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      Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards

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          Abstract

          Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21 st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.

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          The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe

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            Phylogeny And Systematics Of Squamata (Reptilia) Based On Morphology

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              Eocene lower vertebrates from Ellesmere Island, Canadian Arctic Archipelago

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                Author and article information

                Contributors
                Role: ConceptualizationRole: Formal analysisRole: InvestigationRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: InvestigationRole: ResourcesRole: Writing – review & editing
                Role: Data curationRole: Funding acquisitionRole: InvestigationRole: ResourcesRole: SupervisionRole: Writing – review & editing
                Role: ConceptualizationRole: InvestigationRole: Writing – review & editing
                Role: ConceptualizationRole: Formal analysisRole: InvestigationRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: ConceptualizationRole: ResourcesRole: Writing – review & editing
                Role: InvestigationRole: Writing – review & editing
                Role: InvestigationRole: ResourcesRole: Writing – review & editing
                Role: Data curationRole: Funding acquisitionRole: ResourcesRole: Writing – review & editing
                Role: InvestigationRole: ResourcesRole: Writing – review & editing
                Role: InvestigationRole: Writing – review & editing
                Role: ConceptualizationRole: Formal analysisRole: Funding acquisitionRole: InvestigationRole: Project administrationRole: SupervisionRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                5 December 2018
                2018
                : 13
                : 12
                : e0207719
                Affiliations
                [1 ] Dipartimento di Scienze della Terra, Università di Torino, Torino, Italy
                [2 ] Universidad Estatal de la Peninsula de Santa Elena, La Libertad, Santa Elena, Ecuador
                [3 ] Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain
                [4 ] Division of Anthropology, American Museum of Natural History, New York, United States of America
                [5 ] New York Consortium in Evolutionary Primatology, New York, United States of America
                [6 ] School of Earth Sciences, University of Bristol, Bristol, United Kingdom
                [7 ] Laboratory of Geology and Paleontology, Department of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece
                [8 ] Departamento de Paleobiología, Museo Nacional de Ciencias Naturales–Consejo Superior de Investigaciones Cientificas, Madrid, Spain
                [9 ] ARAID–Fundación Conjunto Paleontológico de Teruel-Dinópolis, Teruel, Spain
                [10 ] School of Earth & Environmental Sciences, University of Manchester, Manchester, United Kingdom
                [11 ] Department of Geological Sciences, Faculty of Science, Masaryk University, Brno, Czech Republic
                Ecole Normale Supérieure de Lyon, FRANCE
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Author information
                http://orcid.org/0000-0001-6544-5201
                http://orcid.org/0000-0002-3433-6093
                http://orcid.org/0000-0003-4416-4560
                http://orcid.org/0000-0003-2151-7693
                Article
                PONE-D-18-23407
                10.1371/journal.pone.0207719
                6281198
                30517172
                99b26723-e9f9-4cab-ad51-73ac170e792f
                © 2018 Villa 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.

                History
                : 8 August 2018
                : 5 November 2018
                Page count
                Figures: 18, Tables: 0, Pages: 46
                Funding
                This research has been supported by the Spanish Agencia Estatal de Investigación (Ministerio de Economía, Industria y Competitividad)–European Regional Development Fund of the European Union (CGL2016-76431-P and CGL2017-82654-P, AEI/FEDER EU), by the Generalitat de Catalunya (CERCA Programme) and by the Italian MIUR (PRIN 2009MSSS9L_002). The study of the Batallones material was carried out thanks to the SYNTHESYS Project funded by European Community Research Infrastructure Action under the FP7 ‘Structuring the European Research Area’ Programme (ES-TAF-1975 to MD). Access to the collections of Natural History Museum Wien was also possible thanks to a SYNTHESYS grant (AT-TAF-4591) to AV. AB is a Newton International Fellow (NF160464), funded by the Royal Society. Fieldwork at ACM was funded by CESPA Gestión de Residuos, S.A.U. The Cerro the los Batallones project was funded by the Ministerio de Economía y Competitividad de España (CGL2015-68333-P), the Research Groups CSIC 641538 and CAM-UCM 910607, and the Dirección General de Patrimonio (Comunidad Autónoma de Madrid). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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