Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 127(2): 297-354. July 2021 An overview of the lAte Miocene vertebrAtes froM the fissure fillings of Monticino QuArry (brisighellA, itAly), with new dAtA on non-MAMMAliAn tAxA AndreA VillA1*, GiorGio CArneVAle2, MArCo PAViA2, lorenzo rook3, MArCo SAMi4, zbiGniew SzyndlAr5 & MASSiMo delfino6* 1* Co-corresponding author. Staatlichen naturwissenschaftlichen Sammlungen bayerns - bayerische Staatssammlung für Paläontologie und Geologie, richard-wagner-Straße 10, 80333 München, Germany and dipartimento di Scienze della Terra, Università degli Studi di Torino, Via T. Valperga Caluso 35, 10125 Torino, italy. e-mail: a.villa@unito.it 2 dipartimento di Scienze della Terra, Università degli Studi di Torino, Via T. Valperga Caluso 35, 10125 Torino, italy. e-mail: giorgio.carnevale@unito.it; marco.pavia@unito.it 3 dipartimento di Scienze della Terra, Paleo[fab]lab, Università degli Studi di firenze, Via la Pira 4, 50121 firenze, italy. E-mail: lorenzo.rook@unifi.it 4 Museo Civico di Scienze naturali “Malmerendi”, via Medaglie d’oro 51, 40018 faenza, italy. e-mail: marco.sami@cheapnet.it 5 Polish Academy of Sciences, institute of Systematics and evolution of Animals, Slawkowska 17, 31-016 krakow, Poland. e-mail: szyndlar@isez.pan.krakow.pl 6* Co-corresponding author. dipartimento di Scienze della Terra, Università degli Studi di Torino, Via T. Valperga Caluso 35, 10125 Torino, italy and Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA/ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain. E-mail: massimo.delfino@unito.it To cite this article: Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. (2021) - An overview of the late Miocene vertebrates from the fissure fillings of Monticino Quarry (Brisighella, Italy), with new data on non-mammalian taxa. Riv. It. Paleontol. Strat., 127(2): 297-354. Keywords: fish; Amphibians; reptiles; birds; Messinian. Abstract. The latest Messinian Monticino Quarry fissure fillings, near Brisighella in Italy, are well known for their diverse mammal fauna. Conversely, little is known about other vertebrates from this rich site. beside presenting an overview of the mammals, here we describe fish, amphibian, reptile, and bird remains, identifying three, eight, at least 18, and five taxa, respectively. Some of these represent the oldest occurrences either worldwide (an erycine snake related to Eryx jaculus, Malpolon, and otidids birds) or locally (common toads in the Apennine Peninsula and eremiadine lacertids in Europe, the latter being also the only known occurrence of these lizards in Italy). Altogether, the vertebrate fauna from Monticino Quarry is indicative of a complex palaeoenvironment, which was warm and somehow dry. water bodies were indeed present though, as well as a patchy set of habitats with loose or sandy soils, rocky outcrops, open biotopes, and (even though maybe to a lesser extent) wooded areas. Furthermore, estuarine or lagoonal conditions were likely present near the depositional site. From a palaeobiogeographical point of view, the non-mammal vertebrate assemblage from Monticino Quar- ry shows considerable Afro-Asian affinities at least for some components. In the context of the Italian late Miocene continental vertebrate assemblages, that of Monticino is remarkably similar to those from the Piedmont basin, espe- cially if compared with the Tusco-Sardinian area, Apulian Platform, Calabrian Arc and Sicily. This is consistent with the palaeobiogeographic scenario defined by mammal assemblages, with northcentral Italy, Tuscany and Sardinia, Apulian Platform, and Calabria and Sicily representing separated bioprovinces. IntroductIon ian gypsum in a quarry near brisighella in emilia Romagna, Central Italy (the so called Monticino Costa et al. (1986) first reported the occur- gypsum Quarry; Fig. 1). The strict local and region- rence of a latest Miocene (Messinian) fossil verte- al geological constraints (Marabini & Vai 1989; Vai brate fauna from karst fillings affecting the Messin- 1989) allowed a correlation of the vertebrate assem- blage with the latest Messinian and with the euro- Received: June 16, 2020; accepted: March 12, 2021 pean land Mammal Age Mn 13. 298 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Fig. 1 - Location and geological context of the Monticino Quarry - a) Map of Italy, showing the position of Brisighella, where the Monticino Quarry is located. b) Schematic geological section of the Monticino gypsum quarry. C) Formal geological units and chronological setting of the sediments exposed in the quarry. Abbreviations: BRS, Brisighella; ELMA, European Land Mammal Ages; iM tp, in- tra-Messinian tectonic phase; St., stage. Scale bar in a = 200 km. Modified after Rook & Bernor (2013). The study of the abundant vertebrate fauna turali di faenza and those of the dipartimento di from the fissure fillings exposed in the Montici- Scienze della Terra of the Università degli Studi di no Quarry has been mostly focused on mammals Firenze that were recently expanded thanks to the (De Giuli 1989; Ferretti 2008; Kotsakis & Masini screen washing and selection of unprepared fossil- 1989; Masini & Thomas 1989; Torre 1989; rook et iferous sediments collected in the 1980s. al. 1991; rook 1992; Masini & rook 1993; rook & Masini 1994; Gallai & rook 2011; rook & bernor GeoloGIcal settInG 2013; Pandolfi & Rook, 2017). No detailed infor- mation on the other vertebrates is available, except The exposed succession within the Montici- for a preliminary note on the amphibians and rep- no gypsum Quarry consists of middle Messinian tiles that was published by Kotsakis (1989). In this evaporitic (gypsum) succession (the so-called Ges- note, devoid of figures and descriptions and not soso-Solfifera Formation). Due to emersion and reporting collections numbers, the following taxa tectonization (tilting), the gypsum has been erod- were listed: Rana sp., Trionyx sp., emydidae indet., ed and affected by karstic dissolution. The deposits (?) Testudinidae indet., Gekkonidae indet., Lacerta that infill the fissures intersecting the gypsum and (s.l.) sp., Anguis sp., Ophisaurus cf. pannonicus, Varani- the depressions on the upper surface of the gyp- dae indet., cf. Eryx sp., Colubrinae indet., and natri- sum sequence consist of marly-clays, fine grained cinae indet. conglomerates and sand lenses belonging to the Later on, Delfino (2002) included some of latest Messinian Colombacci formation. They con- the amphibian and reptile remains from Monticino tain partially articulated and fragmented vertebrate Quarry in his PhD thesis and based on these iden- fossils (mostly microvertebrate remains). Several tifications, the amphibians and reptiles were men- fossiliferous collecting points have been identi- tioned and partly figured in a popular scientific pa- fied along the Monticino gypsum Quarry outcrop. per (Rook & Delfino 2003) and two book chapters The succession is capped by up to 20 m of early published in Italian (Rook & Delfino 2007; Rook et Pliocene marine gray marly clays (Argille Azzurre al. 2015). Formation), the base of which falls in the Sphaeroi- The present contribution to the knowledge dinellopsis zone, and which overlie the Colombacci of the vertebrates from the karst fissures of Mon- formation. dismissed in the early 1990s, the Mon- ticino Quarry is focused on the thorough descrip- ticino Quarry was then subject to environmental re- tion of the whole non-mammal vertebrate fauna covery and, because of its geological and palaeonto- but provides also an overview on the mammal taxa logical values, in 2006 it was converted into an open described so far. Therefore, besides focusing for the geological museum and included in the regional first time on fishes and birds, it deals with all the park “Parco regionale della Vena del Gesso” (Sami amphibian and reptile remains currently available in 2007). We refer the reader to Vai (1989) for a more the collections of the Museo Civico di Scienze na- complete discussion of the geological setting of the Late Miocene vertebrates from the fissure fillings of Monticino Quarry 299 The relative scarcity of large animals makes the Monticino faunal assemblage not fully repre- sentative of the mammal communities of the time. For example, among herbivores the proboscideans are represented by only two fragments, just as very few remains represents rhinos, equids, and large bovids. one of the factors responsible for this un- derrepresentation of large animals can be identified in the “size” of the tectonic-karst fissure network, characterized in almost all cases by small fissures/ cracks. In some cases (especially in the largest fis- sures or in the doline-like morphologies on the top of the gypsum succession), the occurrence of skeletal portions in anatomical articulation (Fig. fig. 2 - Partially articulated skull of Plioviverrops faventinus from Mon- 2) suggests that some karst cavities may also have ticino Quarry. functioned as either natural “traps” or as dens in which the animals remained trapped. This hypoth- Romagna Apennine and to Marabini & Vai (1989) esis (again, proposed by e.g., Rook et al., 2015) is for further details on the geology of the Monticino also confirmed by the unusual overabundance of gypsum Quarry. carnivorans remains in the same cracks. An alternative hypothesis to explain the taphonomy and the accumulation of the macrofau- taphonomy nal remains inside the fissures affecting the gypsum has been suggested by some authors (Costa et al. In the Monticino