identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
FF1AFA1CFFE2BF4CFF55FA5BFBA7992F.text	FF1AFA1CFFE2BF4CFF55FA5BFBA7992F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lacerta Linnaeus 1758	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Lacerta Linnaeus, 1758 and thus all species currently referred to  Timon were initially proposed as species of  Lacerta . Only in 1996, was  Timon recognized as a distinct genus (Mayer &amp; Bischoff, 1996), a conclusion that has since been widely accepted (Fu, Murphy &amp; Darevsky, 1997; Harris, Arnold &amp; Thomas, 1998; Fu, 1998, 2000; Harris &amp; Carretero, 2003; Arnold, Arribas &amp; Carranza, 2007; Schmidtler, 2010; Pyron, Burbrink &amp; Wiens, 2013; Ahmadzadeh et al., 2016). </p>
            <p> Six species are currently attributed to  Timon :  T. kurdistanicus (Suchov, 1936) ,  T. lepidus (Daudin, 1802) ,  T. nevadensis (Buchholz, 1963) ,  T. pater (Lataste, 1880) ,  T. princeps (Blanford, 1874) and  T. tangitanus (Boulenger, 1881) (Ahmadzadeh et al., 2012, 2016; Miraldo et al., 2013). Molecular and morphological phylogenetic analyses show that  Timon is the sistergenus to  Lacerta (e.g. Carranza, Arnold &amp; Amat, 2004; Arnold et al., 2007; Kapli et al., 2011; Pyron et al., 2013; Sagonas et al., 2014; Mendes et al., 2016), and that it includes three distinct clades, which are also geographically segregated:  T. lepidus and  T. nevadensis occur on the European side of the western Mediterranean basin;  T. pater and  T. tangitanus inhabit the North-African coast of the western Mediterranean basin; and  T. kurdistanicus and  T. princeps are distributed in the Middle East, with no connection to the Mediterranean (Ahmadzadeh et al., 2016). The divergence of the two distinct eastern and western genetic lineages of  Timon , and the split between the European and African subclades, were estimated at 14.46 Mya and approximately 7.42 Mya, respectively, whereas the evolutionary divergence between  Lacerta and  Timon dates back to approximately 18.6 Mya (Ahmadzadeh et al., 2016). </p>
            <p>*Corresponding author. E-mail: etschopp@amnh.org</p>
            <p> The vast majority of fossil remains reported in the literature were recovered from geographical areas where  Timon still exists today (Mateo, 2009). FosFARbase lists 34 fossil occurrences referable to  Timon , from the Gelasian to the Holocene, and from France, Spain, Portugal and Gibraltar (Böhme &amp; Ilg, 2003). Some additional occurrences are reported by other authors, still from Quaternary localities of France and Spain (e.g. Fernández Eraso et al., 2010; Bañuls Cardona et al., 2012; Benítez de Lugo Enrich et al., 2015). The oldest fossil might be represented by a dentary from the Pliocene of France, which constitutes the holotype of  Lacerta ruscinensis Depéret, 1890 , but was considered closely related to  Timon lepidus by Depéret (1890) himself, Młynarski (1956) and Estes (1983). Estes (1983) even suggested that the specimen might be referred to the extant species, resulting in  L. ruscinensis being a junior synonym of  T. lepidus . The absence of records from Northern Africa and the Middle East is most probably due to the very scarce knowledge of the palaeoherpetofaunas from those regions. </p>
            <p> Reports of fossils outside the current distribution are rare, and none of them has been described or properly identified. A possible  Timon specimen from the late Pleistocene of Germany (Brunner, 1957) was referred to  Lacerta sp. by Estes (1983), and to  L. agilis by Mateo (1988). Large-sized, lacertid bones from Gargano (Italy) have been referred to  Lacerta sp. by Delfino &amp; Bailon (2000), who also mention similarities with  Timon lepidus . Bailon (2004) figured and described a mandible from the Middle Pleistocene of Corsica with affinities to  Timon lepidus . Both the occurrences from Gargano and Corsica have been included in a distribution map of the genus  Timon by Ahmadzadeh et al. (2016), which indicates additional fossil occurrences in Sicily (Italy) and on Malta. These fossils from Sicily and Malta most probably represent the findings of the extinct ‘  Lacerta ’ siculimelitensis by Böhme &amp; Zammit-Maempel (1982), although this is not explicitly stated. This enigmatic large-sized lacertid was considered to belong to  Timon by Mateo (2009), but a detailed reassessment of the species would be needed to confirm this interpretation. </p>
            <p> In order to improve knowledge of the insular lizards that could be related to  Timon , herein we describe and analyse the relationships of fossil material from the Calabrian to Upper Pleistocene fissure fillings of Monte Tuttavista (Orosei, Sardinia, Italy), which was initially referred to  Lacerta sp. on the basis of large size (Abbazzi et al., 2004), but whose morphology was not described in detail. </p>
            <p>INSTITUTIONAL ABBREVIATIONS</p>
