bodies). In addition, the occurrence of low-diversity megafloral assemblages reflects the existence of coastal salt-influenced settings. The palaeobotanical study carried out in this paper which is an integrative work on palynology and palaeobotany, does not only allow the reconstruction of the vegetation that developed in the mid-Creta- ceous fore-erg from the eastern Iberia, in addition, provides new biostratigraphic and palaeogeographic data considering the context of angiosperm radiation as well as the biota inferred in the amber-bearing outcrops of San Just, Arroyo de la Pascueta and La Hoya

(within Cortes de Arenoso succesion). Importantly, the studied assemblages include Afro- pollis, Dichastopollenites, Cretacaeiporites together with pollen produced by Ephedraceae PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.0282178 March 2, 2023 1 / 51 a1111111111 a1111111111 a1111111111 a1111111111 a1111111111 OPEN ACCESS Citation: Barro´n E, Peyrot D, Bueno-Cebollada CA, Kvaček J, A´lvarez-Parra S, Altolaguirre Y, et al. (2023) Biodiversity of ecosystems in an arid setting: The late Albian plant communities and associated biota from eastern Iberia. PLoS ONE 18(3): e0282178. https://doi.org/10.1371/journal. pone.0282178 Editor: Enrique Peñalver, Geological and Mining Institute of Spain, SPAIN Received: July 29, 2022 Accepted: February 8, 2023 Published: March 2, 2023 Peer Review History: PLOS recognizes the benefits of transparency in the peer review process; therefore, we enable the publication of all of the content of peer review and author responses alongside final, published articles. The editorial history of this article is available here: https://doi.org/10.1371/journal.pone.0282178 Copyright: © 2023 Barro´n et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper and its Supporting Information files.