Quarry, most of the de- 1986), according to which the filling of the cracks/ posits yielding fossil vertebrate remains were found fissures studied would have occurred as conse- inside the cavities of a complex karst network de- quence of the phenomenon known as “neptunian veloped on the evaporites (gypsum) of the Gesso- dykes”: the deposits of the Colombacci formation so-Solfifera Formation and filled with lithotypes of would have been injected downwards following the the overlying Colombacci formation. Small ver- opening of cracks in the gypsum due to a tectonic tebrate remains constitute the vast majority of the phase that occurred after the deposition of fossil faunal assemblage, by far dominated by murids. in sediments at the base of the same unit. some cases, the fossiliferous sediments are extreme- ly rich, due to a very high concentration of bones and teeth. This peculiarity (not unusual in the case materIal & methods of karst fissure fillings) could be at least in part at- As reported above, fossils of non-mammalian vertebra- tributed to the accumulation of pellets produced by tes from Monticino Quarry are stored either at the Dipartimento birds of prey that used perches placed within the di Scienze della Terra, Università degli Studi di Firenze (Florence, karst area. Such an interpretation (proposed by e.g., Italy) or at the Museo Civico di Scienze Naturali di Faenza (Faen- Rook et al. 2015) may be in agreement with the fact za, Italy). They were identified using data from the literature and by direct comparisons with prepared skeletons of extant taxa (Marco that no substantial differences were observed in the Pavia Ornithological Collection MPOC and Massimo Delfino Her- presence of both left and right molars and in the petological Collection MDHC, both housed in the Dipartimento di number of first and second molars; moreover, some Scienze della Terra, Università degli Studi di Torino for birds and the herpetofauna respectively). Online resources from the digital repo- bone findings showed a particular type of wear sitory MorphoSource (https://www.morphosource.org/) were also comparable to that produced by the gastric juices of consulted for some taxa. Consulted specimens from MorphoSource the stomachs of some predatory birds (De Giuli et are: Alytes obstetricans cas:sua:21691; Pipa parva uf:herp:37924; Xeno- al. 1988). Fossil remains of medium-large sized ver- pus borealis cas:herp:152778; Xenopus clivii uf:herp:92074; Rhinophrynus dorsalis cas:herp:71766; Barbourula busuangensis uf:herp:70546; Bombina tebrates are mainly present as skeletal portions that maxima uf:herp:96648; and Pelobates varaldii cas:herp:92371. Pictures have been transported and, in any case, in a much of selected specimens were taken with a leica M205 microscope lower percentage in comparison to the microfauna. equipped with the leica application suite v4.10 at the Università degli 300 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Fig. 3 - Fish from Monticino Quarry - Aphanius crassicaudus: abdom- inal vertebra (BRS2/F/1) in anterior (a) and right lateral (b) views. ?Cichlidae indet.: isolated tooth (BRS19/F/1) in lateral view (c). ?Sciaeni- dae indet.: abdominal ver- tebra (BRS19/F/2) in right lateral view (d). Scale bars = 0.5 mm (a, b); 1 mm (c); 5 mm (d). Studi di Torino and with a Canon 5dii 100 mm and 50 mm macro with those of the cyprinodontid Aphanius crassicau- lenses. drawings were made with a camera lucida. The anatomical terminology follows Ratnikov & Litvinchuk (2007) for urodeles, San- dus (see Gaudant, 1979), a species very common in chiz (1998) for anurans, Zangerl (1969) for turtles, Villa & Delfino the Messinian paralic and coastal biotopes through- (2019a) for lizards, Szyndlar (1984) for snakes, and Baumel & Witmer out the Mediterranean (see Carnevale et al. 2006b, (1993) for birds. Snake taxonomy in this paper follows the recent 2018, 2019; reichenbacher & kowalke 2009; Co- work by Zaher et al. (2019). A complete list of the non-mammalian remains from Monticino presented in this study, separated according lombero et al. 2014). to the fissures in which they were recovered, is given in the Supple- mentary information. Family ?Cichlidae Rafinesque, 1818 Acronyms ?Cichlidae indet. brS: brisighella collection hosted at the dipartimento di fig. 3c Scienze della Terra of the Università di firenze; the acronym of the locality is followed by the number of the fissure, then by a letter referred material: One isolated tooth (BRS19/F/1). indicating the group (Av = Aves; H = herpetofauna, F = Fish) and a progressive number; MSf: Museo Civico di Scienze naturali di fa- enza (RA), Italy. description. The tooth is incisiviform, slen- der and laterally compressed. The crown has a lin- gual bulbous structure and is supported by a well-de- systematIc palaeontoloGy veloped pedicel with a nearly rounded cross section. remarks. The configuration of this incisivi- infraclass TeleoSTei Müller, 1845 form tooth is in many ways consistent with that of family Cyprinodontidae Gill, 1865 several members of the family Cichlidae (e.g., Green- Genus Aphanius nardo, 1827 wood 1974; Trewavas 1983), to which it is tentatively referred. Tilapiine cichlids are known based on artic- Aphanius crassicaudus (Agassiz, 1832) ulated skeletal remains in the Messinian deposits of fig. 3a-b the northern Apennines, including the euxinic layers of the basal evaporite cycles of the Gessoso-Solfif- referred material: One abdominal vertebra (BRS2/F/1). era Formation (Carnevale et al. 2003). description. The vertebral centrum is am- Family ?Sciaenidae Cuvier, 1829 phicoelous, thickened, cylindrical and nearly rec- ?Sciaenidae indet. tangular in lateral view. A well-developed rounded fig. 3d parapophysis is present along the lateral surface of the centrum (Fig. 3b). The neural arch and spine referred material: one partially complete abdominal verte- are hypertrophied. The neural spine is conical and bra (BRS19/F/2). obliquely oriented forming an angle of approxi- mately 45° with the main axis of the vertebral cen- description.The vertebra consists of an al- trum (Fig. 3b). There is a prominent neural prezyga- most complete amphicoelous centrum bearing a pophysis at the base of the neural spine. partially preserved left neural prezygapophysis. The remarks. The overall morphology of the neural arches and spine are not preserved at all. in vertebra, and especially the structure and configu- lateral view, the centrum appears massive, subrectan- ration of the neural arch and spine, are consistent gular in outline, longer than high. The anterior face Late Miocene vertebrates from the fissure fillings of Monticino Quarry 301 of the centrum is rather oval, whereas the posteri- a well-developed neck, which separates the main or one is more rounded. The notochordal canal is body from a flat and subvertical anterior condyle. small, characterized by a rounded profile. The max- Ventral crest are mostly broken away (Fig. 4f), but imum height of the anterior face of the centrum they were most likely well-developed in origin. few measures 23.7 mm, the maximum width is 24.1 mm, subcentral foramina are preserved on the left side. whereas the anteroposterior length measured along The transverse processes are missing. The neural the ventral edge of the centrum is 28.8 mm. The arch is also mostly missing, but the preserved an- dorsal prezygapophysis is stout and obliquely orient- terior half appears moderately high and rather flat ed, forming an angle of about 45° with the main axis (most likely rising to some degree posteriorly). A of the centrum. The lateral surface of the centrum neural spine was present, even though it is broken. is extensively ornamented with delicate striae and The spine almost reaches the anterior margin. The ridges and deep pits of different size and shape (Fig. margin is straight in dorsal view (Fig. 4a) and is lo- 3d). What appears to be a moderately large articular cated close to the anterior margin (roughly by the facet (or fossa) for rib articulation can be observed end of the anterior fourth) of the only prezygap- anteroventrally on the lateral side of the centrum. A ophysis preserved, which is the left one. The latter deep groove is present along the ventral surface of is elliptical. The anterior zygapophyseal crests are the centrum. moderately developed. in anterior view, the neural remarks. The overall morphology of the canal is subelliptical in cross section and slightly centrum, the absence of parapophyses, and the pe- larger than the anterior condyle (Fig. 4b). culiar ornamentation are reminiscent of those of remarks. The opisthocoelous trunk ver- many “percoid” percomorphs, especially those of tebra of a small salamandrid is present in the fos- certain croakers (Sciaenidae), jacks (Carangidae), sil material from brisighella. The combination of and groupers (Serranidae). The complete absence subvertical condyle, moderately high neural arch, of parapophyses and the presence of what appears well-developed ventral crest, and anterior margin to be an articular facet for a rib concur to suggest of the neural arch straight and located by the ante- that the abdominal vertebra described herein can be rior fourth of the prezygapophyseal facets hints for interpreted as an anterior trunk element, most likely the referral of this specimen to the genus Lissotriton the third element of the abdominal series. The lack (Ratnikov & Litvinchuk 2007). The very small size of unambiguous distinctive features do not allow would suggest