palevol comptes rendus 2023  22  18 Directeurs de la publication / Publication directors : Bruno David, Président du Muséum national d’Histoire naturelle Étienne Ghys, Secrétaire perpétuel de l’Académie des sciences Rédacteurs en chef / Editors-in-chief : Michel Laurin (CNRS), Philippe Taquet (Académie des sciences) Assistante de rédaction / Assistant editor : Adenise Lopes (Académie des sciences ; cr-palevol@academie-sciences.fr) Mise en page / Page layout : Audrina Neveu (Muséum national d’Histoire naturelle ; audrina.neveu@mnhn.fr) Révisions linguistiques des textes anglais / English language revisions : Kevin Padian (University of California at Berkeley) Rédacteurs associés / Associate editors (*, took charge of the editorial process of the article/a pris en charge le suivi éditorial de l’article) : Micropaléontologie/Micropalaeontology Maria Rose Petrizzo (Università di Milano, Milano) Paléobotanique/Palaeobotany Cyrille Prestianni (Royal Belgian Institute of Natural Sciences, Brussels) Métazoaires/Metazoa Annalisa Ferretti (Università di Modena e Reggio Emilia, Modena) Paléoichthyologie/Palaeoichthyology Philippe Janvier (Muséum national d’Histoire naturelle, Académie des sciences, Paris) Amniotes du Mésozoïque/Mesozoic amniotes Hans-Dieter Sues (Smithsonian National Museum of Natural History, Washington) Tortues/Turtles Walter Joyce (Universität Freiburg, Switzerland) Lépidosauromorphes/Lepidosauromorphs Hussam Zaher* (Universidade de São Paulo) Oiseaux/Birds Eric Buffetaut (CNRS, École Normale Supérieure, Paris) Paléomammalogie (mammifères de moyenne et grande taille)/Palaeomammalogy (large and mid-sized mammals) Lorenzo Rook (Università degli Studi di Firenze, Firenze) Paléomammalogie (petits mammifères sauf Euarchontoglires)/Palaeomammalogy (small mammals except for Euarchontoglires) Robert Asher (Cambridge University, Cambridge) Paléomammalogie (Euarchontoglires)/Palaeomammalogy (Euarchontoglires) K. Christopher Beard (University of Kansas, Lawrence) Paléoanthropologie/Palaeoanthropology Aurélien Mounier (CNRS/Muséum national d’Histoire naturelle, Paris) Archéologie préhistorique (Paléolithique et Mésolithique)/Prehistoric archaeology (Palaeolithic and Mesolithic) Nicolas Teyssandier (CNRS/Université de Toulouse, Toulouse) Archéologie préhistorique (Néolithique et âge du bronze)/Prehistoric archaeology (Neolithic and Bronze Age) Marc Vander Linden (Bournemouth University, Bournemouth) Référés / Reviewers : https://sciencepress.mnhn.fr/fr/periodiques/comptes-rendus-palevol/referes-du-journal Couverture / Cover : Made from the Figures of the article. Comptes Rendus Palevol est indexé dans / Comptes Rendus Palevol is indexed by: – Cambridge Scientific Abstracts – Current Contents® Physical – Chemical, and Earth Sciences® – ISI Alerting Services® – Geoabstracts, Geobase, Georef, Inspec, Pascal – Science Citation Index®, Science Citation Index Expanded® – Scopus®. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 diff.pub@mnhn.fr / https://sciencepress.mnhn.fr Académie des sciences, Institut de France, 23 quai de Conti, 75006 Paris. © This article is licensed under the Creative Commons Attribution 4.0 International License (https://creativecommons.org/licenses/by/4.0/) ISSN (imprimé / print) : 1631-0683/ ISSN (électronique / electronic) : 1777-571X Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island (Columbretes Islands, Castellón, Spain): an unexpected diverse assemblage Rafael MARQUINA-BLASCO Palaeontology of Cenozoic Vertebrates Research Group (PVC-GIUV), Àrea de Palaeontologia, Departament de Botànica i Geologia, Universitat de València, Dr. Moliner 50, 46100 Burjassot (Spain) and Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3), Universitat Rovira i Virgili, 43007 Tarragona (Spain) and Museu Valencià d’Història Natural, L’Hort de Feliu, P.O. Box 8460, 46230 Alginet (Spain) rafael.marquina@uv.es (corresponding author) Ana FAGOAGA Palaeontology of Cenozoic Vertebrates Research Group (PVC-GIUV), Àrea de Palaeontologia, Departament de Botànica i Geologia, Universitat de València, Dr. Moliner 50, 46100 Burjassot (Spain) and Museu Valencià d’Història Natural, L’Hort de Feliu, P.O. Box 8460, 46230 Alginet (Spain) and Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3), Universitat Rovira i Virgili, 43007 Tarragona (Spain) and Centro de Investigação em Ciências Geo-Espaciais (CICGE), Faculty of Sciences, University of Porto, Vila Nova de Gaia (Portugal) Vicente D. CRESPO Departamento de Ciências da Terra (Geobiotec), Universidade Nova de Lisboa, Faculdade da Ciências E (FCT-UNL), 2829-516 Caparica (Portugal) and Museu da Lourinhã, 9 Rua João Luis de Moura, 2530-158 Lourinhã (Portugal) and Museu Valencià d’Història Natural, L’Hort de Feliu, P.O. Box 8460, 46230 Alginet (Spain) Alberto MARTÍNEZ-ORTÍ Museu Valencià d’Història Natural, L’Hort de Feliu, P.O. Box 8460, 46230 Alginet (Spain) and Departamento de Parasitología, Facultad de Farmacia, Universitat de València, Dr. Moliner 50, 46100 Burjassot (Spain) Salvador BAILON Archéozoologie, Archéobotanique : Sociétés, Pratiques, Environnements (AASPE), Histoire naturelle de l’Homme préhistorique (HNHP), Muséum national d’Histoire naturelle (MNHN) - CNRS, case postale 55, 57 rue Cuvier, F-75231 Paris cedex 05 (France) Antonio SÁNCHEZ-MARCO Institut Català de Paleontologia Miquel Crusafont, Campus de la UAB, 08193 Bellaterra (Spain) Daniel GRACIA-MONFERRER Carlos de SANTISTEBAN Palaeontology of Cenozoic Vertebrates Research Group (PVC-GIUV), Àrea de Palaeontologia, Departament de Botànica i Geologia, Universitat de València, Dr. Moliner 50, 46100 Burjassot (Spain) COMPTES RENDUS PALEVOL • 2023 • 22 (18) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr 409 Marquina-Blasco R. et al. Francisco Javier RUIZ-SÁNCHEZ Palaeontology of Cenozoic Vertebrates Research Group (PVC-GIUV), Àrea de Palaeontologia, Departament de Botànica i Geologia, Universitat de València, Dr. Moliner 50, 46100 Burjassot (Spain) and Museu Valencià d’Història Natural, L’Hort de Feliu, P.O. Box 8460, 46230 Alginet (Spain) Submitted on 22 June 2021 | Accepted on 25 May 2022 | Published on 8 June 2023 urn:lsid:zoobank.org:pub:34CD62EB-2344-4119-B516-2AEA9E9D27F7 Marquina-Blasco R., Fagoaga A., Crespo V. D., Martínez-Ortí A., Bailon S., Sánchez-Marco A., Gracia-Monferrer D., de Santisteban C. & Ruiz-Sánchez F. J. 2023. — Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island (Columbretes Islands, Castellón, Spain): an unexpected diverse assemblage. Comptes Rendus Palevol 22 (18): 409-438. https://doi.org/10.5852/cr-palevol2023v22a18 ABSTRACT The Columbretes Islands (eastern Spain) comprise a volcanic archipelago 50 km off the eastern Span- ish coast. Illa Grossa is the main island of the archipelago. After the settlement of humans during the mid-19th century, strong modifications in biodiversity took place, including the reduction of the non-flying vertebrate fauna to a single taxon, Podarcis liolepis atratus (Boscá, 1916). The study of the fossils yielded at the COLT site (dated by AMC between 2780-2724 cal BP and 2600-2492 cal BP) from a loess deposit located at the south of Illa Grossa Island shows that this fauna was more diverse in the past. The assemblage was dominated by squamate reptiles, belonging to two lizards (Chalcides bedriagai (Boscá, 1880) and cf. Podarcis Wagler, 1830) and one snake (Vipera cf. latastei Boscá, 1878). The record in the late Holocene of Columbretes of Vipera cf. latastei settles the existing controversy about the identity of snakes in the archipelago. Regarding Chalcides bedriagai, this is the first record KEY WORDS of its presence in the Columbretes Islands. Other faunal elements recovered from the site include at Vipers, least six species of undetermined birds (passeriforms and procellariforms) and the anthropophilic Mus conservation palaeobiology, cf. musculus Linnaeus, 1758. The origin of the squamates taxa from Columbretes palaeontological small vertebrates, site (COLT) is still unclear, but was probably related to the Late Pleistocene-Greenlandian (11 700 Mediterranean, palaeoecology, to 8300 BP, Early Holocene) sea transgression, natural passive colonization, and/or human intro- islets. duction. We hypothesize implications for the conservation and management of the Natural Reserve. RÉSUMÉ Squamates, rongeurs et oiseaux des dépôts holocènes de l’île Illa Grossa (îles Columbretes, Castellón, Espagne) : un assemblage diversifié inattendu. Les îles Columbretes (Est de l’Espagne) constituent un archipel volcanique situé à 50 km de la côte est de l’Espagne. Illa Grossa est l’île principale de l’archipel. Après l’installation de la population humaine sur celle-ci au milieu du xixe siècle, des importantes modifications de la biodiversité ont eu lieu, notamment la réduction de la faune vertébrée non volante dont un seul taxon persiste, Podarcis liolepis atratus (Boscá, 1916). L’étude des fossiles trouvés sur le site Columbretes (COLT) (daté par AMC entre 2780-2724 cal BP et 2600-2492 cal BP) dans un dépôt de lœss situé au sud de l’île Illa Grossa, montre que cette faune était plus diversifiée dans le passé. L’assemblage était dominé par des reptiles squamates, appartenant à deux lézards (Chalcides bedriagai (Boscá, 1880) et cf. Podarcis Wagler, 1830) et un serpent (Vipera cf. latastei Boscá, 1878). L’enregistrement à l’Holocène supérieur de Columbretes de Vipera cf. latastei, règle la controverse existante sur l’identité des serpents dans l’archipel. En ce qui concerne Chalcides bedriagai, il s’agit du premier signalement de sa présence dans les îles Columbretes. Les autres éléments de la faune vertébrée retrouvés sur le site comprennent, au MOTS CLÉS moins, six espèces d’oiseaux indéterminés (passériformes et procellariformes) et l’anthropophile Mus Vipères, cf. musculus Linnaeus, 1758. L’origine des taxons de squamates de COLT n’est toujours pas claire, paléobiologie de la conservation, mais elle est probablement liée à une transgression marine du Pléistocène supérieur (Groenlandien petits vertébrés, 11 700 à 8 300 BP, Holocène inférieur), une colonisation passive naturelle et/ou à une introduction Méditerranée, paléoécologie, par l’homme. Nous avons également émis des hypothèses sur les implications pour la conservation îlots. et la gestion de la réserve naturelle. 410 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island INTRODUCTION human presence becomes apparent in the archipelago (Serrano Adell 1991), deeply altering its floral and faunal communi- The Columbretes islands are a small volcanic archipelago close ties. Snakes were especially persecuted during this period, to the continental coast (50 km) in the western Mediterranean through direct hunting of specimens, the introduction of (Fig. 1A). This archipelago constitutes the emerged part of predators and, the burning of vegetation, until their total the Gulf of Valencia Volcanic Province (GVVP), extending extinction on Illa Grossa Island, which affected the general 200 km NE-SW over the continental shelf between Tarragona biodiversity and has been related with the extirpation of the and Valencia, mostly as submarine volcanoes. The GVVP is terrestrial gastropods Theba pisana (Müller, 1774) and Pota- located on the western margin of the Valencia Through. This mias elegans (Müller, 1774) (Robles 1991). The last reference area, between the Pyrenean front and the Alboran sector, was of a living snake on the islands was in 1885 (Mestre et al. affected by a rifting process developed after the Oligocene- 2010). Despite the historical high density of snakes on the Lower Miocene alpine orogenic stage (López-Ruiz et al. Illa Grossa Island, their taxonomic identity has remained 2002). This led to the thinning of the crust that extends up unclear (Jiménez et al. 2002). Although most authors iden- to about 50 km on land in the Eastern sector of the Iberian tified it as a viper (e.g. Castilla et al. 1991; van Damme & Chain (Dañobeitia et al. 1992). This context is at odds with Castilla 1996; Brito 2017), no specimens or remains whose the idea of the Rhine-Rhône Rift spreading southwards. origin is unequivocally from the island have been preserved López-Ruiz et al. (2002) propose for this area a geodynamic (Ruiz-Sánchez et al. 2019). model of the volcanism of the SE of Iberia with two phases. At present, the Columbretes Islands area is protected by A first lower Oligocene-Miocene synorogenic phase linked various figures of the European and Spanish legislation due to the genesis of the Valencia Trough, and a second terminal to their high natural heritage value (Special Protection Area Miocene-Quaternary phase related to the functioning of the and Site of Community Importance of the Natura 2000 trans-Moroccan, Western Mediterranean, European Fault network and Natural Reserve and Marine Reserve, among Zone (TMWMEFZ) that crosses the eastern sector of the other). This archipelago also constitutes an important resting Iberian Peninsula. Although, the origin of the archipelago has area for many birds during their pre- and post-nuptial migra- been dated, at least, between 0.8 and 0.3 My (Ancochea & tion routes between Europe and Africa (Mestre et al. 2010). Huertas 2021), the submarine structures associated to the Humans are one of the main factors determining the cur- volcanic field may be much younger, about 13 000 years old rent composition of biological communities in the Medi- (Muñoz