            <p> The material from Monte Tuttavista was initially numbered using ‘MT’ as abbreviation for the locality, ‘VIa’ for specimens from the site ‘Cava VI-antica’, ‘IX’ for the site ‘Cava IX-  Prolagus ’ and ‘BS’ for the site ‘Cava VIII-Blocco Strada’. This resulted in specimen numbers such as ‘MT-IX-054’. All these specimens are deposited in the Soprintendenza Archeologia, Belle Arti e Paesaggio per le province di Sassari e Nuoro, Nuoro, Italy. </p>
            <p> Other institutional abbreviations used herein are the following:  CIPA, Osteoteca, LaboratorioArqueociencias, Lisbon, Portugal ;  COMGR, Collezione Osteologica Mauro Grano, Roma, Italy ;  HUJ-OST, Osteological Collections, Hebrew University of Jerusalem, Israel ;   MDHC, Massimo Delfino Herpetological Collection in the  Museum of Geology and Paleontology of the Department of Earth Sciences of the University of Turin, Italy  ;   MNCN,  Museo Nacional de Ciencias Naturales , Madrid, Spain  ;  MNHN, Muséum National d’Histoire Naturelle, Paris, France ;   MRAC,  Musée Royal de l’Afrique Centrale , Tervuren, Belgium  ;  NHMUK, Natural History Museum, London, UK ;  NHMW, Naturhistorisches Museum Wien, Vienna, Austria ;  PIMUZ, Paläontologisches Institut und Museum der Universität Zürich, Switzerland ;  SRK, Sammlung Ralf Kosma, Staatliches Naturhistorisches Museum Braunschweig, Germany ;  UAM, Universitad Autónoma de Madrid, Spain ;   ZZSiD,  Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland  . </p>
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	https://treatment.plazi.org/id/FF1AFA1CFFE2BF4CFF55FA5BFBA7992F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Tschopp, Emanuel;Villa, Andrea;Camaiti, Marco;Ferro, Letizia;Tuveri, Caterinella;Rook, Lorenzo;Arca, Marisa;Delfino, Massimo	Tschopp, Emanuel, Villa, Andrea, Camaiti, Marco, Ferro, Letizia, Tuveri, Caterinella, Rook, Lorenzo, Arca, Marisa, Delfino, Massimo (2018): The first fossils of Timon (Squamata: Lacertinae) from Sardinia (Italy) and potential causes for its local extinction in the Pleistocene. Zoological Journal of the Linnean Society 184 (3): 825-856, DOI: 10.1093/zoolinnean/zly003, URL: https://academic.oup.com/zoolinnean/article/184/3/825/4953635
FF1AFA1CFFF8BF56FEC8FBE0FD499A19.text	FF1AFA1CFFF8BF56FEC8FBE0FD499A19.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Timon	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> PALEOBIOGEOGRAPHY OF  TIMON</p>
            <p> The herein reported fossils represent the first occurrence of  Timon in Sardinia, which is outside the current geographical range of the genus, and lies in between the eastern and western areas inhabited by the three distinct species-groups of  Timon (Ahmadzadeh et al., 2016) . However, this is perhaps not so surprising given that, according to distribution models based on current environmental factors, all four western species of  Timon would be adapted to inhabit Sardinia (Ahmadzadeh et al., 2016). The occurrence of  Timon in the Pleistocene of Sardinia thus further corroborates a wider distribution of the genus in the recent past, as proposed by Ahmadzadeh et al., (2016). </p>
            <p>POTENTIAL CAUSES FOR LOCAL EXTINCTION</p>
            <p> Timon is not the only reptile known from Monte Tuttavista that became extinct in Sardinia. The fossil record also documents the presence of agamid lizards, vipers and amphisbaenians (Abbazzi et al., 2004; Delfino et al., 2008; Delfino, Bailon &amp; Pitruzzella, 2011), which are absent on the island nowadays. Other localities from the Miocene and Pliocene testify for the former presence of crocodiles, trionychid turtles, non-  Anguis anguid lizards, aniliid and erycine snakes (Kotsakis, Delfino &amp; Piras, 2004; Delfino et al., 2011; Georgalis et al., 2017). </p>
            <p> One of the reasons for local extinction could be the arrival or evolution of new predators. A causal relationship with the first arrival of humans was initially proposed as potential cause for an apparent major faunal turnover during the Pleistocene (Sondaar et al., 1986). Two faunal complexes are recognized based on distinct compositions of the mammalian fauna (the  Nesogoral and  Microtus complexes), with two subcomplexes each (Abbazzi et al., 2004; Palombo, 2006, 2009; Palombo &amp; Rozzi, 2014; Vigne, 2014). The findings at Monte Tuttavista confirm the presence of such a turnover (Abbazzi et al., 2004) but more recent data indicate that this turnover has not been as severe as previously thought (Palombo et al., 2017). Also, it currently seems rather improbable that humans really established sustained settlements on the island before the Holocene, which would have been necessary to have such an important impact on the local fauna (Palombo, 2006, 2009; Masini et al., 2008; Leppard, 2014; Palombo et al., 2017). In any case, at Monte Tuttavista,  Vipera was only found from sites attributed to the  Nesogoral complex, whereas  Timon and the agamids occur in both faunal complexes, and only seem to disappear at the beginning of the so-called ‘Dragonara’ subcomplex within the  Microtus complex (Abbazzi et al., 2004; Palombo, 2006, 2009; Delfino et al., 2008; Palombo &amp; Rozzi, 2014). The extinction of these large lizards does, therefore, not correlate with the main faunal turnover, but rather with the change from the so-called ‘Orosei 2 subcomplex’ to the ‘Dragonara’ subcomplex, confirming a rather gradual faunal change as proposed by Palombo et al. (2017). </p>