its belonging to a small represent- to provide an unquestionable taxonomic assign- atives of the genus, such as the extant Lissotriton ment for the vertebra described above. However, vulgaris and Lissotriton helveticus or the extinct Lisso- the abundance of croaker otoliths (e.g., Leptosciaena triton rohrsi (Herre 1955). However, the poor pres- caputoi, Trewasciaena kokeni) in the deposits of the Co- ervational status and the very short length of the lombacci Formation (Carnevale et al. 2006a; 2018; vertebra even compared to the ranges reported for 2019; Bannikov et al. 2018) may support a tentative the above-mentioned small species hints for a more assignment to the family Sciaenidae. cautious identification limited to the genus level. urodela indet. Class AMPHIBIA linnaeus, 1758 fig. 4g-n order urodela duméril, 1806 family Salamandridae Goldfuss, 1820 referred material: One caudal vertebra (BRS2/H/6); one radius (BRS25/H/244); three ilia (BRS1bis/H/4; BRS25/H/67, Genus Lissotriton bell, 1839 209). Lissotriton sp. description fig. 4a-f Caudal vertebra. The caudal vertebra (Fig. 4g- referred material: One trunk vertebra (BRS25/H/243). l) is small, with a centrum length of 2 mm. The centrum is opisthocoelous, with a well-developed description. The trunk vertebra is very small neck anteriorly. The anterior surface of the condyle (centrum length is less than 1.5 mm) and rather is eroded, but it appears flat and slightly inclined poorly preserved. The opisthocoelous centrum has ventrally. The neural canal is subtriangular in ante- 302 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Fig. 4 - Urodeles from Monticino Quarry - Lissotriton sp.: trunk vertebra (BRS25/H/243) in dorsal (a), anterior (b), left lateral (c), right lateral (d), posterior (e) and ventral (f) views. Urodela indet.: caudal vertebra (BRS2/H/6) in anterior (g), posterior (h), dorsal (i), ventral (j), right lateral (k) and left lateral (l) views; right ilium (BRS25/H/67) in lateral (m) and medial (n) views. Scale bars = 0.5 mm. rior view and about the same size of the condyle cept for BRS1bis/H/4, which is small). They have (Fig. 4g). The prezygapophyses are horizontal in an anterior rod-like portion and expands posterior- anterior view. The postzygapophyses are missing. ly. The acetabulum is rather wide (Fig. 4m), with a The neural arch has a straight anterior margin that distinctly raised anterior margin. is located slightly anteriorly to the midlength of remarks. These remains clearly pertain to the prezygapophyses (Fig. 4i). The neural spine is urodeles. The caudal vertebra is comparable with completely broken away, but its anterior end was the one above referred to Lissotriton in terms of located quite far from the anterior margin of the size, but few morphological differences are present. neural arch (roughly at the level of the posterior end These are the slightly lower anterior portion of the of the prezygapophyses). In lateral view, the neu- neural arch and the slightly more posteriorly located ral arch is quite low anteriorly, but it rises steeply anterior margin of the neural arch. it is not clear posteriorly (Fig. 4k-l). A rather wide and moderately whether these differences might be interpreted as deep posterior notch is present (Fig. 4i). The neural taxonomically significant or they are just individu- spine contacts the notch. The zygapophyseal crests al and/or columnar variation. Because of this, the are rather well developed. The ventral crests, on the caudal vertebra is only referred to an indeterminate other hand, are less developed (Fig. 4j). The later- urodele here. There is no known diagnostic feature al processes of the vertebra are either missing (the for the ilia and radia of urodeles, which are there- left one; Fig. 4l) or poorly preserved (the right one; fore identified only at order level here. They might Fig. 4k). Ventrally, a well-developed hemal arch is pertain to the already recognised Lissotriton as well, present. but the presence of slightly larger specimens (e.g., Radius. The radius is rather small and slender. some of the ilia) could also be evidence of the pres- Ilia. The ilia (Fig. 4m-n) are medium-sized (ex- ence of other, larger urodeles in Monticino. Late Miocene vertebrates from the fissure fillings of Monticino Quarry 303 order Anura fischer, 1813 Superfamily discoglossoidea Günther, 1858 family Alytidae fitzinger, 1843 Subfamily discoglossinae Günther, 1858 Genus Latonia v. Meyer, 1843 ?Latonia sp. fig. 5a referred material: Two urostyles (BRS1bis/H/3; BRS25/H/205). description. The urostyle BRS25/H/205 (Fig. 5a) is small. It has two anterior cotyles, which are clearly separated in the middle. due to poor preservation, their exact shape in anterior view can- not be stated with confidence. The neural arch bears no crista dorsalis, but it has two transverse process- es located near the anterior end. The processes are broken on both sides, but they clearly have a rather enlarged base and narrow distally. in spite of being more damaged than the left one, the right process clearly displays a long posterior laminar extension, which is less developed on the left side of the uro- style. The neural arch is dorsally closed only by its anterior end, being still open posteriorly. Just poste- rior to the closed part, the left margin of the aper- Fig. 5 - Discoglossoid anurans from Monticino Quarry - ?Latonia sp.: urostyle (BRS25/H/205) in dorsal view (a). ?Discoglos- ture is broken, thus resulting in a moderately large soidea indet.: atlas (BRS25/H/203) in anterior (b), posterior space. More posteriorly, the aperture is represented (c) and ventral (d) views; sacral vertebra (BRS25/H/207) in dorsal view (e). Scale bars = 1.5 mm. by a narrow space. in anterior view, the neural canal is suboval. The other urostyle, BRS1bis/H/3 is less preserved, but it shares a comparable morphology. Latonia. Moreover, these laminae can even be asym- in this specimen, the cotyles retain their original metrically developed on the two sides in some cas- shape, being subcircular but slightly mediolaterally es (Hossini 1993; Rage & Hossini 2000). A certain elongated. affinity of these two specimens with Latonia can remarks. following the diagnostic key pro- therefore be suggested, even though the absence of vided by Bailon (1999), the combination of bicotylar other, more taxonomically significant bones in the sacrourostilar articulation, presence of transverse material hinders a completely confident attribution. processes, and absence of a crista dorsalis identi- fies urostyles of Alytes and Discoglossus. The same ?Discoglossoidea indet. morphology is also found in Latonia (Roček 1994; fig. 5b-e Biton et al. 2016), being therefore probably diag- referred material: One atlas (BRS25/H/203); one sacral nostic for Alytidae as a whole. The dorsal fissure vertebra (BRS25/H/207). is found in both members of the subfamily dis- coglossinae, Discoglossus and Latonia (Roček 1994; description Biton et al. 2016), but not in Alytes. These urostyles Atlas. BRS25/H/203 is a fragment of a rather are therefore evidence for the presence of a small large, opisthocoelous atlas (Fig. 5b-d). Anteroposte- discoglossine anuran in the herein-studied material. riorly, the centrum is short (Fig. 5d). Only part of Biton et al. (2016) stated that Discoglossus lacks hori- the left anterior cotyle is preserved (Fig. 5b) and it zontal laminae related to the transverse processes, is not possible to observe whether it coalesced with which are on the other hand present sometimes in the right one or not. The posterior cotyle is circular 304 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. (Fig. 5c). There is no sign of a notochordal canal. opisthocoelous nature of the vertebrae. ranids also on the ventral surface of the atlas, there is no keel have sacral vertebrae with an anterior condyle, but (Fig. 5d). The rest of the vertebra is missing. they display subcircular posterior condyles. Sacral vertebra. The sacral vertebra (Fig. 5e) is moderately large, preserving only the centrum. it Superfamily Hyloidea Stannius, 1856 has a subelliptical anterior condyle and two medio- family bufonidae Gray, 1825 laterally elongated posterior condyles. Genus Bufo laurenti, 1768 remarks. Among anurans reported as ei- ther fossils or living in laurasia, opisthocoelous cf. Bufo gr. bufo (Linnaeus, 1758) vertebrae are found in some members of the su- fig. 6a-e perfamilies Discoglossoidea and Pipoidea (Roček referred material: One humerus (BRS25/H/214); one 2013). The only pipoid family that is known to have ilium (BRSNoNumber/H/12); two femurs (BRS25/H/208, 248). occured in Europe, the extinct Palaeobatrachidae, had procoelous vertebrae, whereas opisthocoelous description ones are found in the Afroamerican Pipidae and Humerus. BRS25/H/214 is a large and robust the American rhinophrynidae. Vertebrae of pipids humerus (Fig. 6a-b). The proximal half of the shaft have a strongly dorsoventrally compressed centrum, is missing, as well as most of the distal end of the which contrast with the uncompressed, posteriorly bone. The eminentia capitata is not preserved, but circular one of the herein-described atlas. Rhinophr- judging from the position of the olecranon scar ynus, the only extant rhinophrynid, has hour-glass (Fig. 6b), it was shifted laterally compared to the shaped vertebrae, which might be more compara- main axis of the bone. The preserved part of the ble with the Monticino atlas. However, these verte- scar appears rather wide. The dorsal surface of the brae