et al. 2005). terranean region, and this is particularly evident in island Traditionally, the archipelago is divided in four groups of faunas (Blondel & Aronson 1999; Corti et al. 1999). The islands, from north to south: Illa Grossa, Ferrera, Foradada study of these biocenosis and the disturbances produced by and El Carallot (Fig. 1B). As Ancochea & Huertas (2020) human colonization has been studied in the main insular and Huertas & Ancochea (2020) pointed out these groups areas in the Mediterranean, such as the Balearic Islands, do not correspond to a particular volcanic phase, and there Corsica, Sicily, Malta, Crete, and Cyprus (Seguí & Alcover are some islets than belong to a different phases (i.e., Ferrera 1999; Alcover 2000; Marra 2005; van der Geer et al. 2010). group). Thus, there are recognised two alignments according However, this kind of studies has been scarce on small islands to two strikes: a main N170° E direction (formed by more (e.g. Cherry & Leppard 2017). than seven volcanic centres of phonolitic composition and In 2005 a palaeontological survey of Illa Grossa was car- which, at present, appears as small islets) and a second direc- ried out to locate vertebrate fossil remains, in order to char- tion, N55° E (which belongs to the two volcanic centres of acterize the biological communities of the island prior to basic composition, one of which is Illa Grossa) (Ancochea & the human settlement, and to assess human impact in this Huertas 2020; Huertas & Ancochea 2020). Among them, the area (Barnosky et al. 2017). As a result, the first palaeonto- Illa Grossa group (Fig. 1C) would correspond to a magmatic logical site of the island, named as Columbretes (COLT), outcrop older (0.7-0.8 My) than the other island groups was located in an extensive aeolian deposit in the southern (0.2-0.3 My) (Ancochea & Huertas 2021). Basanite and part of Illa Grossa Island fossil remains of vertebrates and phonolite rocks mainly constituted the Illa Grossa island gastropods (Fig. 1C). In that area, Hernández-Pacheco & (Aparicio & García 1995). The current ring-shape of the Asensio Amor (1966) reported the presence of numerous Illa Grossa Island is the result of three successive emission bones, although they identified them as belonging to rab- centres (Ancoechea & Huertas 2020). bits, a species introduced by humans several times along the The name of the archipelago (Columbretes, from the latin 19th century AC (Salvator 1895). word “coluber”, which derived in the Spanish word “culebra”, In the present work, the fossil remains of reptiles, birds, and meaning “snake”) comes from the high density of snakes in rodents from the COLT site are studied within a radiocarbon the Illa Grossa Island. In fact, in the Antiquity, the island dated deposit, thus providing new data about the palaeobio- was called Ophiusa by the Ancient Greeks, and Colubraria geographical and palaeoecological context of the Columbretes by the Romans (Bernis 1968). The large number of snakes islands in the time interval between 2 780 and 2 492 BP. For present on Illa Grossa prevented the human settlement on the first time, we have obtained snake remains unmistakably the island until the mid-19th century, when the construction originating from the Columbretes archipelago. The conserva- of the lighthouse began. Starting from this point, in theory, tion implications of these findings are also discussed. COMPTES RENDUS PALEVOL • 2023 • 22 (18) 411 Marquina-Blasco R. et al. –50 A B 1 –25 2 –25 –50 –75 3 –25 N N –25 100 km 500 m 4 C N 100 m Fig. 1. — A, Location of Columbretes Islands in the western Mediterranean; B, location of Illa Grossa Island on the archipelago, Island’s group: 1, Illa Grossa group; 2, Ferrera group; 3, Foradada group; 4, El Carallot group; C, location of the Columbretes palaeontological site (COLT) on Illa Grossa (white star). GEOLOGICAL CONTEXT redeposited pyroclasts and levels of calcareous palaeosoils (Fig. 2). Hernández-Pacheco & Asensio Amor (1966) con- The Illa Grossa Island is a circular structure open to the NE sider that the top of the fine grained materials of this terrace consisting of the emerged rim of a volcano with three SW to have a wind source. The paleontological remains studied in NE overlapping craters (Fig. 1C). It is composed of pyroclastic this paper have been sampled in this terrace, located 30 m agglomerates, lapilli and ash (Vidal Romaní et al. 1972), in high in the southern sector of the interior of the crater that layers inclined both to the outside of the volcanic cone and forms the Puerto Tofiño. The sample has been extracted to to the inside of the crater. Most of the island has a relief with a depth of 95 cm (Fig. 2). slopes and locally vertical cliffs (Las Covachas, Mascarat). Most of the materials are volcanic pyroclasts with the excep- tion of a horizontal terrace between 26 and 36 metres above MATERIAL AND METHODS mean sea level. It is a horizontal sedimentary terrace described by Hernández-Pacheco & Asensio Amor (1966). This sedi- The material studied in the present work consists mainly of mentary terrace is 6 m thick and lies on lapilli and altered disarticulated elements collected during the 2005 palaeontologi- ashes. It consists of brown and yellowish silts, with traces of cal prospection campaign at Illa Grossa (Columbretes Islands, 412 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island Spain). An amount of 40 kg of sediment was extracted in a SSW-NNE first sampling, washed and sieved, yielding a small vertebrate collection of 239 identifiable remains. This material is stored at the Museu de la Universitat de València d’Història Natural (Burjassot, Spain), with the acronym COLT- (Columbretes). The sediment was water-screened using superimposed 1.5 and * 0.5 mm-mesh screens. The fossils were processed, sorted and classified using a Leica MS5 binocular microscope. Measure- 30 m a.m.s.l. ments were taken on a Leica MZ75 binocular microscope, by means of displacement of a mechanical stage, connected to a 2m Sony Magnescale measuring equipment. Photographs were taken with a HITACHI 4800 scanning electron at the Servei Central de Suport a la Investigació Experimental (SCSIE) of the Universitat de València. Aeolian silt Squamata bones remains were assigned to different taxa Lime mud and silt following the criteria given by Szyndlar (1984), Barbadi- llo (1989), Bailon (1991), Barahona (1996), Barahona & Volcanic clasts ­Barbadillo (1997), Szyndlar & Rage (2002), Villa & Delfino Volcanic agglomerates, (2019a) and Čerňanský et al. (2020). The obtained fossils lapilli and ash were compared with the materials housed in the collec- tions of dry skeletons of the Museo Nacional de Ciencias Fig. 2. — Lithostratigraphy of the sedimentary terrace and emplacement of the sample (asterisk). Abbreviation: a.m.s.l., above mean sea level. Naturales-Consejo Superior de Investigaciones Científicas (MNCN-CSIC; Madrid, Spain), Muséum national d’Histoire naturelle (Paris, France), Gabinet de Fauna Quaternària of COLT-BONES was pretreated with alkali (sodium hydrox- the Museu de Prehistòria de València (Valencia, Spain) and ide, NaOH). The obtained ages were calibrated to calendar Departament de Botànica i Geologia from the Universitat de years (cal BC/AD) and calibrated radiocarbon years (cal BP). València (Burjassot, Spain). The osteological nomenclature Calibration was calculated using one of the databases associ- follows Roček (1984), Szyndlar (1984), Bailon (1991), and ated with the 2013 INTCAL program (Reimer et al. 2013). Russell & Bauer (2008). We follow the taxonomic nomen- clature of Speybroeck et al. (2020) for reptiles and Pasquier Abbreviations (1974) for mammals. All the measurements have been taken c1 posterior accessory cusp; following Barahona (1996) and Barahona & Barbadillo (1997) cal BP calibrated radiocarbon years before present; CL vertebral centrum length, measured ventrally for lacertids, Blain (2009) and Georgialis et al. (2021) for and including the condyle; snakes and Pasquier (1974), Darviche & Orsini (1982) and CL/NAW ratio between the centrum length and the mini- Darviche et al. (2006) for mammals. mum width of the neural arch; In order to identify the agent responsible of the fossil accu- COLT Columbretes palaeontological site; mulation two preliminary taphonomical studies were carried CTH cotyle height; out, one based on the small mammals and the other one only CTW cotyle width; CTW/CTH ratio between width and height of the cotyle; on the squamates fossils. In this sense, digestion marks on small CTW/NAW ratio between cotyle width and the minimum mammal teeth (molars and incisors) have been studied based width of the neural arch; on the descriptive-systematic method developed by Andrews I upper incisor; (1990) and Fernández-Jalvo et al. (2016). Regarding reptiles, L length of the tooth; the study has been based on vertebrae. The digestion marks M1 first upper molar; M2 second upper molar; were observed on the zygosphene, the prezygapophyseal pro- m1 first lower molar; cess, condyles and the neural spine. We follow the categories m2 second lower molar; of digestion established by Lebreton et al. (2020): MNCN-CSIC Museo Nacional de Ciencias Naturales-Consejo 0, non digested; Superior de Investigaciones Científicas; 1, diapophysis and/or parapophysis affected; NAW minimum width of the neural arch, measured at 2, prezygapophyseal processes also affected; the maximum intrazygapohyseal constriction; PO-PO width of the external borders of the articular 3, condyle also affected; surfaces of the postzygapophysis; 4, loss of lamellar bone. PO-PO/NAW ratio between the width of the external borders Two samples, one consisting of bone fragments (COLT- of the articular surfaces of the postzygapophysis BONES) and the other of land snails remains (COLT-SNAILS), and the minimum width of the neural arch; were selected for AMS radiocarbon dating and sent to the PR-PR width of the external borders of the articular surfaces of the prezygapophysis; laboratory of Beta Analytic Inc. located in Miami (United PR-PR/NAW ratio between the width of the external borders States). COLT-SNAILS was pretreated with acid etch (diluted of the articular surfaces of the prezygapophysis hydrogen chloride, HCl). For the extraction of the collagen, and the minimum width of the neural arch; COMPTES RENDUS PALEVOL • 2023 • 22 (18) 413 Marquina-Blasco R. et al. SCSIE Servei Central de Suport a la Investigació Exper- surface between these two processes is concave. On the medial imental; margin, the insertion surface with the frontal and parietal is W width of the tooth; t1, t7, tE tubercles 1, 7 and E; well marked, while on the lateral margin, the impression with TL total length; the postorbital is less deep. ZW zygosphene width; COLT-20 is the preserved anterior part of a right pterygoid ZW/NAW ratio between the width of the zygosphene and (Fig. 3D). The quadrate (posterior), transverse (anterolateral) the minimum width of the neural arch. and palatine (anteromedial) processes are partially broken. However, the latter is subrectangular and shows a thicker SYSTEMATIC PALAEONTOLOGY area in the middle of its platform. The transverse process is subtriangular and has well marked impressions of the ectopter- Class REPTILIA Laurenti, 1768 ygoid on its dorsal surface. The pterygoid recess is deep and Order SQUAMATA Oppel, 1811 U-shaped. The basis of the quadrate process is broken, for Family Scincidae Oppel, 1811 this reason it is not possible to describe it. In lateral view, the Genus Chalcides Laurenti, 1768 transverse process presents a wide and short insertion surface of the pterygoideus muscle. No pterygoid teeth are present. Chalcides bedriagai (Boscá, 1880) The dentaries bear pleurodont, isodonts, and cylindrical (Fig. 3A-H) teeth with flattened or slightly concave lingual surfaces of the crowns and apices oriented in lingual direction (Fig. 3E). In Gongylus ocellatus bedriagai Boscá, 1880: 50 (original description labial view, the teeth appear monocuspid (a slightly pointed or of the species, holotype designated). blunt labial cuspid), but in lingual view, underneath the labial cuspid, a smaller lingual cuspid may be present. The antrum Chalcides bedriagae – Boulenger 1887: 402 (current taxonomy intercristatum is a generally well-marked groove located on established). — Salvador 1998: 152 (lectotype designed). one side and on the other, between these two cuspids. Every Material examined. — 1 right premaxilla (COLT-4); 4 right max- tooth apex shows a transversal groove, only some teeth show illae (COLT-7 and 8); 2 left maxillae (COLT-56); 1 left postfrontal a vertical striation. The most complete dentaries (COLT-1 (COLT-23); 2 right pterygoids (COLT-20 and 24); 3 right dentaries and COLT-3) are characterized by the presence of 17 dental (COLT-1 and 9); 4 left dentaries (COLT-3 and 11); 11 dorsal ver- positions (Fig. 3E). The width/height ratio for all the teeth tebrae (COLT-12 and 13); 4 caudal vertebrae (COLT-14); 1 right coxal (COLT-45); 1 right femur (COLT-47). equals 0.3 (n = 31; δ = 0.07). The coronoid process is subtri- angular, straight and posterodorsally projected. In labial view, this process does not show any impression of the coronoid Description on the wall of the dentary. The studied dentaries present four The preserved premaxilla (COLT-4) is the left one (Fig. 3A). lingual foramina. In lingual view, the