            <p> Whereas an influence of humans on the local extinction of  Timon and agamids appears unlikely, several carnivore mammals first occur on Sardinia during the ‘Dragonara’ subcomplex. These include three members of Lutrinae, and the small-sized canid  Cynotherium sardous (Abbazzi et al., 2005; Palombo, 2006, 2009; Palombo &amp; Rozzi, 2014). The lutrines are specialized to prey on aquatic animals (Willemsen, 1992), and can thus probably be excluded as predators of the large lizards. However, the appearance of the endemic canid  Cynotherium sardous is of particular interest with respect to the extinction of  Timon and possibly also the vipers and agamids.  Cynotherium sardous likely evolved from larger canids that first arrived on the island during the Early Pleistocene (just before or during the main faunal turnover: Abbazzi et al., 2005; Lyras &amp; van der Geer, 2006; Lyras, van der Geer &amp; Rook, 2010; Palombo &amp; Rozzi, 2014). Throughout the Pleistocene, these earlier forms became progressively smaller and adapted to prey on small-sized animals like the small lagomorph  Prolagus (Lyras &amp; van der Geer, 2006; Lyras et al., 2010), and thus possibly also large-sized lizards such as  Timon , whereas smaller lizards remained safe from predation by this newly evolved predator (see below). Larger canids, as well as hyaenids, are reported from the early Pleistocene of Sardinia (Rook et al., 2004; Abbazzi et al., 2005; Lyras &amp; van der Geer, 2006), but these were most likely less adapted to feed on smaller prey like  Prolagus , large lizards and snakes, and became extinct during the faunal turnover (Rook et al., 2004; Palombo, 2006, 2009; Palombo &amp; Rozzi, 2014). </p>
            <p> The body size decrease in  Cynotherium , and thus the evolution of  C. sardous , can also be traced among the sites at Monte Tuttavista (Abbazzi et al., 2005). Whereas elements attributable to the genus are found nearly throughout all stratigraphic layers, only bones from the two stratigraphically highest sites, ‘VI-Banco 6’ and ‘VII-2’, can be tentatively (  C. cf. sardous at VI-Banco 6; Abbazzi et al., 2005) and definitively referred to the small-sized  C. sardous (VII-2; Abbazzi et al., 2005). This size decrease apparently correlates, in particular, with the disappearance of  Timon and agamids: VI-Banco 6 produced six remains attributable to  Timon , and VII-2 none (see Fig. 1 and Supplementary Material), and the last occurrence of agamids is from ‘XI-canide’, which is just below the two sites hosting  C. sardous . Indeterminate snakes and the small-sized, possible  Podarcis continued to thrive throughout the ‘Dragonara’ subcomplex, and were among the most frequent finds in VI-Banco 6 (Abbazzi et al., 2004). Around the Pleistocene–Holocene boundary,  Vulpes vulpes also arrived on Sardinia (Palombo, 2006), which has been shown to prey on lizards and snakes (Díaz-Ruiz et al., 2013). </p>
            <p> The Pleistocene was a time of marked climatic oscillations between warmer and colder periods. The end of the Pleistocene corresponds to a cold period (the Younger Dryas; Miraldo et al., 2013), which seems to have had an important impact on the fauna in Corsica (Vigne, 2014) and on the distribution of  Timon lepidus on the Iberian Peninsula (Miraldo et al., 2013). However, current  Timon populations inhabit variable climatic zones around the Mediterranean, and some species seem to have speciated quite fast to adapt to new environmental conditions and ecological niches (Ahmadzadeh et al., 2016). Therefore, it seems most plausible that the cause for extinction of large lizards on Sardinia was a combination of the evolution or introduction of new predators preying on small-sized animals (in particular  Cynotherium sardous ) and climatic changes. </p>
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	https://treatment.plazi.org/id/FF1AFA1CFFF8BF56FEC8FBE0FD499A19	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Tschopp, Emanuel;Villa, Andrea;Camaiti, Marco;Ferro, Letizia;Tuveri, Caterinella;Rook, Lorenzo;Arca, Marisa;Delfino, Massimo	Tschopp, Emanuel, Villa, Andrea, Camaiti, Marco, Ferro, Letizia, Tuveri, Caterinella, Rook, Lorenzo, Arca, Marisa, Delfino, Massimo (2018): The first fossils of Timon (Squamata: Lacertinae) from Sardinia (Italy) and potential causes for its local extinction in the Pleistocene. Zoological Journal of the Linnean Society 184 (3): 825-856, DOI: 10.1093/zoolinnean/zly003, URL: https://academic.oup.com/zoolinnean/article/184/3/825/4953635