are notochordal (Walker 1938), unlike the lat- humerus is flat. Ventrally, a deep fossa cubitalis ven- ter. The morphology of the atlas from Monticino tralis is present (Fig. 6a). The cristae medialis and is more similar to at least some of the opisthoc- lateralis are marked, but not strongly developed. oelous discoglossoids. it clearly differs from bom- Ilium. This large and robust ilium (Fig. 6c) binatoridae because of the wider, shorter and less is poorly preserved. The preserved portion of the compressed centrum, which also does not display a shaft shows no dorsal crest. The dorsal tubercle is flattened ventral surface. The large and circular cen- dorsally rounded and shows some irregularities dor- trum is more comparable with alytids, even though sally. There is no preacetabular fossa. The posterior differences are evident also when BRS25/H/203 is half of the body of the bone is missing. compared with genera included within this group. Femurs. The femurs are large and robust. it does not show horizontal and elongated anterior They bear a well-developed crista femoris. in cotyles as in Alytes (Bailon 1999), whereas it differs BRS25/H/208 (Fig. 6d), the crista is sharp and does from both Discoglossus and Latonia because of the not clearly split into a triangular structure proximal- absence of a ventral keel (Roček 1994; Bailon 1999). ly. However, the absence of the triangle could be The identification of BRS25/H/203 appears there- due to the preservation of the bone, given that the fore rather complicated. Affinities with discoglos- most proximal part is missing and a certain degree soids seems more plausible than with pipoids, but of erosion of the bone surface is recognisable. This all the above-mentioned differences hinder a confi- might be confirmed by BRS25/H/248 (Fig. 6e), dent referral to any of the group included in this su- which seems to display the starting of the split (rep- perfamily. even though alytids seem to be the most resented by a clear thickening of the crista) by the similar group in term of atlas morphology, we here proximalmost preserved portion of the bone. refrain to push the identification that far and prefer remarks. The presence of a large bufonid to tentatively assign the atlas to an indeterminate is testified by few bones, all displaying a compara- discoglossoid taxon. The future recovery of other ble size. The dorsally rounded dorsal tubercle, the fossil material may help to better understand the na- absence of a preacetabular fossa, and the apparent ture of this large anuran. The sacral vertebra is here split of the crista femoris support a referral to the referred to the same taxon because of the similar Bufo bufo species complex (Bailon 1999). However, size and the anterior condyle, indicating a probable the poor preservational status of these elements Late Miocene vertebrates from the fissure fillings of Monticino Quarry 305 fig. 6 - bufonids from Monticino Quarry - cf. Bufo gr. bufo: right humerus (BRS25/H/214) in ventral (a) and dor- sal (b) views; left ilium (BRSNoNumber/H/12) in lateral view (c); right fe- mur (BRS25/H/208) in medial view (d); left femur (BRS25/H/248) in medial view (e). Bufotes gr. viridis: left ilium (BRS15/H/9) in medial view (f); right il- ium (BRS25/H/174) in medial view (g); left ilium (BRS25/H/206) in medial view (h). Scale bars = 1 mm. does not allow a completely confident assignment Femurs. few small femurs have a sharp and and so we treat this identification as tentative here. well-developed crista femoris, which clearly does not split anteriorly. BRS25/H/212 is very small, whereas Genus Bufotes Rafinesque, 1814 the others are distinctly larger (even though still not as large as the femur above referred to cf. Bufo gr. bufo). Bufotes gr. viridis (Laurenti, 1768) remarks. A second, smaller bufonid is also fig. 6f-h present. it displays features of the Bufotes viridis spe- cies complex, such as the deep preacetabular fossa, referred material: Two scapulae (BRS1bis/H/5; BRS3/H/5); nine ilia (BRS5tg2/H/43; BRS15/H/9; BRS19/H/30-31; the bilobed dorsal tubercle, and the not clearly undi- BRS24/H/28; BRS25/H/174, 206; BRSNoNumber/H/24); seven vided crista femoris (Bailon 1999). The supraglenoidal femurs (BRS5tg2/H/41; BRS18/H/8; BRS19/H/29; BRS25/H/212, fossa is also present in Epidalea calamita (Bailon 1999), 216, 223). but the similar size and the absence of other remains clearly referable to the natterjack toad allows to favor description an attribution of the scapulae to the green toads as Scapula. BRS3/H/5 is a small and poorly pre- well. served scapula that clearly display a small supragle- noidal fossa. There is no crista anterior along the bufonidae indet. anterior margin. both ends of the bone are almost completely missing, but the fossa glenoidalis is referred material: Two trunk vertebrae (BRS5bis/H/27; BRS25/H/27); one sacral vertebra (BRS25/H/92); two urostyles clearly posteroventrally directed. BRS1bis/H/5 is (BRS18/H/9; BRS?/H/20); seven humeri (BRS4/H/28; BRS9/H/20; also small, but better preserved. it is rather slender BRS25/H/179; BRS?/H/19). and display a ventrally directed fossa glenoidalis. it also has no crista anterior and is provided with a description small supraglenoidal fossa. both the pars acromialis Trunk vertebra. BRS25/H/27 is a well-preserved and glenoidalis are rather robustly built, in propor- trunk vertebra, whereas BRS5bis/H/27 is fragmen- tion. tary and preserves only part of the vertebral centrum Ilia. These ilia (Fig. 6f-h) have a shaft with and part of the left half of the neural arch. The ver- no dorsal crest and a deep preacetabular fossa. tebrae are robustly built and small- (BRS25/H/27) or The dorsal tubercle is well developed and displays medium-sized (BRS5bis/H/27). The centrum is pro- a small accessory lobe anteriorly (except maybe for coelous and well developed. The base of the trans- BRS25/H/206, but its absence there could be a verse processes is located posteriorly to the prezyga- preservational artifact; Fig. 6h). The posterior por- pophysis. tion of the body of the bone is usually missing, but Sacral vertebra. The sacral vertebra is small. it has in BRS25/H/174 (Fig. 6g) and BRS25/H/206 (Fig. a wide and elliptical anterior cotyle and two subcircu- 6h) it preserves a rather short dorsal acetabular ex- lar posterior condyles, which appear slightly mediolat- pansion. erally elongated. The vertebra misses both transverse 306 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. processes. dorsally, it displays a well distinct carina neuralis on the dorsal surface of the neural arch. Urostyles. The small urostyles have two well-dis- tinct articular cotyles anteriorly, which appear me- diolaterally elongate despite being partly damaged. Posteriorly, these bones have a moderately high cris- ta dorsalis, which bears no transverse processes. The neural canal is subtriangular in anterior view. Humeri. few small humeri have a poorly ossi- fied eminentia capitata, a deep fossa cubitalis ventralis, a wide and rather short olecranon scar and a rather flat dorsal surface. The cristae flanking the eminentia are not developed. The proximal half of the bone is always missing. remarks. These remains can be referred to in- determinate bufonids following the diagnostic key of Bailon (1999). A more precise identification cannot be provided, either because the bones represented do not bear significant diagnostic features or because the preservation is too poor. The small size of all these elements may suggest that they pertain to the smaller Bufotes gr. viridis, but their belonging to juvenile indi- viduals of the larger cf. Bufo gr. bufo should also be taken into account as a possibility. Family Hylidae Rafinesque, 1815 Genus Hyla laurenti, 1768 cf. Hyla sp. fig. 7a-b referred material: One ilium (BRS25/H/15). description. This very poorly preserved ilium bears only a small portion of the shaft, which clearly lacks a dorsal crest. The dorsal tubercle is present, Fig. 7 - Hylids and pelobatids from Monticino Quarry - cf. Hyla sp.: left ilium (BRS25/H/15) in lateral (a) and medial (b) views. but it is very poorly preserved. Anteroventrally to Pelobates sp.: left maxilla (BRS25/H/8) in medial (c) and lat- the acetabulum, the ilium is expanded and shows no eral (d) views. ?Pelobates sp.: right ilium (BRS25/H/71) in lateral (e) and medial (f) views. Scale bars = 1 mm. preacetabular fossa. The medial surface of the pre- served portion of the body of the ilium is rather flat. remarks. despite the very poor preservation, this ilium still displays some features that are useful Superfamily Pelobatoidea bonaparte, 1850 for its identification. Following Bailon’s (1999) key, it family Pelobatidae bonaparte, 1850 shares affinities with Hyla, as highlighted by the fol- Genus Pelobates wagler, 1830 lowing combination of features: absence of dorsal crest; presence of dorsal tubercle; anteroventral ex- Pelobates sp. pansion of the body of the bone. because of these, fig. 7c-d but also adopting a cautious approach due to the preservation, we here tentatively identify this fossil referred material: Six maxillae (BRS5tg2/H/44; as cf. Hyla sp. BRS25/H/8, 16, 141, 213; BRS?