Meckelian groove is This remain has three dental positions, but only two teeth open in its whole length and narrower between the eighth- are present. These are pleurodont, isodont, cylindrical, and ninth dental position and the anterior end of the tooth row. monocuspid with a blunt apex. In frontal view, the pos- The subdental shelf is slightly arched and the ventral margin terodorsal process is short and leaf-shaped. No foramen is of the dentary is straight. observable in the bone. The vertebrae are procoelous and longer than wide (verte- COLT-7 is the distal half of a right maxilla (Fig. 3B). bral centrum length (CL), CLmed = 1.94 mm; n= 5; σ = 0.22; Thus, only the posterior and prefrontal processes, although CLmin = 1.69 mm; CLmax = 2.23 mm; minimum width of partially broken, are preserved. In labial view, the prefrontal the neural arch, measured at the maximum intrazygapohyseal process seems to be subpentagonal, with an anterior margin constriction (NAW), NAWmed = 0.98 mm; n = 4; σ = 0.08; broken, a dorsal margin roughly subdivided in anterodorsal NAWmin = 0.86 mm; NAWmax = 1.05 mm). In dorsal view, they and posterolateral portions and a sigmoidal posterior mar- show a few marked interzygapophyseal constrictions (Fig. 3F). gin. A dorsal corner and a posterodorsal corner are clearly The neural arch shows a rudimentary zygosphene-zygantrum visible. The labial surface of the bone is smooth and only articulation, sensu Hoffstetter (1969) and Hoffstetter & Gasc one labial foramen is visible; however, the surface is highly (1969), with two shorts lateral facets (zygosphene) but with damaged so an additional foramen might also be present. In a barely outlined posterior zygantrum. The neural spine runs lingual view, the maxilla bears pleurodont, isodont, cylindri- along the entire length of the neural arch, finishing in a point cal, and monocuspid teeth with a more or less blunt apex. that overtakes the posterior end of the postzygapophysis. In The teeth show a more or less visible ornamentation, with lateral view, the neural spine is low. The synapophyses are delicate vertical striation limited ventrally by a transverse oval-shaped and slightly projected anteroposteriorly. In ven- groove. The supradental shelf is arched and partially broken tral view, the centrum is subcylindrical with its anterior part at the palatine process. slightly wider than the posterior one. It shows neither haemal The postfrontal is Y-shaped (Fig. 3C). The anteromedial and keel nor foramen, except in the case of one vertebra stored anterolateral processes are short and pointed. Both processes under the number COLT-13, which presents a pair of slightly have similar size; however, the anterolateral process is more marked large subcentral foramina. In anterior and posterior robust than the anteromedial one. In ventral view, the ventral view, the condyle and cotyle are dorsoventrally flattened. 414 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island A B C D E F G H K G’ I J L M Fig. 3. — Squamates remains from COLT (Illa Grossa, Columbretes Island, Eastern Spain: A-H, Chalcides bedriagai (Boscá, 1880): A, left premaxilla (COLT-4), in lingual view; B, right maxilla (COLT-7), in lingual view; C, left postfrontal (COLT-23) in ventral view; D, right pterygoid (COLT-21) in dorsal view; E, right den- tary (COLT-1) in lingual view; F, G, dorsal vertebra (COLT-12) in dorsal (F) and anterior (G) views; H, left coxal (COLT-44) in left lateral view; I-M, cf. Podarcis sp. Wagler, 1830: I, J, premaxilla in lingual (I) (COLT-43) and labial (J) (COLT-16) views; K, left maxilla (COLT-19) in lingual view; L, right dentary (COLT-25) in lingual view; M, right coxal (COLT-42) in right lateral view. The white arrow denotes the presence of the rudimentary zygosphene. Scale bars: 0.5 mm. COMPTES RENDUS PALEVOL • 2023 • 22 (18) 415 Marquina-Blasco R. et al. COLT-14 includes two vertebrae, which have a pair of dor- Ecology soventrally flattened transversal processes typical of caudal Chalcides bedriagai is an Iberian endemic species localized vertebrae (Bailon 1991). throughout the Iberian Peninsula, except its northermost COLT-45 is the anterior edge of a coxal, a large elongated area. The distribution range of this taxon is included into the bone (Fig. 3H). In lateral view, on the ilium the preacetabular Mediterranean bioclimatic region, except for some localities spine and dorsal crest are absent. The acetabulum is rounded. of northwestern Iberia; although C. bedriagai always occupies Only the proximal end of the femur is preserved. It shows thermal areas or Mediterranean-like environments. Chalcides a well-developed, ventrally expanded femoral condyle. The bedriagai frequents a wide variety of habitats, which are internal trochanter is small and is not well defined. Both mainly characterized by sandy or earthy substrate with an structures are separated by a large and deep intertrochanteric abundance of shelters (rocks, roots, leaves and shrubs) and fossa in ventral view. high sun exposure, which constitutes a limiting factor of their presence (Pollo 2015). Remarks The traits described on the material (unfused premaxilla, morphology of the teeth, straight ventral end of the den- Family Lacertidae Oppel, 1811 taries, open Meckelian groove, presence of a thicker area in the palatine process of the pterygoid, vertebrae with the end Genus Podarcis Wagler, 1830 of the neural spine overtaking the posterior of the postzyga- pophysis, and absence of the preacetabular spine and dorsal Lacerta (Podarcis) Wagler, 1830: 155. crest in the coxal) are consistent with the genus Chalcides Podarcis – Arnold 1973: 343. (Barbadillo 1989; Bailon 1991; Caputo et al. 1995; Blain 2009; Villa & Delfino 2019a; Čerňanský et al. 2020). At Type species. — Seps muralis Laurenti, 1768, subsequent designa- tion by Fitzinger (1843). present, there are three species of that genus in the Iberian Peninsula, the autochthonous Chalcides striatus (Cuvier, 1829) and Chalcides bedriagai, and the introduced Chalcides Notes ocellatus Forsskal, 1775 (Bisbal-Chinesta et al. 2020). In the Böhme (1997) regarded the gender of Podarcis to be masculine; latter species, teeth are robust with enlarged crowns (Kosma Arnold (2000) proposed to retain the gender of the genus as 2004; Villa & Delfino 2019a). This morphology differs femenine; and Böhme & Köhler (2005, current taxonomy) strongly from other members of the genus Chalcides (Kosma established it as masculine gender. 2004). As we describe above, the COLT fossils do not show these traits on the teeth. Their morphology and width/high proportion do not differ from those of the modern species cf. Podarcis sp. Chalcides bedriagai, which is characterized by 15 to 18 dental (Fig. 3I-M) positions and a width/height ratio of 0.3 (Barbadillo 1989; Caputo 2004); in contrast to C. striatus, which is character- Material examined. — 4 premaxillae (COLT-15, 16 and 43); 4 left maxillae (COLT-18, 19 and 39); 2 right maxillae (COLT-17); ized by 18 to 21 dental positions (Caputo 2004) and a tooth 2 left pterygoids (COLT-21 and 22); 1 dorsal vertebra (COLT-40); width/height ratio of 0.22 (López-Ruiz et al. 2002). 1 possible sacral vertebra (COLT-53); 2 caudal vertebrae (COLT-41); The fossil record of the genus Chalcides in the Mediter- 1 centrum of caudal vertebra (COLT-48); 1 right coxal (COLT-42); ranean islands is scarce. In this sense, cf. Chalcides has been 2 left humeri (COLT-45 and 57); 1 right humerus (COLT-46). reported from the late Miocene/early Pliocene of Menorca (Bover et al. 2014) and indeterminate species of the genus Description had been reported from the Pleistocene of Menorca (Estes The premaxillae of this species are represented in the Colum- 1983), and Mallorca (Holman 1998). Skinks are highly capa- bretes Islands by four small-sized remains, with an average ble of transmarine migration, as shown by the diversification with of the premaxillary shelf of 1.23 mm (n = 3; σ = 0.13). of the genus Chalcides in the Canary Islands. However, in the The posterodorsal process is thin and has parallel lateral mar- case of the Balearic Islands sites, it has been proposed that gins (Fig. 3I). The most complete specimen (COLT-43) has Chalcides colonized the archipelago during the Messinian seven dental positions, but only three teeth are present. These Salinity Crisis (MSC), when these islands were connected to are pleurodont, isodont, cylindrical, and monocuspid with a the mainland (Carranza et al. 2008). A similar scenario was blunt apex. COLT-16 shows the distal end of the posterodorsal proposed for the origin of the endemic herpetofauna of the process slightly expanded (Fig. 3J). The nasal foramina are Balearic Islands (Pinya & Carretero 2011), and the spread present at the base of the posterodorsal process. In posterior of Chalcides bedriagai in the Iberian Peninsula (Carranza view, this process has a well-defined medial crest. et al. 2008). The MSC event had a strong influence in the The maxilla bears pleurodont, isodont, cylindrical, mono or Mediterranean herpetofauna (e.g. Georgialis et al. 2019a). bicuspid teeth with a blunt apex. In dorsal view, the remains Currently, C. bedriagai inhabits numerous islands close to both have a wide palatal shelf with a large superior dental foramen. the Atlantic and Mediterranean coasts of the Iberian Peninsula The anterior margin of the maxilla has short anteromedial (Pollo 2015; and references in Ruiz-Sánchez et al. 2019). and anterolateral processes. The two processes are similar in 416 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island size; however, the anterolateral process seems to be slightly procoelous vertebra with the centrum transversally convex larger than the anteromedial one. The notch formed on the and the neural spine not exceeding the posterior limit of anterior concavity between both processes is shallow. A wide the postzygapophysis; coxal with preacetabular spine but vomeronasal foramen is located in the basis of the anterior no dorsal crest, humerus with a laterally flattened radial margin of the dorsal process. In lingual view, the anteromedial condyle, rounded ulnar condyle, and the presence of an process does not show any lappet. However, in COLT-39 the ectepicondylar foramen (Bailon 1991; Barahona 1996; Rus- anteromedial process, although broken, seems to be elevated sell & Bauer 2008). Although the osteology of the lacertids and, probably, formed a lappet. It is impossible to evaluate is poorly studied, some taxonomic ascription can be tried. the degree of development of the eroded prefrontal process. Thus, the presence of a step in the posterior process of the The posterior process is short and with a step on its dorsal maxilla is a characteristic reported by Barahona & Barbadillo margin (Fig. 3K). In labial view, the surface of the maxilla is (1997) and Villa & Delfino (2019a) for the lacertid genera smooth and shows three labial foramina. Iberolacerta Arribas, 1997, Lacerta Linnaeus, 1758, and The dentaries are fragmentary (Fig. 3L). The recovered Podarcis. Regarding the premaxilla, the parallel position of remains bear pleurodont, isodont, cylindrical, bi- and mono- lateral margins of the posterodorsal process and the number cuspid teeth, with a more or less pointed principal cuspid. In of dental positions exclude some lacertid species, reducing lingual view, the Meckelian groove is open and the subdental the options to Podarcis, Iberolacerta, Psammodromus Fitzinger, shelf is arched. 1826 and Acanthodactylus Wiegmann, 1834 (Barahona & The vertebrae are procoelous and longer than wide. In Barbadillo 1997). The last two can be discarded due to the dorsal view, they show a marked interzygapophyseal constric- absence of a step in the posterior process of the maxilla tion. In the foremost part, the remains show a rudimentary (Barahona & Barbadillo 1997; Villa & Delfino 2019a). The zygosphene-zygantrum articulation, sensu Hoffstetter (1969) mean size of the premaxilla shelf is included in the range of and Hoffstetter & Gasc (1969). The neural spine runs along the Podarcis species provided by Barahona (1996), except the entire length of the neural arch, except its first quarter. The for Podarcis bocagei (Seoane, 1884), considered as Podarcis neural arch terminates in a point, which does not overtake the bocagei bocagei in the refereed work, and Iberolacerta bonnali postzygapophysis. In lateral view, the neural spine is low. The (Lantz, 1927), in which sizes are larger than those recorded vertebra has an oval-shaped synapophysis, which is slightly for the COLT lacertid premaxilla. On the other hand, the anteroposteriorly projected. The ventral margin of the cen- number of premaxillary teeth (7) restrains the options to trum is convex. In ventral view, the centrum is subtriangular Podarcis muralis (Laurenti, 1768), P. bocagei, Podarcis his- in shape with its anterior part wider than the posterior one. panica sensu lato (Steindachner, 1870) and Iberolacerta The haemal keel is absent. A pair of small not well-defined bonnali (Lantz, 1927) (Barahona & Barbadillo 1997). Fol- subcentral foramina is present. In anterior and posterior view, lowing biogeographic criteria, the presence of a species of the condyle and the cotyle are small and have a subcircular Iberolacerta seems unlikely because these lacertids are almost shape. COLT-48 consists in a centrum of a procoelous caudal entirely confined to small widely separated mountain areas vertebra, slightly wider than longer with well-developed and (e.g. I. bonnali only inhabited the Central Pyrenees) and they dorsoventrally flattened transverse processes. tend to be taxa adapted to cold environments (Ortega et al. The coxal is a large elongated triradiate compound bone 2016). Thus, all these characteristics permit an attribution (Fig. 3M). It is formed by the complete fusion of the ilium, of the COLT fossils to cf. Podarcis sp. ischium, and pubis, without any trace of suture line. In the The fossil record of lacertids is known in the larger Medi- posterior region of the ilium, a hooked preacetabular spine terranean islands (Bailon 2004). The insular record of the is dorsally projected. The coxal does not show any dorsal genus Podarcis is limited to Crete, Sicily, Sardinia, Corsica, crest. The acetabulum is oval and large. The pubis is broken. Mallorca, Ibiza, and Menorca (Bailon 2004; Villa & Delfino Although the ischium is broken, it seems to be large and 2019b). Several works (Bover et al. 2014; Rodríguez et al. wide. It forms a posterior angle of about 90° with the ilium. 