/H/7). Late Miocene vertebrates from the fissure fillings of Monticino Quarry 307 description. All maxillae preserve only the no. The tubercular (pustular) ornamentation is, on posterior portion, except for BRS5tg2/H/44 and the other hand, typical of the Pliocene-recent spe- BRS25/H/141. In those preserving the posterior cies (see Venczel 2004), even though its presence part, the lateral surface is almost completely covered is already attested since at least the late Miocene by a dense dermal ornamentation (Fig. 7d), which is (MN 11) of Northern Caucasus (Syromyatnikova firmly attached to the bone (i.e., not connected to 2017). The maxillae from Monticino differ from Pe- it by a trabecular layer as in species of Latonia with lobates fuscus in the strongly concave dorsal margin ornamented maxillae; Roček 1994; Syromyatnikova and in the dense ornamentation, appearing in this et al. 2019). This ornamentation is composed by sense more closely comparable with Pelobates cultripes tubercles and ridges. In BRS25/H/8, the best-pre- and Pelobates syriacus (Bailon 1999; Blain et al. 2016). served specimen, tubercles are mostly found in the Maxillae attributed to Pelobates aff. praefuscus from middle of the surface, whereas ridges are mostly lo- the late Miocene of Northern Caucasus (Syromyat- cated towards the margins. In BRS25/H/16, on the nikova 2019) also have a strongly concave dorsal other hand, the tubercles are connected by ridges margin, but they display a less dense ornamentation. also in the middle of the surface. from what can be Pelobates varaldii, on the other hand, share with the observed on the preserved portion, the dorsal mar- Monticino specimens a dense ornamentation, but it gin of the bone is strongly concave. Medially (Fig. seems to have a rather less concave dorsal margin. 7c), a narrow lamina horizontalis is visible. There Taking all of this into consideration, these maxillae is no processus pterygoideus. The alveolar border, are here referred only as Pelobates sp., even though provided with at least 20 closely spaced tooth posi- we can highlight their affinities with the youngest tions in BRS25/H/8, extends far beyond the pos- group of Pelobates species. terior end of the lamina horizontalis. dorsally to the lamina, a large, posteriorly-directed cavity is vis- ?Pelobates sp. ible. The cavity of BRS25/H/8 is enclosed by two fig. 7e-f osseous laminae, which however do not contact, referred material: Three ilia (BRS25/H/71, 105, 282). leaving a narrow slit visible in medial view (Fig. 7c). Anteriorly, this slit is still visible, even though the two laminae contact and fuse. BRS5tg2/H/44 and description. All ilia are poorly preserved. BRS25/H/141 preserve a more anterior portion of The main body is almost entirely missing. The shaft the bone (even if not the anterior end). These spec- bear no dorsal crest and, judging from the preserved imens also present dermal ornamentation on the portion of the bone, the dorsal tubercle is also miss- lateral surface similar to the former ones, but this is ing. Most of the shaft is missing and therefore it is less developed in BRS25/H/141. The lamina hori- not possible to state whether it was straight or more zontalis is less narrow in medial view and is grooved curved. There is no preacetabular fossa. dorsally. At least 21 and 18 tooth positions are pre- remarks. The absence of both a dorsal crest served or partially preserved in BRS5tg2/H/44 and of a dorsal tubercle is diagnostic of Pelodytes and BRS25/H/141 respectively, some of them still and Pelobates according to Bailon (1999). Following hosting complete pleurodont and cylindrical teeth the diagnostic key proposed by the same author, the in the latter. discrimination between ilia of these two genera is remarks. The dermal ornamentation firmly based on structures that are not preserved in the attached to the lateral surface of these maxillae al- specimen described above. However, we tentatively lows their attribution to a pelobatid (Bailon 1999; assign them to Pelobates with doubt given that no Roček 2013). An ornamentation composed by tu- clear remains of Pelodytes are known from Montici- bercles and ridges is found in Pelobates, contra the no Quarry, whereas the former genus is testified pit-and-ridge ornamentation that characterizes Eo- also by other elements. pelobates (Roček 2013; Syromyatnikova 2017, 2019). Some oligo-Miocene species of Pelobates also have a pit-and-ridge ornamentation (Venczel 2004; Syro- Superfamily Ranoidea Rafinesque, 1814 myatnikova 2017, 2019), thus excluding affinities Family Ranidae Rafinesque, 1814 between these forms and the taxon from Montici- Genus Pelophylax fitzinger, 1843 308 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. fig. 8 - ranids from Monticino Quarry - Pelophylax sp.: right ilium (BRS3/H/3) in lateral view (a); right ilium (BRS6/H/23) in lateral (b) and medial (c) views. Scale bars = 1 mm. Pelophylax sp. view. This morphology may somehow recall that of fig. 8 a bufonid (a large taxon, such as cf. Bufo gr. bufo), but the poor preservational status does not allow to be referred material: Three ilia (BRS3/H/3-4; BRS6/H/23). more confident with this identification. description. These three ilia preserve only Class rePTiliA laurenti, 1768 part of the body of the bone and the base of the order testudines batsch, 1788 shaft. Their preservational status is, on the whole, family Testudinidae batsch, 1788 rather poor. The dorsal crest was present in origin, Genus Testudo linnaeus, 1758 even though now it is broken in all specimens. it started from an elongated and well distinct dorsal Testudo s.l. tubercle. In BRS3/H/3, which preserves a small fig. 9a-k dorsal acetabular expansion, an angle close to 90° between the latter and the tubercle is recognis- referred material: Two nuchals (MFS.551; BRS25/H/254); able (Fig. 8a). A similar angle can be inferred for two neurals (BRS25/H/255); two undetermined peripherals (MFS.550,2; MFS.550,3); one left peripheral 7 sutured to the pe- BRS6/H/23 (Fig. 8b-c), even though its dorsal ex- ripheral 8 (MFS.550,4); two bridge peripherals 3 or 7 (MFS.537; pansion is less preserved. BRS?/H/26); one suprapygal (BRS?/H/25); two pygals (MFS.549; remarks. These three ilia can be referred to BRS?/H/27); one entoplastron (BRS?/H/24); one left hyopla- stron (MSF.553,2); one right hypoplastron (MSF.553,1); one tibia Pelophylax based on the following combination of (BRS25/H/274); 29 limb osteoderms (BRS25/H/3,13, 64, 98, 225, characters (Bailon 1999): presence of a dorsal crest; 256, 265, 272, 273; BRS28/H/5). well-distinct dorsal tubercle; angle between the tu- bercle and the dorsal acetabular expansion close to description. Two large, partially preserved 90°. Blain et al. (2015) demonstrated that precise nuchals (MFS.551, Fig. 9a, and BRS25/H/254) identification of Pelophylax species is impossible share the presence of a relatively narrow cervical with a small number of ilia. and a vertebral sulcus with an arched, deeply con- vex anterior profile. The convexity of the vertebral Anura indet. makes the cervical very short on the dorsal surface but it is long (nearly the double in BRS25/H/254) remarks. See the Supplementary informa- on the inner surface. There are no signs of pleu- tion for a complete list of remains of indeterminate ral overlapping on the nuchal. The two neurals anurans. Two specimens are particularly interest- BRS25/H/255 are quadrangular and crossed by an ing and deserve further comments. brSnonum- intervertebral sulcus. The best preserved peripher- ber/H/8 is a very large humerus preserving only als as MFS.550,2 (Fig. 9b) and MFS.550,3 show that the spherical eminentia capitata. it has a rather the pleuro-marginal sulcus overlaps the costo-pe- elongated olecranon scar. These features suggest ripheral suture or is very close to it. The elliptic su- that this bone does not pertain to a bufonid. it prapygal BRS?/H/25 is markedly convex dorsally might belong to the large indeterminate possible but only weakly ventrally. it hosts the vertebro-mar- discoglossoid recognised in the material, but this ginal sulcus very close to, but separated from, the cannot be supported by any clear evidence at the suprapygal-pygal suture. The two pygals are both moment. BRS25/H/181 is the procoelous cen- rather flat and trapezoidal in shape. BRS?/H/27 trum of a large vertebra, which is dorsoventrally (Fig. 9c-d) is slightly wider than long and shows a compressed and slightly ventrally concave in lateral sagittal intermarginal sulcus on its external surface Late Miocene vertebrates from the fissure fillings of Monticino Quarry 309 Fig. 9 - Chelonians from Monticino Quarry - Testudo s.l.: nuchal (MFS.551) in external view (a); peripheral (MFS.550,2) in external view (b); pygal (BRS?/H/27) in external (c) and internal (d) views; pygal (MFS.459) in external view (e); entoplastron (BRS?/H/24) in external view (f) ; left hyoplastron (MSF.553,2) in external view (g); flat osteoderm (BRS25/H/272) in external (h) and lateral (i) views; con- ical osteoderm (BRS25/H/273) in lateral (j) and external (k) views. Mauremys sp.: costal (MSF.70) in external view (l); entoplastron (BRS25/H/257) in external view (m). Scale bar = 5 mm. only; its dorsal (anterior) edge is slightly concave, an hypo-xiphiplastral suture or a hinge. The several whereas the ventral (posterior) edge is straight. limb osteoderms have a general morphology that MFS.459 (Fig. 9e) is about as long as wide and has varies from nearly flat to distinctly conical. The flat no sagittal (intermarginal) sulcus. The incomplete ones (Fig. 9h-i) are generally smaller than the con- entoplastron BRS?/H/24 (Fig. 9f) is nearly circu- ical ones (Fig. 9j-k) that can be up to 10 mm long lar in ventral view. it is modestly overlapped by the and 5 mm tall. Most of them show a sort of base, gulars anteriorly, but in the posterior area it does in some cases partly encircled by a sulcus parallel not host the humero-pectoral sulci. The ventral to the margin, and have a tip that can project out- surface is slightly concave in the anterior area; the side the base or elevate on it. The position of tip is humeral surfaces show longitudinal growth marks. therefore eccentric in all cases. The elliptic base can The visceral surface is totally altered. The left hyo- be concave. plastron MSF.553,2 (Fig. 9g) is compatible with the remarks. The morphology of the shell ele- entoplastron BRS?/H/24 because of the position ments described in this section is broadly congru- of the humero-pectoral sulcus that is located poste- ent with that of a group of Miocene to Holocene riorly to the ento-hyoplastron suture. The partially tortoises from the western Palearctic traditional- preserved right hypoplastron MSf.553,1 hosts the ly referred to the genus Testudo (Lapparent et al. markedly arched lateral end of the abdomino-fem- 2006a,b), but the phylogenetic relationships of oral sulcus. because of the incompleteness of this which have been only partly explored and resolved. element, it is not possible to assess the presence of Since the extinct taxa do not constitute a monophy- 310 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. letic group along with the extant ones (among oth- remarks. The most diagnostic element, the ers, Corsini et al. 2014; Vlachos & Rabi 2018), the entoplastron BRS25/H/257 clearly indicates the material herein described is generically referred to presence of a second turtle taxon in the Monticino a Testudo-like taxon. Worth mentioning is that the assemblage. The position of the sulci (the hume- late Miocene species from Tuscany (Italy) Testudo ro-pectoral sulci crossing the entoplastron in par- amiatae Pantanelli, 1893 has never been formally ticular) is congruent with the morphology of Mau- described, nor its phylogenetic relationships were remys spp. (Hervet 2000; Chesi et al. 2009; Vlachos evaluated in a modern context. The presence or et al. 2019). The peculiar arrangement of the sulci absence of an intermarginal sulcus on the pygal is on the costal MSf.70 is somewhat reminiscent of a character, among others, generally used to distin- Mauremys aristotelica Vlachos et al., 2019 recently de- guish Testudo hermanni Gmelin, 1789 from Testudo scribed for the late Miocene-Pliocene of northern graeca linnaeus, 1758, but there is also a certain Greece (Vlachos et al. 2019) that is characterized degree of intraspecific variation (Lapparent et al. by very broad vertebral shields. The broadness of 2006a,b) and therefore MFS.459 and BRS?/H/27 the vertebral shield and the degree of concavity of could perhaps belong to the same taxon (but the the vertebral lateral edges in MSf.70 seems to be difference in terms of general shape should also intermediate between the one of M. aristotelica and be taken into consideration). The limb osteoderms that of Mauremys campani (Ristori, 1891) (see Chesi have been tentatively referred to Testudo s.l. because et al. 2009) hindering a specific identification of the of their overall resemblance with those of extant geoemydid from Monticino Quarry that is there- species of this genus. fore simply assigned at genus level. The small neu- ral BRS25/H/258 is tentatively referred to the same family Geoemydidae Theobald, 1868 taxon on the basis of its broad dorsal keel. Genus Mauremys Gray, 1869 Testudines indet. Mauremys sp. fig. 9l-m remarks. See the Supplementary informa- referred material: One neural (BRS25/H/258); one costal tion for a list of the remains identified only as Tes- 6 (MSF.70); one entoplastron (BRS25/H/257). tudines indet. description. The incomplete, very small order crocodylia Gmelin, 1789 neural BRS25/H/258 is characterized by a sulcus crocodylia indet. crossing transversally its anterior portion and by a fig. 10 marked constriction posterior to it. The external surface of the constricted portion has a broad keel. referred material: Two teeth (BRS3/H/2; BRS6/H/6). The costal MSF.70 (Fig. 9l) is rather complete and curved (likely being a right sixth element). Its ex- description. The two isolated, conical teeth ternal surface hosts the y-shaped sulci left by the BRS3/H/2 and BRS6/H/6 are both characterized vertebral shield and by the two pleurals. The lateral by being represented by the crown only and by hav- sides of the vertebral shield are markedly concave ing a conical depression at their base. BRS6/H/6 and therefore the lateral corner of the vertebral (Fig. 10a) is slightly taller than BRS3/H/2 (10.5 and has an acute angle. The triple junction of the sulci 8.5 mm respectively) and has a larger base (8.1 and is located approximately in the middle of the ele- 5.2 mm respectively). The major breakage shows ment. The small entoplastron BRS25/H/257 (Fig. several overlapping inner layers. The outer surface 9m) is anteriorly incomplete. It is crossed by near- of BRS6/H/6 is significantly damaged and in the ly straight humero-pectoral sulci in its posterior only preserved area it shows few shallow, longitu- area (but far from the posterior edge of the en- dinal groves separating small ridges. Two evident toplastron). Careful inspection with a microscope mesiodistal carinae separate the lingual and the la- indicated that the anterior breakage followed the bial surfaces. BRS3/H/2 (Fig. 10b-c) is slender and gulo-humeral sulci that therefore likely crossed the more curved than the previous teeth. its surface is anterior most portion of the entoplastron. also better preserved and shows more ridges and Late Miocene vertebrates from the fissure fillings of Monticino Quarry 311 teeth are significantly larger and more massive (for an overview on its presence in the Mediterranean area see Piras et al. 2007). During the late Miocene, crocodylians were rather common in the Mediterra- nean area of what is now europe, but they are often represented by isolated teeth (among others, Geor- galis et al. 2016b; Colombero et al. 2017; zoboli et al. 2019) and therefore they were rarely identified at genus level. order squamata oppel, 1811 Family Agamidae Spix, 1825 Subfamily Agaminae Spix, 1825 Agaminae indet. fig. 11 referred material: Two fragmentary maxillae (BRS25/H/4, 5); one dentary (MSF 67). description Maxillae. The fragmentary maxillae are robust and very poorly preserved. both specimens car- ry acrodont teeth, the number of which is two in BRS25/H/4 and five in BRS25/H/5 (Fig. 11a-b). The teeth are triangular and closely spaced. despite the wearing, a very small cusp is visible in some of them, both anteriorly and posteriorly to the main Fig. 10 - Crocodylians from Monticino Quarry - Crocodylia indet.: tip. The tooth base is expanded towards the medi- isolated tooth (BRS6/H/6) in labial view (a); isolated tooth al surface of the alveolar border, rather than being (BRS3/H/2) in labial (b) and mesial (c) views. Scale bar = 2 mm. located on its edge (Fig. 11a). The size of the teeth increases towards one of the end of both bones, which is probably the posterior one. in lateral view, grooves than that of the previous one. The lingual there are no interdental grooves (Fig. 11b). surface, slightly concave, hosts a larger number of ridges than the nearly smooth labial surface. only one of the mesiodistal carine is preserved. remarks. The two isolated teeth BRS3/H/2 and BRS6/H/6 clearly show a reabsorbed root in- dicating that they were lost before the death of their owner. Since isolated crocodylian teeth are, with very few exceptions, not diagnostic even at family level, the remains from Monticino Quarry are simply re- ferred at order level despite they likely belong to Crocodylus laurenti, 1768, the only crocodylian with similar teeth so far identified in the central Medi- terranean sector during the late Miocene (Delfino et al. 2007; Delfino & Rook 2008; Delfino & Rossi 2013). Young long snouted crocodylians belonging to Tomistoma Müller, 1846 have teeth that are usual- Fig. 11 - Agamids from Monticino Quarry - Agaminae indet.: left ly more slender than those from Monticino Quar- maxilla (BRS25/H/5) in medial (a) and lateral (b) views; left ry, whereas later in ontogeny at least some of the dentary (MSF 67) in lateral view (c). Scale bars = 1 mm. 312 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Dentary. The dentary (Fig. 11c) is almost larger part of the anterior half of the bone. in spite complete, but still plunged in a block of sediment. of the anterior portion being always absent, a mid- only the smooth lateral surface is visible, showing dle constriction seems to be recognizable in dorsal four mental foramina. The tooth row is roughly 17 view. The dorsal surface is covered by a dermal or- mm long and carries 19 teeth. The two anterior- namentation made up of a crust covered by small most ones are subpleurodont, conical and pointed, pits and low ridges (Fig. 12a, e). Three grooves are whereas all other teeth are acrodont, closely spaced visible on it, separating the frontal shield from the and subtriangular. distinct interdental grooves are frontoparietal ones and the latter from one another. recognisable. The ventral margin of the bone is In BRS6/H/3, however, a small and triangular area straight. The posterior end is broken off. (the vertex is directed posteriorly) is visible in the remarks. in spite of the possible presence middle of the posterior end of the bone, between of variability in the main characters used to sep- the frontoparietal shields. Moreover, the ornamen- arate the two families (see Rage & Bailon 2011), tation is slightly less developed in this same speci- fragmentary tooth-bearing bones of Agamidae are men. The ornamentation is more developed on the usually distinguished from those of Chamaeleoni- frontoparietal shields, whereas it is poorly defined dae because of the closely spaced acrodont teeth on the frontal one. The grooves separating frontal extending onto the lingual side of the alveolar bor- and frontoparietal shields are located posteriorly to der (Evans et al. 2002; Delfino et al. 2008; Villa & the middle constriction. by the posterior margins Delfino 2019a). The presence of an agamid in the of the preserved portion of the specimens, the an- late Miocene of Monticino is further supported by terior end of the articulation surfaces with the post- the isolated dentary MSf 67, which carries both frontals are visible (Fig. 12c-d). A portion of those pleurodont and acrodont teeth (Delfino et al. 2008; with the prefrontals is visible by the anterior end of Blain et al. 2014; Villa & Delfino 2019a). The num- the preserved portion of BRS5bis/H/11 (Fig. 12d), ber of subpleurodont teeth suggests that MSf 67 BRS6/H/3, BRS24/H/22, and BRS?