2017; Spilani et al. 2019) proposed that the ancestors of the The recovered humeri have only the distal end preserved. current Podarcis species that inhabit these islands reached them The epiphysis is wider than the diaphysis. In ventral view, it via land bridges. However, the populations of Podarcis siculus shows a deep radioulnar fossa. The radial condyle is oval in (Rafinesque-Schmaltz, 1810), present in Menorca, Sardinia shape and transversely compressed, whereas the ulnar condyle and Corsica, have been introduced by humans in historical is slightly larger and rounded. The ectepicondyle is not well times (Silva-Rocha et al. 2018). differentiated. The entepicondyle is reduced. The entepicon- dylar foramen is large. Current ecology Podarcis is a circummediterranean genus that is widespread in Remarks Southern Europe, the Anatolian Peninsula and Northern Africa The fossils described here show typical traits of Lacertidae: (Spreybroeck et al. 2016). At least, 12 species are endemic fused premaxilla with a slender posterodorsal process; pleu- of the Mediterranean islands, especially in the Aegean Sea rodont, isodont and mono or bicuspids teeth; ventral edge (Spreybroeck et al. 2020). At present, the Columbretes archi- of the dentary concave in lingual view; Meckelian groove pelago is inhabited by Podarcis liolepis atratus (Boscá, 1916), of the dentary open along its whole length in lingual view; until recently considered as an endemic insular s­ubspecies COMPTES RENDUS PALEVOL • 2023 • 22 (18) 417 Marquina-Blasco R. et al. A B C D Fig. 4. — Vipera cf. latastei Boscá, 1878 from COLT (Illa Grossa, Columbretes Island, Eastern Spain): A, solenoglyph fang (COLT-2); B-D, precaudal vertebra (COLT-5) in anterior (B), posterior (C) and right lateral (D) views. Scale bars: 1 mm. (Ruiz-Sánchez et al. 2019 and references therein). This lizard Description is present in all islands and islets of the Columbretes, except The fangs are strongly curved (Fig. 4A). Their anterior surface in the Ferrera group (Castilla et al. 1998a). The absence of shows an elongated, slender venom discharge distal orifice, the species in Ferrera is not understood, since this island which is long and narrow. The pulp cavity and a canal, with include potential habitats for this lizard. The presence of a circular cross-section, can be observed in the teeth. The fossils of a Wall Lizard (Podarcis sp.), or related taxa, in a canal is completely closed, without any groove or suture in palaeontological context confirms the ancient occupation of the external surface of the teeth. the Columbretes Islands by this taxon. The open taxonomi- COLT-49 is a fragment of dentary bone with three dental cal attribution proposed for the lacertid fossils does not allow positions, but only two teeth preserved. The teeth, which are further inferences about the palaeoecological context during inserted on small alveoli, are conical and curved posteriorly. the formation of the site. The precaudal vertebrae are small-sized (CLmed = 2.76 mm; n = 5; σ = 0.43; CLmin = 2.10 mm; CLmax = 3.21), procoelus and longer than wide. In anterior view, the zygosphene is Sub-order SERPENTES Linnaeus, 1758 straight (Fig. 4B). The cotyle is rounded and large. A pair of Clade CAENOPHIDIA Hoffstetter, 1939 large and well-defined paracotylar foramina are present. The Family Viperidae Oppel, 1811 articular facets of the prezygapophyseal process is dorsolaterally Genus Vipera Laurenti, 1768 inclined. In posterior view, the neural arch is dorsoventrally flattened (Fig. 4C). The condyle is rounded and large. Its Vipera cf. latastei Boscá, 1878 diameter is smaller than the neural arch one. In dorsal view, the (Fig. 4) vertebrae show a trilobated zygosphene, where the central and the lateral ones are of similar size. The neural spine runs along Vipera latastei Boscá, 1878: 121 (original description of the spe- the whole extension of the neural arch. The prezygapophyses cies). — Boscá 1879: 30. are anterolaterally projected, whereas the postzygapophyses Vipera berus aspis var. latastei – Camerano 1889: 231. are posterolaterally projected. The prezygapophyseal process is short and pointed. In ventral view, the centrum is triangular Vipera latastii [sic] – Boulenger 1896: 484. in shape, wider anteriorly than posteriorly and with not well- defined lateral margins. The subcentral foramina are situated Rhinaspis (Latastea) latastei Reuss, 1930: 69. in the anterior part of the centrum and are large and well defined. In lateral view, the neural spine is high (Fig. 4D). Vipera ammodytes latastei – Saint-Girons 1977: 605 (the author pointed out that V. latasti was a lapsus calami, thus he proposed that The lateral margins of the neural arch are not well defined. the correct name is V. latastei; designation of a lectotype). — ICZN The prezygapophyses are anterolaterally projected, whereas 2017: opinion 2381 (case 3629), confirmation that V. latastei is the the postzygapophyses are posterolaterally projected. The correct specific name for the taxon. parapophyseal process is well defined and antero-ventrally projected. The inferior margin is well defined. The centrum Material examined. — 15 venom fangs (COLT-2 and 10); 1 left dentary (COLT-49); 8 precaudal vertebrae (COLT-5, 6, 50 and is transversally convex. The hypapophysis is broken, but 52); 8 centra of precaudal vertebrae (COLT-51); 1 caudal vertebra seems to be long and straight. In ventral view, the centrum (COLT-53). is triangular in shape. It is wider anteriorly than posteriorly. 418 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island A B C Fig. 5. — Mus cf. musculus Linnaeus, 1758 from COLT (Illa Grossa, Columbretes Island, Eastern Spain): A, right m1 (COLT-31); B, left m1 (COLT-37); C, right M1 (COLT-38). Both pictures are in occlusal view. Scale bars: 0.5 mm. The lateral margins of the centrum are not well defined. The from the COLT fossils are broken, and the posterior edge of subcentral foramina are situated in the anterior part of the the neural spine is preserved only in COLT-5, where it is high centrum and are large and well defined. The condyle and the (higher than the centrum in lateral view) as in the “V. aspis cotyle are large and rounded. In posterior view, the diameter complex” members. For morphological, palaeobiogeographi- of the condyle is smaller than the diameter of the neural cal, and biostratigraphical reasons, the species present at the arch. The presence of a pair of haemapophysis indicates that Illa Grossa Island most probably was Vipera latastei. Therefore COLT-53 belongs to a caudal vertebra. we propose here an open attribution to this taxon until new fossils can definitively settle this problem. This attribution Remarks is more extensively discussed in the section on the identity The recovered fossils are attributable to Vipera by: the pres- of the Columbretes snakes below. ence of a venom fang of solenoglyph type without any groove or suture in the external surface of the tooth; the presence of a closed venom canal which originated from the dentine Class MAMMALIA Linnaeus, 1758 floding throughout ontogeny; dorsal vertebrae with hypa- Order RODENTIA Bowdich, 1821 pophysis, neural arch dorsoventrally depressed, vertebral Family Muridae Illiger, 1811 centrum longer than wide, ventrally convex centrum with Genus Mus Linnaeus, 1758 diffuse lateral margins, well developed condyle and cotyle, parapophyseal processes present, and the articular surface Mus cf. musculus Linnaeus, 1758 of the pre and postzygapophyses dorsally inclined (Szynd- (Fig. 5) lar 1984; Bailon 1991; Bailon et al. 2002; Georgalis et al. 2019a). According to the size of the examined vertebrae (CL: Mus musculus Linnaeus, 1758: 62 (original description of the species). 2.76 mm), the material from Illa Grossa belongs to a “Euro- Mus domesticus Rutty, 1772: 281. pean Viper” (CL < 5mm). Within the latest group, there are two complexes that differ in their vertebral morphology: the Mus abbotti Watherhouse, 1837: 77. “Vipera berus complex” and the “V. aspis complex” (Szynd- Mus nordmanni – Keyserling & Blasius 1840: 330 (nomen nudum). lar & Rage 1999, 2002). Their isolated trunk vertebrae may be distinguished by their shape and size. The vertebrae of the Mus molossinus Temminck, 1845: 51. “V. berus complex” are small, elongated, and provided with low neural spine and hypapophyses, whereas the vertebrae of Mus varius – Fitzinger 1867: 70 (nomen nudum). the “V. aspis complex” bear more developed hypapophyses and Mus musculus yonakuni Kuroda, 1924: 8. — Kaneko & Maeda neural spine (Szyndlar & Rage 1999). In the material from 2002: 12. COLT, the hypapophyses are broken but they appear to be straight and well-developed; this trait, together with the pres- Mus albula Kishida, 1924: 143. — Kaneko & Maeda 2002: 12. ence of a high posterior edge of the neural spine (higher than Mus bactrianus tantillus Allen, 1927: 9. — Schwarz & Schwarz the centrum in lateral view), allow us to attribute the remains 1943: 62 (reviewed along with the rest of the genus). — Ellerman & to the “V. aspis complex”. Unfortunately, the hypapophyses Morrizon-Scott 1951: 607. COMPTES RENDUS PALEVOL • 2023 • 22 (18) 419 Marquina-Blasco R. et al. Material examined. — 3 I (COLT-33 to COLT-35); 1 M1 Description (COLT-30); 1 M2 (COLT-29); 2 m1 (COLT-28, COLT-31); 1 m2 Among the recovered vertebrate remains, a total of 35 bone (COLT-32). fragments could be attributed to birds, all small-sized, and 30 of them belonging to small passerines. Unfortunately, Description these bone remains lack diagnostic features allowing a more Upper incisors are rounded and curved. precise taxonomic ascription. Among the recovered bones is The recovered first upper molars are brachyodont and buno- a proximal part of humerus, but its poor preservation state dont, and do not present the t7. The t1 is placed towards the prevents its identification. The remaining five bones belonged posterior part of the tooth. to seabirds. Three fragments of pedal phalanges belonged to The first lower molars are brachyodont and bunodont and shearwaters (Procellariidae), and a wing phalanx and a cranial there is no anterocentral tubercle. On the anterior region, part of scapula to very small species of other families within the tE is not well individualized, and it develops a trilobed the procellariiforms group. All the recovered bones correspond morphology in one specimen (COLT-31). COLT-28 has its tE to adult individuals. broken, while COLT-37 is quite worn. No external cingular margin with a well-developed secondary cusp (c1), is present Remarks in any of the three specimens. The recovered avian fossils comprise at least six undetermined The second molars are also brachyodont and bunodont, with species of birds (two of seabirds and four passerines). Currently, no trace of the antero-labial tubercle. The labial cingulum is many of the birds that make their migratory trips along the not developed, and the posterior tubercle is big, round or oval. eastern coast of the Iberian Peninsula arrive to the Colum- bretes due to adverse weather conditions, particularly, with Remarks westerly winds. For this reason and despite their small surface, The absence of the t7 tubercle in the upper first molars is these islands are home to a varied ornithocenosis, made up of characteristic of the genus Mus, contrary to what it is seen in dozens of passerine species. They constitute also a safe place the genus Apodemus Kaup, 1829, in which the t7 is always for breeding colonies of several marine species of birds (Díaz present (Chaline 1974). Moreover, the first lower molars et al. 1996). It can be assumed that the avian ranges did not recovered from in Columbretes lack the anterocentral tubercle, undergo major changes in the region during the late Holocene. as it is the case in Mus (Pasquier 1974; Gosàlbez 1987). The trilobed morphology observed in the anterior part of the first lower molars is characteristic of Mus musculus (Darviche & DISCUSSION Orsini 1982). In addition, the clear absence of an external cingular margin with a well-developed secondary cusp in Chronology the first lower molars also points to M. musculus, whereas it The two radiocarbon (14C) dated samples of the site yielded is quite common in Mus spretus Lataste, 1833 (Darviche & similar chronologies: 2780-2724 cal BP for COLT-SNAILS Orsini 1982; Darviche et al. 2006). Regarding the metrics of and 2600-2492 cal BP COLT-BONES. In general, terrestrial the first lower molars, the mean length and width values fall snails provide older 14C shell dates (Evin et al. 1980), which is between Mus spretus lowest mean values (Fig. 6A), although attributed to the fact that many taxa incorporate 14C-deficient if we take into account specimens values, those are placed in carbon from limestone and other carbonate rocks when build- an overlapping area between both species values (Fig. 6B) ing their shells, the so-called “Limestone Problem” (Good- (Darviche & Orsini 1982; Domínguez García et al. 2019). friend & Stipp 1983). However, this phenomenon seems to Species within the genus Mus can be distinguished based be related with large specimens (Brennan & Quade 1997) on morphotypes percentages. Unfortunately, the very low and is strongly dependant of the species (Pigati et al. 2010). population size does not allow us to extract percentages. How- Moreover, small land snails recovered from fossil deposits in ever, all morphologic characters are typical of Mus musculus. arid environments, as it is the case of the Columbretes Islands On the other hand, metric values are compatible with some (Castilla & Bauwens 1991), may yield reliable 14C ages (Pigati big individuals from a Mus musculus population as shown in et al. 2010). The sample of terrestrial gastropods used in the the record (Darviche & Orsini 1982). Due to the aforemen- present study was composed of a mixture of shell fragments tioned reasons, we ascribe the studied mammal material to of small and large species. Mus cf. musculus. Fossil assemblage The COLT site has provided a rich fossil assemblage of ter- Class AVES Linnaeus, 1758 restrial gastropods and vertebrates (fishes, reptiles, birds and mammals). The only vertebrates previously described here are Aves indet. spp. dermal denticles of Raja clavata Linnaeus, 1758 or a very close taxon (Ferrón et al. 2015). The gastropod remains recovered Aves Linnaeus, 1758: 12, 78-80. from the COLT site are currently under study. A preliminary examination reports the occurrence of several families of ter- Material examined. — 58 undetermined remains (COLT-58 to 92). restrial snails: Ferussaciidae Bourguignat, 1883; Geomitridae 420 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island A Populations B 1.00 Individuals M. spretus 14 9 0.96 M. spretus 8 0.95 COLT-31 13 0.93 COLT 12 COLT-28 7 W (mm) W (mm) 15 COLT-37 0.90 11 6 0.90 10 0.87 M. m. domesticus5 4 1 0.85 0.84 2 3 M. m. domesticus 0.81 1.41 1.44 1.47 1.50 1.53 1.56 1.59 1.62 1.45 1.50 1.55 1.60 1.65 L (mm) L (mm) Modern N. Africa Modern France L Modern Spain Holocene Spain L. Pleistocene Morocco W Fig. 6. — Scatterplot comparing mean length (L) and mean width (W) of first lower molar from Mus m. domesticus Linnaeus, 1758 (yellow) and Mus spretus Lataste, 1883 (blue) in several localities from the Late Pleistocene until recent time in the western mediterranean area: A, mean values for populations (modi- fied from Domínguez-García et al. 2019); B, values for individuals (modified from Darviche & Orsini 1982). Numbers: 1, Azemour (Morocco) (n = 14); 2, Toulouse (n = 16); 3, ­Algeria (n = 14); 4, Tunisia (n = 19); 5, Southern France (n = 29); 6, Granada (n = 20); 7, Montpellier (n = 22); 8, Southern France (n = 33); 9, Southern France (n = 7); 10, Tunisia (n = 25); 11, Morocco (n = 40); 12, Morocco (n = 41); 13, Doukkala (n = 17); 14, El Harhoura II (n = 263); 15, Cerro de la Encantada (n = 14). Abbreviation: CB, Castillejo del Bonete (n = 164). Sources: 1-4, 7, 10, 11, 13, Darviche et al. 2006; 5, 8, Darviche & Orsini 1982; 6, Morales Muñiz 1986; 9, Gerasimov et al. 1990; 12, 14, Stoetzel 2009; Stoetzel et al. 2010. Credits: modified from Domínguez-García et al. 2019. Boettger, 1909; Helicidae Rafinesque, 1815; Lauriidae Steen- as snakes by non-expert observers due to of their elongated berg, 1925; Pomatiidae Newton, 1891; Punctidae Morse, body. Their extirpation on the island most probably took 1864; and Vertiginidae Fitzinger, 1833. Fossils of marine place before the start of the scientific study of the islands (last gastropods (Littorinidae Children, 1834) are also present. third of the 19th century) (Ruiz-Sánchez et al. 2019) and, in A total of 239 remains of reptiles, mammals, and birds have general, could be related with the profound human-induced been studied in this paper. The fossil assemblage determined changes that affected the terrestrial ecosystems. in the present work is composed by two lizards (Chalcides The description of the vegetation of the Illa Grossa Island in bedriagai and cf. Podarcis sp.), a snake (Vipera cf. latastei), a some classical texts (Smyth 1831; Salvator 1895; Bru 1913) rodent (Mus cf. musculus) and, at least, six species of birds pointed out the presence of Opuntia ficus-indica Miller, 1754, belonging to seabirds (two) and passerines (four). an invasive species in Mediterranean and semiarid areas (Sanz- The COLT association clearly differs from the present Elorza et al. 2004), which is still present in the Columbretes terrestrial fauna of Illa Grossa, which is limited to a single Islands (Mestre et al. 2010). The colonization of the archi- non-flying vertebrate species: the Columbretes’ Wall Lizard, pelago by this species could be early, since O. ficus-indica was Podarcis liolepis atratus (Jiménez et al. 2002; Mestre et al. already present in the coastal areas of eastern Spain in the 2010). Castilla & Bauwens (2000) noted that the divergence 18th century (Sanz-Elorza et al. 2004). However, the nega- within the Columbretes’ lizard population took place at most tive impacts of the modifications in the terrestrial biological one million years ago. The oldest reference about Wall Lizards communities (fires and introduction of exotic species) that in the Columbretes Islands is provided by Salvator (1895), were produced in the Illa Grossa during the construction of whereas the occurrence of snakes was already reported by the the lighthouse in the 19th century cannot be ruled out. In Ancient Greeks (Bernis 1968; Silva-Rocha et al. 2018). In this this sense, some populations of C. bedriagai have undergone sense, the remains described here are the oldest specimens of strong recessions or have disappeared due to the human- Wall Lizards in the Columbretes Islands. induced habitat alteration in different areas of the Iberian One of the most remarkable discoveries in the COLT Peninsula (Jiménez et al. 2002; Pollo 2015). association is the presence of Chalcides bedriagai, since no historical reports the presence of skinks in the Columbretes About the identity of the snakes Islands. This lack of reports in the Illa Grossa Island might from the Columbretes archipelago be related to the cryptic behaviour of this species, which Although the historical abundant presence of snakes in the remains hidden most of the time (López-Jurado et al. 1978). Columbretes archipelago is a widely known fact, their identity Alternatively, some specimens could have been misidentified remains unclear and is a matter of debate since the middle COMPTES RENDUS PALEVOL • 2023 • 22 (18) 421 Marquina-Blasco R. et al. of the 20th century (Mertens 1957; Klemmer 1961; Bernis resulting in an easy confusion between both species. Moreo- 1968; Ruiz-Sánchez et al. 2019). Herpetologists have worked ver, there are islands populations close to the mainland in the with two hypotheses: Atlantic and Mediterranean coast of Europe and North Africa Colubrid hypothesis: in the middle of the last century, (Mateo 1990, 1997; Mateo et al. 2009) with the presence of Mertens (1957) and Klemmer (1961) proposed that the snakes this species. On the other hand, the N. maura populations from from the Columbretes Islands were actually colubrids. These larger islands, such as Mallorca or Menorca, were probably researchers based its proposal on: 1) absence of bite reports. introduced by humans (Corti et al. 1999). Additionally, it With such a high density of alleged vipers, it is difficult to is known that N. maura is capable to exploit marine resources explain why none of the lighthouse construction workers were in many areas of its distribution range (Fuentes & Escoriza ever bitten. However, according to Salvator (1895), the her- 2015). A few years ago, a snake vertebra was recovered from petologist Eduard Boscà claimed that, in fact, these incidents the sediment form the around of the cistern area in the did occur; 2) the high densities reported in the historical texts Columbretes Islands. Altough preliminarily attributed to the could only be supported by a wide availability of terrestrial genus Coronella Laurenti, 1768 (Jiménez et al. 2002), this prey, but theoretically this does not seem to be the case at remain is currently lost and, to the best of our knowledge, the Illa Grossa Island. Therefore, the colubroid species must no photograph or description that can support or refute this have exploited other resources of marine origin (fishes and determination (I. Lacomba, comm. pers.). crustaceans); and 3) there are no cases of insular populations Viperid hypothesis: The classical reports identified the of vipers in islands similar to Illa Grossa. At present, there are species as a viper (Vicent 1885; Bru 1913; Bernis 1968), several island populations of vipers in the Mediterranean Basin some of them with doubts (Salvator 1895; Español 1958). (Speybroeck et al. 2016). Thus, in the Aegean Sea, Vipera In this sense, no direct rest is available to confirm or discard ammodytes (Linnaeus, 1758) is present on numerous Cycladic this identification. The MNCN-CSIC houses a supposedly islands, Macrovipera schweizeri (Werner, 1935) inhabits the preserved specimen (reference number 468 in the herpetologi- eastern Cyclades, Macrovipera lebetinus (Linnaeus, 1758) is cal collection), but the history of its origin is controversial. present in Cyprus, but its autochthony is still under debate The original identification label of the specimen is currently (Torres-Roig et al. 2020), and, finally, Montivipera xanthina lost, and the copy presents some typographic errors. In fact, (Gray, 1849) is present on numerous Greek islands off the the collector’s name (“Sr. Mojaelos”) is erroneous because Turkish coast. All seem to have an origin related to natural nobody with that name is present in any workers’ list during dispersal from the Anatolian mainland to the islands after the construction of the lighthouse in the Illa Grossa Island. the formation of the mid-Aegean Trench (Poulakakis et al. As Bernis (1968) points out, it is possible that, actually, 2015). In contrast, the origin of viper populations from the this name was an erroneous transcription of Mojades, the western Mediterranean islands is much less clear. While it surname of one of the engineers who were in charge of the has been demonstrated that the population of Vipera aspis works between 1856 and 1857. Bernis (1968) did not found (Linnaeus, 1758) on the island of Montecristo was introduced any differences between the conserved specimen and Lataste’s by humans around the 5th century, it is not clear whether viper individuals from south-eastern Spain. The only direct the origin of the populations on the island of Elba is due to description of living individuals, which is compatible with natural dispersal during episodes of marine transgression or vipers, was written by Smyth (1831). to human-induced dispersal (Masseti & Zuffi 2011). As for As it has been described above, the morphology of the the colonization of Sicily by V. aspis, it seems to be result recovered material in the COLT site belongs to a viperid of from the low-stand sea level during the Pleistocene (Barbanera the genus Vipera. During the Pleistocene, the fossil record of et al. 2009). In this sense, the anthropogenic introduction of vipers in the southern half of the Iberian Peninsula is limited snakes in western Mediterranean islands seems to be a recur- to the Vipera aspis complex, while the remains of the Vipera rent event in the historical times (i.e., Insacco et al. 2015; berus group is located in the northern Iberian region (Szyn- Silva-Rocha et al. 2018). However, natural colonization has dlar & Rage 2002; Böhme & Ilg 2003). The same pattern also been documented on Western Mediterranean islands. is observable in the current distribution of the three Iberian This is the case of the today extinct Vipera latastei ebusitana vipers (Pleguezuelos et al. 2002). In this sense, the only viper Torres-Roig, Alcover & Bailon, 2020, from the Late Pleis- that inhabits southern Iberia is Vipera latastei, and it is the tocene-Holocene of Es Pouàs (Ibiza, Balearic Islands, Spain) only species reported from palaeontological and archaeological (Torres-Roig et al. 2020). These authors proposed that the sites during the Pleistocene. Although there are some reports arrival of the ancestor of this taxon occurred through oversea of Vipera aspis in some localities of north-eastern (North of rafting between 1.31 Ma and c. 30 000 BP. the province of Barcelona, Catalonia) and north-central Spain Thus, basing on the three reasons described above, Mertens (North of the province of Burgos, Castilla y León) (Bailon (1957) and Klemmer (1961) ascribed the snakes from the 1991; Blain & Bailon 2006; Blain 2009; Bisbal-Chinesta & Columbretes Archipelago to a natricine colubroid; specifically Blain 2018), none of these reports are located below 40°N Klemmer (1961) suggested that it probably was Natrix maura latitude. Thus, basing on those criteria, together with the his- Linnaeus, 1758, one of the most common snake species in torical reports and the specimen stored in the MNCN-CSIC, the Iberian Peninsula. In this sense, N. maura displays a viper- it is highly probable that the vipers that inhabited the Illa like defensive behaviour mimicry (Aubret & Mangin 2014), Grossa Island until the 19th century belonged to V. latastei. 