/H/14. The belongs to a member of the subfamily Agaminae two articulation surfaces are far from each other. (Maul et al. 2011). The length of the specimens is 2 mm in brS5t- g27h/2 (which, however, is only a very small frag- family lacertidae batsch, 1788 ment), 3 mm in BRS24/H/22, 4 mm in BRS6/H/3 Subfamily lacertinae oppel, 1811 and BRS?/H/14, 5.3 mm in BRSNoNumber/H/14, Tribe eremiadini Shcherbak, 1975 5.5 mm in BRS4/H/23, and reaches 6 mm in BRS- eremiadini indet. 5bis/H/11. BRS25/H/87 is represented only by a fig. 12a-f fragment of the anterior portion, but its morphol- ogy and its size are similar to those of the other referred material: eight pairs of fused frontals specimens. (BRS4/H/23; BRS5bis/H/11; BRS5tg2/H/2; BRS6/H/3; BRS24/H/22; BRS25/H/87; BRSNoNumber/H/14; BRS?/H/14). remarks. Among lacertids, fused frontals lacking a longitudinal suture line in ventral view description. each specimen represents a (meaning that they fused in an early stage of the pair of fused frontals. The anterior end is always ontogeny) are found only in some members of the missing, whereas the posterior one is preserved tribe Eremiadini (Arnold 1989; Arnold et al. 2007; partially in BRS4/H/23 (which misses the posterior Villa & Delfino 2019a). Following Arnold (1989) margin and the posterolateral processes; Fig. 12a-c), and Hipsley & Müller (2017), fused frontals with a BRS5bis/H/11 (Fig. 12d-f), and BRS6/H/3 (which strong middle constriction are related to adaptation both lack the right posterolateral process and the to arid environments in lacertids. The poor preser- tip of the left one). In ventral view, no suture line vation of the specimens from Monticino, the ab- is visible in the middle (Fig. 12b, f). Two low and sence of other taxonomically significant elements moderately robust cristae cranii run along the lat- clearly referable to this tribe of lacertids and the eral margins. The cristae are more sharpened and fact that no other eremiadini-like frontals to which slightly more developed in the anterior portion the herein considered remains could be compared of BRS5bis/H/11, BRS5tg2/H/2, BRS6/H/3, are known from Miocene localities in europe pre- BRS24/H/22, and BRS?/H/14, which preserve a vent a more precise identification. Late Miocene vertebrates from the fissure fillings of Monticino Quarry 313 Fig. 12 - Lacertids from Monticino Quarry - Eremiadini indet.: fused frontals (BRS4/H/23) in dorsal (a), ventral (b) and right lateral (c) views; fused frontals (BRS5bis/H/11) in left lateral (d), dorsal (e) and ventral (f) views. Lacertidae indet.: right jugal (BRS25/H/187) in lateral (g), medial (h) and dorsal (i) views; left dentary (BRS4/H/21) in medial view (j); left dentary articulated with the splenial (BRS25/H/148) in medial view (k); right lower jaw (BRS25/H/251) in medial (l) and lateral (m) views. Scale bars = 1 mm. lacertidae indet. nate a very short medial process (Fig. 12i). A large fig. 12g-m foramen opens on the posteromedial surface of the bone, ventral to this latter process. The lateral referred material: Four fragmentary maxillae (BRS4?/H/5; surface is completely covered by a light but distinct BrS24/H/17; BRS28/H/17); one jugal (BRS25/H/187); two dentaries (BRS4/H/21; BRS?/H/13); 22 fragmentary denta- vermicular ornamentation (Fig. 12g). ries (BRS4/H/20; BRS5tg2/H/16; BRS8/H/1; BRS18/H/6; Tooth-bearing bones. The tooth-bearing bones BRS19/H/11; BRS24/H/18, 21, 35; BRS25/H/184; BRS27/H/12; bear cylindrical, slender and pleurodont teeth, BRS28/H/18); four fragmentary dentaries articulated with the among which the best preserved ones show a splenial (BRS5tg2/H/18; BRS25/H/11, 12, 148); 71 fragments of tooth-bearing bones (BRS4/H/19; BRS4?/H/6; BRS5bis/H/10; mono-, bi- or tricuspid crown. The best preserved BRS5tg2/H/17; BRS8/H/2; BRS15/H/6; BRS16/H/4; specimens, BRS4/H/21 (Fig. 12j) and BRS?/H/13, BRS18/H/5; BRS19/H/10, 12; BRS24/H/19; BRS25/H/10, 47, 84, are dentaries, with preserved portions reaching 147, 183; BRS27/H/13); two teeth (BRS5tg2/H/1; BRS25/H/48); one fragmentary lower jaw (BRS25/H/251). about 6.5 mm in length and carrying 18 and 17 tooth positions respectively; the posterior portion, description. All specimens are very frag- however, is missing in both of them. The lateral mentary, poorly preserved and small- or medi- surface of both maxillae and dentaries is smooth um-sized. and the one of BRS28/H/17, a maxilla, is also dis- Jugal. The medium-sized jugal, BRS25/H/187 tinctly concave. Moreover, on the dorsal surface (Fig. 12g-i), preserves only the middle portion, of the palatal shelf of the maxillae, there is a wide where the anterior and the posterodorsal processes and posteriorly directed superior dental foramen, meet. both processes seem to be moderately robust continuing posteriorly in a wide groove. when pre- and the base of a well-developed quadratojugal pro- served, the mandibular symphysis of the dentaries cess is present, though its tip is missing. The palatal is narrow and subhorizontal, whereas the Meckelian process is also well developed and seems to origi- fossa is wide and medially opened. In BRS24/H/21 314 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. and BRS?/H/13, a convex ventral margin is also recognised indeterminate member of the tribe er- recognizable. In BRS5tg2/H/18, BRS25/H/11, emiadini or they represent a second lacertid taxon. BRS25/H/12, and BRS25/H/148, the fragment Nevertheless, it has to be noted that in all extant of dentary is still articulated with a fragment of European Eremiadini the palatal process of the ju- the splenial. Though poorly preserved, the splenial gal is poorly developed (Villa & Delfino 2019a), in is wide and narrows towards the anterior end. in contrast with the condition seen in BRS25/H/187. BRS25/H/148 (Fig. 12k), a large anterior inferior foramen (dorsally) and a very small anterior mylo- Scincidae Gray, 1825 hyoid foramen (ventrally) are visible in the middle Scincidae indet. of the bone in medial view. fig. 13a Lower jaw. BRS25/H/251 preserves only part referred material: one fragmentary dentary of the middle portion of the articulated right low- (BRS25/H/142). er jaw (Fig. 12l-m). Its preservational status is very poor and it is partially covered by matrix. The to- description. This specimen is a poorly pre- tal length of the specimen is 6.5 mm. Anteriorly, served fragment of a small dentary, carrying pleu- the posterior portion of the dentary is preserved. rodont, cylindrical and slender teeth. only two teeth It has a wide and medially opened (but covered by are preserved, but they are in good preservational the splenial) Meckelian fossa and bears four pleu- conditions. The crowns are bicuspid, with parallel rodont, cylindrical and closely-spaced teeth. The lingual and labial cusps, and appear pointed in me- tooth crowns are eroded, but a bicuspid condition dial view. The tip curves slightly in posteromedial is recognisable in the two posteriormost teeth. The direction and light striae are visible only on the lin- lateral surface of the dentary is smooth and its ven- gual surface of the tooth crowns. tral margin appears convex. The posterior process- remarks. BRS25/H/142 is referred to Scin- es are not recognisable. The splenial has a strongly cidae because of the pleurodont, bicuspid teeth concave medial surface, the concavity of which is provided with parallel labial and lingual cusps, a maybe enhanced as a taphonomic artifact. on the slightly bending crown and striae on the lingual sur- anterior preserved part of this bone, a large anteri- face (Townsend et al. 1999; Caputo 2004; Villa & or inferior foramen is present, flanked ventrally by Delfino 2019a), but the very poor preservational the small anterior mylohyoid foramen. The posteri- status prevent a more precise attribution. or half of the specimen is composed by the ante- rior end of both the compound bone and the free ?Scincidae indet. angular, but the preservational status hinders rec- fig. 13b-c ognition of main morphological features of these bones. dorsally, part of the coronoid is visible. it referred material: one fragment of tooth-bearing bone has a moderately short and pointed labial process. (BRS27/H/15). Medially, the anteromedial process is largely cov- ered by dentary and splenial, whereas the postero- description. BRS27/H/15 is a small frag- medial process displays a well-developed coronoid ment of tooth-bearing bone carrying four pleuro- ridge. The coronoid process is not preserved. dont, cylindrical and slender teeth. Teeth are well remarks. The tooth morphology of the preserved and show a bicuspid crown, with parallel above-described tooth-bearing bones, as well lingual and labial cusps. The cusps are only slight- as the convex ventral margin of BRS24/H/21, ly distinguishable, but this could be due to erosion. BRS25/H/251, and BRS?