422 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island Table 1. — Fossil record of Viperidae in Mediterranean islands. In bold, taxa included in the Vipera aspis group. Taxon Locality Chronology Island Reference Vipera sp. (Vipera aspis group) Oschiri Early Miocene Sardinia Venczel & Sanchíz 2006 Viperidae indet. Aliveri Early Miocene (MN4) Euboea Georgalis et al. 2019b Viperidae indet. Punta Nati 2 Middle Miocene Menorca Bailon et al. 2002 Viperidae indet. (Oriental viper) Gargano palaeoisland Late Miocene-Early Pliocene – Delfino 2002 Vipera sp. Punta Nati 3 and 12 ?Pliocene Menorca Bailon et al. 2002 Vipera natiensis Punta Nati 12 Pliocene Menorca Bailon et al. 2002 Viperidae indet. (Oriental viper) Caló den Rafelino Pliocene Mallorca Bailon et al. 2010 cf. Vipera natiensis Caló den Rafelino Pliocene Mallorca Bover et al. 2014 Viperidae indet. (Oriental viper) Na Burguesa-1 Zanclean (Early Pliocene) Mallorca Torres et al. 2014 Vipera sp. Capo Mannu D1 LF Late Pliocene Sardinia Delfino et al. 2011 Vipera sp. Monte Tuttavista Late Pliocene-Early Pleistocene Sardinia Abbazzi et al. 2004 Vipera sp. (Vipera berus group) Laghada B Pleistocene Kos Szyndlar 1991 cf. Vipera sp. Cova de ca na Reina Calabrian (Early Pleistocene) Ibiza Torres-Roig et al. 2020 Viperidae indet. (Oriental viper) Latomi Middle Pleistocene Chios Szyndlar 1991 Vipera sp. (Vipera aspis group) Contrada da Pianetti Late Pleistocene Sicily Delfino 2002 Vipera sp. (Vipera aspis group) K22 (San Vito Lo Capo) Late Pleistocene/Holocene Sicily Delfino 2002 Vipera latastei ebusitana Es Pouàs Late Pleistocene to Holocene Ibiza Torres-Roig et al. 2020 Viperidae indet. (Oriental viper) Khirotikia Holocene Cyprus Watson et al. 1977 Macrovipera lebetina Akrotiri-Aetokremnos Holocene (Mesolithic) Cyprus Bailon 1999 Macrovipera lebetina Kisonerga-Myloudia Holocene (Neolithic) Cyprus Croft et al. 2017 Vipera ammodytes Sanctuary of Poseidon Holocene (Archaic-Hellenistic) Poros Lymberakis & Iliopoulos at Kalaureia 2019 Vipera sp. (Vipera aspis group) Vallone Inferno Holocene (Early Bronze-Late Sicily Forgia et al. 2013 Roman) Vipera cf. latastei Columbretes Holocene (Iron Age) Illa Grossa This work As mentioned above, we propose an open specific attribution Vipera aspis group has been reported in the Late Pleistocene until more remains are recovered and studied. The fossils here of Contrada da Pianetti; the Late Pleistocene/Holocene from studied differ from the recently established Vipera latastei K22 (San Vito Lo Capo), and in the Early Bronze and Late ebusitana, from the Late Pleistocene–Holocene of Es Pouàs Roman levels (3.1 and 3.4, respectively) of Vallone Inferno (Ibiza, Balearic Islands, Spain) (Torres-Roig et al. 2020) in (Forgia et al. 2013), all these sites from Sicily (Delfino 2002), the following traits: presence of a trilobate zygosphene with and, finally, the above mentioned Vipera latastei ebusitana of the central lobe of similar size to the lateral ones in dorsal Es Pouàs (Ibiza) and cf. Vipera sp. from the Cova de ca na view; well-defined parapophyses, and, well-defined subcen- Reina (Ibiza) (Torres-Roig et al. 2020). tral ridge. All these features are similar to those described for Vipera latastei latastei Boscà, 1878 (Torres-Roig et al. 2020). Taphonomical approach However, the COLT fossils may belong to small specimens The low number of mammal remains recorded at Illa Grossa (with CL slightly longer than 3 mm), with biometric indexes exhibit digestion traits in some of the molars (absent, light in the size range of V. l. ebusitana and not of V. l. latastei and moderate categories) that allow us to establish a preda- (see Appendix). tion origin for those remains but prevent us from any further The palaeontological record of vipers from the Mediterranean conclusion about the taxonomic identity of the predator agent islands is scarce, only 23 palaeontological sites are known on that produced the fossil association. The preliminary results eleven islands (Table 1; a complete review of the Mediterra- of the taphonomical analysis shows that in the studied mate- nean Islands fossil record of snakes is presented in Georgialis rial the digestion marks are mainly absent, with only a few et al. 2019b). Fossils with a morphology attributable to the specimens having light and moderate damage. The presence of Vipera aspis group are restricted to Western Mediterranean intact bones is rare. Discerning how this breakage has occurred islands, except for the remains of Vipera ammodytes recovered is complicated, therefore, and to avoid over-representing the from the Sanctuary of Poseidon at Kalaureia (Poros island, fragmentation degree in the analysis we do not study in detail Greece) (Lymberakis & Iliopoulos 2019). The oldest report the fragmentation of reptile’s bones. corresponds to the Vipera sp. fossils from the early Miocene The vertebrate assemblage of the COLT site is clearly of Oschiri (Sardinia) (Venczel & Sanchíz 2006). Also, Bover dominated by reptiles (81% of the remains), followed by et al. (2014) reported cf. Vipera natiensis from Na Burguesa-1 birds (35%). Mammals only represent 11% of the studied (Mallorca, Balearic Islands; late Miocene/early Pliocene). From material. Moreover, the diversity of species is low compared the Pliocene localities, the V. aspis group is cited as Vipera sp. in to mainland. This reduced biodiversity is typical of island in Capo Mannu D1 site in Sicily (Italy), MN15 (early Pliocene; contrast to continental areas (Grant 1988). On islands, reptiles Delfino et al. 2011) and as Vipera natiensis, an endemic taxon are often abundant and reach sometimes elevated densities in Punta Nati 12 site (Menorca, Balearic Islands, Spain, early (Pérez-Mellado et al. 2008), whereas terrestrial mammals are Pliocene) (Bailon et al. 2002). In Quaternary deposits, the infrequent (Lawlor 1986). COMPTES RENDUS PALEVOL • 2023 • 22 (18) 423 Marquina-Blasco R. et al. Following Andrews (1990) and Pinto Llona & Andrews Island (Balearic Islands), and the attacks recorded to lizard (1999), the digestion degree of the assemblage could cor- models can rather be attributed to the exploratory behaviour respond with a predator category 1. Within this category of the Yellow-legged Gull (Pérez-Mellado et al. 2014). On have been included Tyto alba (Scopoli, 1769) and Asio flam- the other hand, all the small mammal species present on the meus (Pontoppidan, 1763) (Andrews 1990). No breeding Colom Island were preyed by L. michahellis (Pérez-Mellado population of nocturnal bird of prey is present currently in et al. 2014). As generalist feeders, seagulls prey on snakes too; the Columbretes Islands. However, the existence of repro- however, there are few reports in this sense in the scientific ductive colonies of owls is possible during historical times. literature (i.e., Martín & López 1990). To our knowledge, no The Short-eared Owls (Asio flammeus) is a common winter- study has been carried out yet on the degree of taphonomic ing species in Spain, with the eastern peninsula as one of its alteration of prey bones caused by gulls. main Iberian wintering areas (Onrubia et al. 2003). Until the 1990s, breeding of A. flammeus in Iberia was occasional, Origin of the historical faunal community but the expansion of Microtus arvalis (Pallas, 1778) in the The origin of the vertebrate terrestrial fauna of Columbretes Northern Plateau, and its periodic population explosions, has been explained by three hypotheses: 1) isolated popula- have allowed the settlement of a variable number of nesting tions on the islands due to the sea-level rising (Castilla 2000; Short-eared Owls (Onrubia et al. 2003). As for the barn owl Ruiz-Sánchez et al. 2019); 2) colonization through marine (T. alba) it is a widespread species, with breeding popula- dispersion on floating rafts (Harris & Sá-Sousa 2002); or tions both on the mainland and on numerous islands in the 3) human introduction (Bernis 1968). The latter prob- Mediterranean and Macaronesia (Janžekovič & Klenovšek ably explains the origin of the terrestrial mammals, as it is 2020). Although reptiles are not an important part of the diet reflected in the historical references. Salvator (1895) was the of this species (Roulin & Dubey 2012), their consumption first author who reported the presence of mammals, point- are higher on islands, especially on the smaller ones, than ing out that they has recently appeared on the island, around on the continental areas (Janžekovič & Klenovšek 2020). In 1888, and went extirpated in an indeterminate moment of this sense, Delgado (1993) pointed out that reptiles (mainly the 20th century (Jiménez 1991). Thus, the mammal fossils the single species: Tarentola angustimentalis Steindachner, described here belong to an older colonization phase, previ- 1891) are the main trophic resource of the eastern canarian ous to the historical reports. The Columbretes Islands, due endemic subspecies Tyto alba gracilirostris (Hartert, 1905) in to their geographical position between Ibiza and the mouth the Alegranza Islets (Canary Islands); particularly in sum- of the Ebro river, are present in the Western Mediterranean mer time, when T. angustimentalis represents 69.7% of the commercial routes since the Antiquity. The seabed around consumed preys. A similar situation has been described for the Columbretes archipelago has preserved numerous ship- the T. alba populations of the Branco and Raso islets (Cabo wrecks since the early Roman Period. The oldest archaeo- Verde Islands) (Naurois 1982). Worth considering are several logical remains have been dated around the 3-4th centuries preliminary results in Macaronesia pointing to a shift in the BC (Fernández Izquierdo 2013), i.e., within an age range trophic spectrum of T. alba, from reptiles to small mammals, similar to the COLT radiocarbon datings. This material, a coinciding with the human colonisation of the islands, which B3-type Iberian amphora, was probably manufacturated by reflects the trophic flexibility of the species (Siverio et al. 2007, the Laietani (Fernández Izquierdo 2013), an ancient Iberian 2008). Another strategy reported from insular populations population who occupied the coast of the present Barcelona of T. alba is breeding on small islets (s’Espalmador, Balearic province between the Llobregat and Tordera rivers (Balsera Islands) and foraging (mainly small mammals, but also birds Moraño 1997). It is interesting to note that the closest Iberian and reptiles) on larger islands (Ibiza and Formentera) and people to the Columbretes Islands were not the Laietans but other islets surrounding s’Espalmador (Guerra et al. 2014). the Ilercavones, located in the area occupied by the current These owls fly at least 11 km on each occasion (s’Espalmador provinces of Castellón and Tarragona (Crespo et al. 2017). to Ibiza) (Guerra et al. 2014). Nevertheless, the sea stretch Since Mus musculus is a strongly human-dependant taxa that separates Illa Grossa from the continent is significantly (Palomo et al. 2007), its ocurrence could be treated as an wider (around 50 km). indicator of some kind of human presence or/and activity in Other potential biologic taphonomic agents are seagulls. the area, as for example voyaging (Vigne 2015). In 1976, Mus Breeding colonies of two taxa: Larus michahellis Naumann, musculus and Mus spretus were recognized by genetic criteria 1840, and Larus audouinii Payradeau, 1826 are located in the as different species (Britton et al. 1976; Britton & Thaler archipelago (Mestre et al. 2010). Experimental data based 1978). According to Domínguez García et al. (2019), the on the use of plasticine models of lizards demonstrated that genus Mus could have arrived to the southwestern European the Yellow-legged Gull (Larus michahellis) is the main preda- coasts around 6 000 BP or even before, but evidence is not tor of Podarcis liolepis atratus in Illa Grossa (Castilla & completely solid. In the case of Mus spretus, an Iberomagrebian Labra 1998). In the literature, there are some reports of species, a colonization of Europe linked to anthropogenic L. michahellis preying on lizards, especially in insular contexts translocation from North Africa to the Iberian Peninsula (Martín & López 1990; Carretero et al. 1993; Vervust et al. through the Strait of Gibraltar, probably during the Chal- 2007; Piorno et al. 2016). However, lizards do not seem to colithic, has been proposed (Domínguez García et al. 2019; be a common prey in L. michahellis diet, at least in the Colom Olalde et al. 2019). On the other hand, Mus musculus would 424 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island A 19 000-22 000 years BP B 14 000-15 000 years BP Oropesa del Mar Oropesa del Mar Benicàssim Benicàssim Columbretes Island Columbretes Island Mediterranean Sea Mediterranean Sea 10 km 10 km C 11 000-12 000 years BP D 9 000-10 000 years BP Oropesa del Mar Oropesa del Mar Benicàssim Benicàssim Columbretes Island Columbretes Island Mediterranean Sea Mediterranean Sea 10 km 10 km Present emerged land Sea Emerged coastral plains Fig. 7. — Palaeogeographical scenario of Columbretes Islands at: A, 19 000-22 000 years BP; B, 14 000-15 000 years BP; C, 11 000-12 000 years BP; D, 9 000- 11 000 years BP. The reconstruction is based in the model proposed by Lambeck & Bard (2000). have arrived with human migrations from South-East Asia, than 1.5 Myr ago. Given that the last connection between the following the “Mediterranean Route” sometime after 3 000 BP Balearic Islands and the European mainland occurred during (Auffray et al. 1990; Bonhomme & Searle 2012). According the Messinian Salinity Crisis (5.97-5.32 Mya) (Bover et al. to this chronology, the most plausible hypothesis would be 2014), the arrival of the species to Ibiza must have taken place that mice would have been introduced on the islands by the through floating rafts, probably from the northeastern Iberian Phoenicians, who established an intense commercial activ- region (Torres-Roig et al. 2020). Regarding the Columbretes ity since the 6th century BC in the nearby coast of the cur- vipers, Bernis (1968) suggested a possible anthropic intro- rent Castellón Province (Elias Ramos & Fernández 2012). duction or a very recent colonization of Illa Grossa. In fact, In agreement with this, Bisbal-Chinesta et al. (2020) have this authors did not identify any difference in the external pointed out that a recently discovered population of Chal- morphology between the specimen presumably found on the cides ocellatus in Serra del Molar (SE Spain), was probably island and the vipers of the eastern Iberian Peninsula. The introduced by Phoenicians. only characteristic noted by Bernis (1968) is the remarkable Vipera latastei is a species capable of transmarine dispersal, length of the examined specimen (72-73 cm), which makes as recently demonstrated by Torres-Roig et al. (2020). These it the largest specimen of Vipera latastei known to date (Brito authors, based on a phylogenetic analysis of an almost complete 2017). It is however dubious whether this is a case of island mitochondrial genome obtained from fossil material of Vipera gigantism due to the scarcity of material (only one specimen). latastei ebusitana (Late Pleistocene-Holocene, Es Pouàs, Ibiza, In addition snakes exhibit indeterminate growth throughout Balearic Islands, Spain), proposed that this taxon colonized their life (Andrews 1982), so this specimen could also be just the island of Ibiza crossing the seaway from the Peninsula less an old individual. However, as Boback (2003) pointed out, COMPTES RENDUS PALEVOL • 2023 • 22 (18) 425 Marquina-Blasco R. et al. viperids show a body size decrease trend in island popula- mainland, thus originating the insular populations of these tions. This author based its conclusions mainly in pitvipers of species. According to the model proposed by Lambeck & Bard the genus Crotalus, and only three of the 11 analyzed species (2000) for the sea-level change along the French Mediterra- belonged to the genus Vipera. Recently, this insular dwarfism nean coast for the Late Pleistocene and Holocene, between has been reported for the Vipera aspis population from the approximately 14 000 and 15 000 years ago, the Columbretes Montecristo Island (Luiselli et al. 2015) and for the Middle/ Islands were already separated from the mainland, at which Late Pleistocene-Holocene fossil remains of V. latastei ebusi- time the coastline was 70-75 m below the current sea level. tana from Ibiza (Balearic Islands) (Torres-Roig et al. 2020). At that time, the current different islands and islets could The fossil viper specimens recovered from the COLT site are be included in a single island (Fig. 7B). Around 11 000- also small-sized; however, their small number together with 12 000 years BP, the Illa Grossa group was isolated from the the large size of the specimen stored at the MNCN-CSIC rest of groups (Ferrera, Foradada and El Carallot), which prevent us from inferring the existence of an insular dwarfism were connected (Fig. 7C). Muñoz et al. (2005) reported the in the Illa Grossa population. existence of submarine volcanic structures without evidence About the possible anthropic origin for the V. latastei popu- of erosion, which could be originated at depths low enough to lation of Illa Grossa, Bernis (1968) hypothesised that some be eroded by waves, or even, as was proposed by Ancochea & specimens were released by a local chief to avoid the use of Huertas (2021), originated in subaerial context. The age of the island by pirates. The use of snakes as war weapons during these structures has been proposed as 13 000 BP based on the Antiquity is far knowledge (Mayor 2003). In this sense, different considerations regarding the speed of sea level rise the habit of launching earthenware jars filled with snakes (Muñoz et al. 2005). At 9 000-10 000 years ago the different against rival ships to spread chaos among the enemies was island groups were already isolated from each other (Fig. 7D). the proposed origin for the Montecristo Island Vipera aspis Finally, the present sea level was reached approximately at population (Bruno 1985). However, there are no registered 6 000 years ago (Lambeck & Bard 2000). naval battles in the vicinity of this island (as in the case of the The taxa identified in the fossil record of the COLT site Columbretes Islands). Genetic studies in the Montecristo Island have been recorded in localities from the central and northern demonstrated a more plausible origin for the snakes popula- Mediterranean Spanish coasts from the Piacenzian (MN17, tion around the middle of the first millennium AD, when the late Pliocene) up to the present time (Blain 2009). Therefore, island was inhabited by anchorites, either due to involuntary the composition and origin of those taxa can be tested. introduction or for medical purposes (Masseti & Zuffi 2011). Regarding wall lizards, Harris & Sá-Sousa (2002) proposed Due to the relatively low latitude range of the Iberian Penin- a possible colonization of the islands by rafting following sula (36-43° N), far from the European glacial zones, and the the southerly sea currents. In this sense, the genus Podarcis tempering effect of the proximity of the sea, the eastern and contains species with markedly different dispersion capabil- south-eastern coast of Spain only manifested subtle changes in ity across the sea. Thus, Podarcis peloponnesiacus (Bibron & temperature, during the Late Pleistocene-Holocene boundary Bory, 1833), Podarcis filfolensis (Bedriaga, 1876), Podarcis except in those area located at high altitude. Major changes cretensis (Wettstein, 1952) and Podarcis levendis Lymbera- seem to be related with eustatic oscillations as well as humid- kis, Poulalakis, Kaliontzopoulou, Valakos & Mylones, 2008 ity variations (Aura et al. 1998). Sea level oscillations have show limited over-sea dispersal capacity (Salvi et al. 2014; been associated mainly with exchanges between the ocean Spilani et al. 2019). On the other hand, species such as and the ice sheets during hot and cold periods (Lambeck & Podarcis vaucheri, seem to have spread from Morocco-West Bard 2000). A sea-level decrease of approximately 120 m has Algeria to the south of the Iberian Peninsula by crossing been documented during the Last Glacial Maximum (LGM) the Gibraltar Strait around 2.81 Mya (Pinho et al. 2006). (22 000-19 000 years before present) (Shackleton et al. 1984). Other species that could have dispersed over sea is Podar- Furthermore, according to Shackleton et al. (1984), the sedi- cis gaigeae (Werner, 1930), in which a gene flow has been ment layer produced during the Holocene must be thin, given reported between the subspecies Podarcis gaigeae weigandi the presence of few large rivers and the negligible effect of (Gruber & Schultze-Westrum, 1971), from the Piperi Island vertical tectonic movements throughout the Mediterranean and Podarcis gaigeae gaigeae (Gruber, 1986), which inhabits Sea basin. At present, the Columbretes Islands are located the Skyros Island (40 km north from Piperi) and satellite between the 75 and 100 m isobates (Ancochea & Huertas islets (Runemark et al. 2012). The capacity of transmarine 2021). Considering its bathymetric context, it is likely that dispersal in Podarcis liolepis in general and P. l. atratus in the Columbretes archipelago was part of the continent during particular, is unknown. The absence of this species in the the LGM (Fig. 7A). At that moment, Illa Grossa was located Ferrera group (Castilla et al. 1991, 1998a) might be due within the mainland area, 9.47 km from the contemporane- either to its weak oversea migration capacity or to its later ous coastline (the current mainland shoreline is situated at extirpation at an undetermined moment. The hypothesis 61.69 km from the Illa Grossa, Fig. 7A). According to this of colonization of the Illa Grossa Island via rafting is sup- hypothesis, Chalcides bedriagai, Vipera cf. latastei and cf. Podarcis ported by the close presence of a large river (Ebro), and the sp. reached Illa Grossa in an undetermined moment when it dominant southerly sea current in this area (Font 1990); was part of the continent. Later, with the gradual Holocene however, the small size of the island implies small colo- transgression, the locality was once again isolated from the nization rates. Since this process is totally random, small 426 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island islands receive fewer dispersion events than the bigger ones cies exploited on the archipelago, which comprise plants, (MacArthur & Wilson 1967). On the other hand, human arthropods (including marine isopods in the intermareal introduction (voluntary or not) cannot be discarded, at least zone), carcasses and congeneric specimens (Castilla & Bau- for the wall lizards. In fact, human-introduced populations wens 1991; Castilla et al. 1991, 2008, 2009; Castilla & van of Podarcis species have been detected in 222 Mediterranean Damme 1996). Moreover, “cross predation” between the islands in recent years (Silva-Rocha et al. 2019). Moreover, Wall lizard and the scorpion Buthus occitanus (Amoreux, the high leves of genetic divergence in the Podarcis liolepis 1789) has been reported; both species are abundant in the populations from the different islands of the Columbretes Illa Grossa Island (Castilla et al. 2009). In the recent past, archipelago (Castilla et al. 1998b), suggest several phases when the vertebrate community was more diverse, as shown of colonisation of the different islands by various genetic by the study of the COLT fossils, the interactions between lines of mainland P. liolepis or even by other Iberian Podarcis the different species might have been more complex. In rep- species (Harris & Sá-Sousa 2002). If this is confirmed, it is tile communities, several constraints, such as evolutionary not inconceivable that at least, some of these colonisation history, habitat use and intrinsic morphology, determine the waves were human-induced. Castilla et al. (1998b) pointed relationship between the different species (Capula & Luiselli out that the wall lizard population of the Foradada Group 1994, Carretero et al. 2010). Thus, it is expected that at least Islands (Foradada and Lobo Islands) possibly derived from one of those constraints was a conditioning factor in the pal- an early colonization wave. The same authors proposed that aeoherpetological community of the Columbretes islands. For the divergence between the wall lizards of these islands and example, the lizard microcommunity inhabiting the Lampione those from the Illa Grossa Island took place 80 000 years ago. islet (2.1 ha; Pelagian islands, South Italy) is composed by However, these results should be taken with caution since the skink Chalcides ocellatus (Forskål, 1775) and the lacertid they are based on a small sample of continental individuals Podarcis filfolensis (Bedriaga, 1886) (Carretero et al. 2010; Lo from the central Spanish Mediterranean region (only eight Cascio & Pasta 2012). Both species show a moderate trophic individuals from Valencia, Burjassot and Grau de Castelló). overlap, and while C. ocellatus consumes hard medium to large In summary, the origin of the fossil assemblage is unclear. prey, P. filfolensis prefers more diverse and smaller sized preys It is even probable that the different taxa recovered from than the former (Carretero et al. 2010). Plant consumption the COLT site colonised Illa Grossa through different ways, and cannibalism have been reported in both species (Carret- i.e., during the probable connection of the archipelago with ero et al. 2010). The exploitation of both trophic resources the mainland, by means of marine dispersal or via human- increases in insular lizards due to the scarcity of other sup- facilitated introduction. In this sense, the arrival of Mus plies and high population densities (Pérez-Mellado & Corti cf. musculus to the island was probably mediated by human 1993; Carretero 2004). However, there is a clear size differ- activity. Future genetic studies based on mainland and insular ence between C. ocellatus (TL: 30 cm) and P. filfolensis (TL: populations of Podarcis liolepis could shed light about this 25 cm) (Carretero et al. 2010; Speybroeck et al. 2016). In issue. The same could be carried out for the vipers, provided this case, the reported cases of predation of P. filfolensis by that the genetic material of the stored specimen of Vipera C. ocellatus (Carretero et al. 2010) could be explained by the latastei at the MNCN-CSIC is not degraded. The case of higher size of the latter (Cohen et al. 1993). However, Chal- Chalcides bedriagai is more problematic since the recovering cides bedriagai is smaller than the European Podarcis taxa, of ancient DNA from the fossils is unlikely due to the high except for the Podarcis hispanicus species complex, which is temperatures recorded on the island. similar in size (Speybroeck et al. 2016). Thus, the trophic relationships in Illa Grossa were presumably different than Palaeoecology those described for the Lampione islet between Chalcides and According to the palynological study of Expósito & Burjachs Podarcis. Chalcides bedriagai is considered an opportunistic (2007) the palaeoflora of Illa Grossa consisted of a shrubland and euryphagous predator (Seva & Escarré 1976; López- dominated by herbaceous plants and low bushes, not differ- Jurado et al. 1978). In an insular context, C. bedriagai preys ent from the current vegetation. The sample “Mostra 3” of on a wider spectrum of arthropods than in mainland locali- Expósito & Burjachs (2007), and recovered from the same ties (López-Jurado et al. 1978). Most of the preys consumed COLT deposit, is dominated by herbaceous species, mainly by C. bedriagai are typically ground-dwelling taxa or live in Poaceae Barnhart (45.5%) and Apiaceae Lindl. (27.3%); other the lower parts of the vegetation (Seva & Escarré 1976). The reported taxa are Artemisia L. (4.5%) and other ligulliflora adaptation to plant consumption requires some evolutionary (4.5%). On the other hand, arboreal pollen is constituted by time (Carretero 2004) and this may explain why it has not Pinus sp. (9.1%) and Quercus ilex-coccifera (9.1%). Thus, the been reported in the islands of Nova Tabarca (Seva & Escarré described landscape based on pollen analyses correspond to 1976; López-Jurado et al. 1978); where this species has been an open Mediterranean maquia (Expósito & Burjachs 2007), isolated for only a short time interval. On the other hand, which is in agreement with the vertebrate species identified the Mediterranean small lacertid Podarcis is considered as a in the present work. food generalist and an active forager (Arnold 1987). In par- The present herpetological community on Columbretes is ticular, as mentioned above, Podarcis liolepis atratus shows a composed by a single taxon, Podarcis liolepis atratus. Several wide range of food sources (Castilla & Bauwens 1991; Cas- works showed the varied trophic resources that this spe- tilla et al. 1991, 2008, 2009; Castilla & van Damme 1996). COMPTES RENDUS PALEVOL • 2023 • 22 (18) 427 Marquina-Blasco R. et al. Although, both species seem to feed on the same preys, they perspective (Barnosky et al. 2017), primarily in protected select different microhabitats. Thus, whereas C. bedriagai areas. Moreover, the palaeontological record is essential for forages under rocks and other elements on the ground (Hai- the theoretical body of any translocation or reintroduction ley et al. 1987), members of the Podarcis hispanicus species project (Faith 2012). complex, in which P. l. atratus is included, are active foragers Our results demonstrate the presence of vipers in Illa on a wide spectrum of substrates (rocks, tree trunks, human Grossa, providing a solid empirical basis for a hypothetical structures, etc.) (Desfilis et al. 1993). It is possible that the reintroduction project of the species on the Columbretes extirpation of C. bedriagai on Illa Grossa made it possible archipelago. The main cause of the extirpation of Vipera for P. l. atratus to exploit new trophic resources previously latastei on the island is clear: active human hunting. This factor unavailable to the species. has resulted in the extirpation on numerous populations of Vipera latastei (Vipera cf. latastei in the fossil record of vipers from different areas of Europe (see Brito et al. 2001). the COLT site) may have been the terrestrial top preda- This persecution is derived from the consideration of snakes tor in Illa Grossa. In fact, skinks and wall lizards are part as disgusting, harmful and feared animals (Brito et al. 2001). of the prey spectrum of the species (Santos et al. 2007a). Given the absence of a permanent human population, with However, the frequency of predation varies according to the the exception of the technical staff of the Nature Reserve, age classes of vipers and local faunal community. Predation this problem is currently totally absent in the Columbretes over skinks seems to be anecdotal, whereas wall lizards are Islands. Therefore, it could be interesting to carry out a study the most consumed reptiles (Santos et al. 2007a). There are on the possibility of reintroducing the species in Illa Grossa, variations in the diet according to the age of the specimen; assessing any potential impacts on the current community of in juveniles the consumption of reptiles and arthropods is vertebrates, especially on the endemic Podarcis liolepis atratus. higher than in adults, which prey mainly on rodents and After the evaluation of potential risks and benefits for the birds (Bea & Braña 1988; Santos et al. 2007a). In islands ecological system of the island, as indicated by the IUCN/ with low density or absence of small mammals, viperids SSC (2013) guidelines, a further point would be to discern seem to be specialized in feeding mainly on birds (Nilson which population is genetically the closest to the one that was et al. 1999; Shine et al. 2002; Luiselli et al. 2015). As in present in the archipelago. For this purpose, samples from the mainland, an ontogenetic shift in the selected prey has the specimen deposited in the MNCN-CSIC might be used; been recorded on islands. For example, only lizards have given that, in principle, the contrasting temperatures recorded been reported in the stomach content of road-killed young in dry areas, as Illa Grossa, provoked a high degradation of individuals of Macrovipera schweizeri, an insular species from the ancient DNA potentially present in the recovered fossils the western Cyclades Islands, whereas in adults specimens remains from COLT (Bollongino & Vigne 2008). The fact only bird feathers were detected (Nilson et al. 1999). Bothrops that it is the only known insular population of V. latastei in alcatraz, an insular viper species from the Alcatrazes Islands historical times together with the high number of locali- (35 km off the coast of southeastern Brazil), is considered a ties where this species has disappeared (Santos et al. 2007b) paedomorphic taxon based in, among other traits, its diet further supports its reintroduction. However, previous to composed of centipedes and lizards (Marques et al. 2002). this, an in-depth study of the fossil record of the species on As Bernis (1968) proposed the Illa Grossa vipers fed on this island and others in the archipelago together with the invertebrates during their juvenile stage, whereas as adults search and study of new potential sites are necessary. In this they fed on lizards and, during the migration epoch, on respect, Hernández-Pacheco & Asensio Amor (1966) report passerine birds. According to Castilla (2000), the observed the existence of loess deposits in other sites on Illa Grossa, difference in scalation, color-pattern and body dimensions and Boira & Carretero (1991) cite the existence of a poorly between the P. l. atratus of Illa Grossa those of the rest of developed formation at Ferrera. The same question, a possible the islands could be partly due to the intense selective pres- reintroduction of the taxon in Illa Grossa, arise for Chalcides sures exerted by vipers. Moreover, laboratory experiments bedriagai, although the causes of its extinction are unclear. have shown that individuals of P. l. atratus responded to the chemical signal of V. latastei, even though this snake became extirpated more than 100 years ago (van Damme & CONCLUSIONS Castilla 1996). The study of the fossil assemblage from the COLT site yielded Implications for the conservation an unexpected diverse community of vertebrates composed In the last decades, the application of palaeontological data by two lizards (Chalcides bedriagai and cf. Podarcis sp.), one to conservation biology has greatly increased (Lyman 2012). snake (Vipera cf. latastei), one rodent (Mus cf. musculus) and, This new field, known as Conservation Palaeobiology, has at least, six undetermined species of birds (two seabirds and numerous applications (Barnosky et al. 2017). Palaeontology four passerines). The historical presence of vipers in the Illa provides valuable information about which species were pre- Grossa Island is confirmed, thus settling the controversy about sent in the past in a determined region, especially in disturbed the identity of the snakes of the Columbretes archipelago. For habitats (Faith 2012). The knowledge of a baseline biological the first time, the historical presence of Chalcides bedriagai is community is strongly interesting for the land-management reported in the archipelago. 428 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island Two different samples: COLT-SNAILS (fragments of ter- Pavia M., Piras P., Rook L., Torre D., Tuveri C., Valli A. & restrial gastropods) and COLT-BONES (fragments of unde- Wilkens B. 2004. — Plio-Pleistocene fossil vertebrates of Monte Tuttavista (Orosei, Eastern Sardinia, Italy), an overview. Rivista termined bones) were AMS radiocarbon-dated. Both yielded Italiana di Paleontologia e Stratigrafia 110: 681-706. strongly similar chronologies: COLT-SNAILS has been dated Alcover J. A. 2000. — Vertebrate Evolution and Extinction on at 2780-2724 cal BP and COLT-BONES at 2600-2492 cal BP. Western and Central Mediterranean Islands. 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Submitted on 22 June 2021; accepted on 25 May 2022; published on 8 June 2023. 436 COMPTES RENDUS PALEVOL • 2023 • 22 (18) Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island APPENDIX Appendix 1. — Measurements (in mm) and biometric indices of the trunk vertebrae of Vipera cf. latastei Boscá, 1878 (COLT), mainland Vipera latastei (IMEDEA 106925 and MNHN-ZA-AC-2020-1), and Vipera latastei ebusitana Torres-Roig, Alcover & Bailon, 2020. Data of mainland V. latastei and V. l. ebusitana obtained from Torres-Roig et al. (2020). Abbreviations: σ, standard deviation; CL, vertebral centrum length, measured ventrally and including the condyle; CL/NAW, ratio between the length and the width of the neural arch; CTH, cotyle height; CTW, cotyle width; CTW/CTH, ratio between width and height of the cotyle; CTW/ NAW, ratio between cotyle width and the width of neural arch; Max, maximum value; Min, minimum value; N, number of specimens; NAW, width of the neural arch, measured at the maximum interzygapophyseal constriction; PO-PO, width of the external borders of the articular surfaces of the postzygapophysis; PO- PO/NAW, ratio between the width of the external borders of the articular surfaces of the postzygapophysis and the width of the neural arch; PR-PR, width of the external borders of the articular surfaces of the prezygapophysis; PR-PR/NAW, ratio between the width of the external borders of the articular surfaces of the prezygapophysis and the width of the neural arch; ZW, zygosphene width; ZW/NAW, ratio between the width of the zygosphene and the width of the neural arch. COLT IMEDEA 106925 MNHN-ZA-AC-2020-1 Vipera latastei ebusitana CL Mean 2.76 3.74 4.31 2.74 N 5.00 – – 18 σ 0.43 – – 0.27 Min 2.10 – – 2.29 Max 3.21 – – 3.19 NAW Media 1.81 2.53 2.87 1.96 N 4.00 – – 18 desv 0.29 – – 0.24 Min 1.41 – – 1.63 Max 2.08 – – 2.4 CTW Media 1.00 1.84 2.22 1.22 N 4.00 – – 18 desv 0.55 – – 0.08 Min 0.19 – – 1.07 Max 1.35 – – 1.33 CTH Media 1.17 1.41 1.56 1 N 4.00 – – 18 desv 0.09 – – 0.09 Min 1.11 – – 0.85 Max 1.30 – – 1.13 PR-PR Media 3.22 5.45 5.82 3.62 N 3.00 – – 18 desv 0.50 – – 0.46 Min 2.70 – – 2.97 Max 3.70 – – 4.39 PO-PO Media 3.11 5.27 5.57 3.5 N 2.00 – – 15 desv 0.73 – – 0.45 Min 2.59 – – 2.97 Max 3.62 – – 4.3 ZW Media 1.16 2.18 2.80 1.74 N 1.00 – – 16.00 desv – – – 0.15 Min – – – 1.48 Max – – – 1.98 CTW/CTH Media 0.84 1.30 1.42 1.23 N 4.00 – – 18 desv 0.45 – – 0.06 Min 0.17 – – 1.14 Max 1.16 – – 1.35 PO-PO/NAW Media 1.85 2.08 1.94 1.8 N 2.00 – – 15 desv 0.02 – – 0.07 Min 1.84 – – 1.65 Max 1.87 – – 1.9 ZW/NAW Media 0.82 0.86 0.98 0.9 N 1.00 – – 16 desv – – – 0.05 Min – – – 0.8 Max – – – 0.98 CTW/NAW Media 0.47 0.73 0.77 0.63 N 3.00 – – 18 desv 0.33 – – 0.05 Min 0.09 – – 0.54 Max 0.68 – – 0.73 COMPTES RENDUS PALEVOL • 2023 • 22 (18) 437 Marquina-Blasco R. et al. Appendix 1. — Continuation. COLT IMEDEA 106925 MNHN-ZA-AC-2020-1 Vipera latastei ebusitana CL/ZW Media 1.81 1.72 1.54 1.56 N 1.00 – – 16 desv – – – 0.08 Min – – – 1.44 Max – – – 1.69 PR-PR/NAW Media 1.88 2.15 2.03 1.84 N 3.00 – – 18 desv 0.05 – – 0.06 Min 1.83 – – 1.67 Max 1.91 – – 1.94 CL/NAW Media 1.47 1.48 1.50 1.4 N 4.00 – – 18 desv 0.02 – – 0.08 Min 1.45 – – 1.24 Max 1.49 – – 1.55 438 COMPTES RENDUS PALEVOL • 2023 • 22 (18)