/H/13, are clear evidence The tip of the crown is strongly curved in postero- of them being attributable to the family lacertidae medial direction and distinct striae are clearly visible (Delfino et al. 2011; Villa & Delfino 2019a). The on both the labial and lingual surfaces. jugal, on the other hand, can be assigned to lacer- remarks. The tooth morphology of tids because of the vermicular ornamentation and BRS27/H/15 recalls the one of scincids (see the overall morphology not as slender as in scincids above), but the strongly curved crown and the stri- and anguids (Villa & Delfino 2019a). It is not clear ae on the labial surface distinguish this specimen whether these specimens belong to the previously from the one attributed above to this latter family, Late Miocene vertebrates from the fissure fillings of Monticino Quarry 315 as the posterior one and there is no lateral process. The articular surface with the nasal is clearly distin- guishable by the anterior end. Moreover, the frontal is not constricted in the middle and its lateral mar- gin is straight. The posterolateral process is short, moderately wide and roughly pointed. The posteri- or margin is straight, but slightly irregular. in later- al view, both the articulation surfaces with the pre- frontal and the postfrontal are visible (Fig. 14a). The former is very wide and surpasses the middle of the length of the bone, whereas the latter is smaller and covers only the posterior fifth of the lateral margin. The two surfaces are distinctly separated from one another. A wide crista cranii is visible on the ventral surface: it is low in its posterior half, but becomes more ventrally developed anteriorly. The ventral Fig. 13 - Scincids from Monticino Quarry - Scincidae indet.: right portion of its anterior half, however, is missing. A dentary (BRS25/H/142) in medial view. ?Scincidae indet.: fragment of tooth bearing bone (BRS27/H/15) in medial well-developed dermal ornamentation is present on (b) and lateral (c) views. Scale bars = 1 mm. the dorsal surface, but the orbital margin remains smooth (Fig. 14b). The frontal shield covers almost BRS25/H/142. According to Kosma (2004), the entirely the bone, but the small areas covered by the parallel cusps and the distinct lingual striation are frontoparietal (laterally) and the interfrontal (medi- also present in the dentition of Cordylidae, but teeth ally) shields are visible by its posterior end. The or- of this family are usually more robust and lack striae namentation is less developed on these latter shields on the labial surface. BRS27/H/15 is here tenta- and the grooves separating them from the frontal tively attributed to an indeterminate scincid (maybe one are recognisable. different from BRS25/H/142), but the possibility Parietal. BRS25/H/119 is a fragmentary pa- that it belongs to a different taxon cannot be totally rietal (Fig. 14d-e), preserving only the right part ruled out. of the shelf. A short, rounded and slightly lateral- ly developed anterolateral process is present by the anterolateral corner. There are no distinct interdig- family Anguidae Gray, 1825 itations on the anterior margin. The dorsal surface Subfamily Anguinae Gray, 1825 is covered by a strong dermal ornamentation (Fig. Genus Ophisaurus daudin, 1803 14d), which does not reach the lateral margin. The ornamentation is made up of very low ridges and Ophisaurus sp. pits. A smooth area levis is also present posteriorly. fig. 14 The right lateral, interparietal, and occipital shields are recognizable, whereas the right frontoparietal referred material: One frontal (BRS25/H/160); one parie- tal (BRS25/H/119); one fragmentary parietal (BRS25/H/121); one one is absent since the lateral one reaches the an- fragmentary maxilla (BRS5tg2/H/7); two dentaries (BRS25/H/7, terior margin. The occipital shield is small, whereas 135); seven fragments of tooth-bearing bone (BRS4/H/22; the interparietal one is very large. Anteriorly, the lat- BRS4?/H/4; BRS5tg2/H/6; BRS25/H/6, 137; BRS27/H/14); 12 ter occupies a wide portion of the anterior margin, teeth (BRS5tg2/H/4; BRS24/H/10; BRS25/H/136, 271); two trunk vertebrae (BRS5bis/H/35; BRS28/H/15). without reaching the anterolateral corner though. The lateral margin of the lateral shield is straight. description The parietal foramen is large and circular and it is Frontal. BRS25/H/160 (Fig. 14a-c) is a me- located anteriorly to the middle of the shelf. The dium-sized and well-preserved frontal (length = 9 ventral surface is smooth (Fig. 14e), except for a ro- mm), missing only the tip of the medial process and bust and well-developed anterolateral ventral crest the anterior part of the crista cranii. in dorsal view running along the lateral margin. There is no facies (Fig. 14b), the anterior end is roughly half as wide muscularis (sensu Klembara et al. 2010) laterally to 316 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. fig. 14 - Ophisaurus from Montici- no Quarry - Ophisaurus sp.: left frontal (BRS25/H/160) in lateral (a), dorsal (b) and ventral (c) views; pa- rietal (BRS25/H/119) in dorsal (d) and ventral (e) views; right maxilla (BRS5t- g2/H/7) in medial view (f); left dentary (BRS25/H/135) in lateral view (g); trunk ver- tebra (BRS28/H/15) in dor- sal (h), ventral (i), anterior (j) and right lateral (k) views. Scale bars = 1 mm. the crest. The specimen is large-sized, being 9.5 al status. The specimen is moderately robust and mm in length. BRS25/H/121 is a fragment of a 12.5 mm in length. it carries at least 10 tooth po- robust and large-sized parietal, representing only sitions. The teeth are large, subpleurodont, close- the left anterolateral corner of the shelf. A poor- ly spaced and canine-like, provided with a slightly ly developed anterolateral process is recognizable posteromedially bending tip. no distinct striae are and very little developed interdigitations are pres- visible neither on the lingual, nor on the labial sur- ent along the anterior margin. The ventral surface face of the teeth. Two distinct cutting edges run shows the anterior end of the left anterolateral along both the anterior and posterior margins of ventral crest, whereas the dorsal one bears a strong the crown, with the anterior one being strongly dermal ornamentation. A portion of the interpari- more developed than the posterior one. etal and of the left lateral shields are recognisable, Dentaries. BRS25/H/135 preserves only the since they are separated by a distinct groove. The anterior half of a left dentary, still plunged in the latter shield reaches the anterior margin and so the sediment (Fig. 14g). Only the external surface of left frontoparietal shield is not visible. the bone and the dorsal portions of four teeth are Maxilla. BRS5tg2/H/7 is a very fragmentary visible. five labial foramina are visible on the lat- maxilla (Fig. 14f), preserving only part of the tooth eral surface and a sixth one is cut by the poste- row. The posterior process is completely missing, rior breakage. The teeth are conical, slender and whereas only the anterolateral portion of the an- slightly posteromedially bending by their pointed terior premaxillary one is preserved. The antero- tip. There are no striae on their exposed surface. lateral process is long, pointed in dorsal view and The second dentary, BRS25/H/7, includes the rather rounded in lateral view. The lateral surface posterior half, even if the angular process and the is smooth and the ventrolateral foramina are not ventral margin are missing. Teeth are morphologi- recognizable because of the poor preservation- cally similar to those of BRS5tg2/H/7 and a total Late Miocene vertebrates from the fissure fillings of Monticino Quarry 317 of eight tooth positions is present. A small and physis. it is wide, subelliptical and dorsally inclined pointed splenial spine is present by the fifth tooth of about 45°. in anterior view, the neural canal is position from the end of the tooth row. The open- higher than the cotyle (Fig. 14j). ing of the alveolar canal is visible, in ventral view, remarks. The tooth morphology of the under the third posteriormost tooth position. be- above described tooth-bearing bones recalls that cause of the absence of the ventral half of the of Ophisaurus (Klembara et al. 2014). Despite be- posterior end, it is not possible to see whether a ing very fragmentary, BRS25/H/119 is clearly not free posterior projection of the intramandibular attributable to Pseudopus (the only other anguid septum was present or not. The superior posterior identified in the remains from Monticino; see be- process is made up by two projections: the dorsal low) because of the absence of a facies muscularis one (the coronoid process) is pointed and distinct- (Klembara 1986; Klembara et al. 2017). Moreover, ly longer than the more rounded ventral one (the parietals of both Anguis and Pseudopus usually dis- surangular process). Because of this, the notch be- play small frontoparietal ornamented shields by tween the two projections, the coronoid incisure, the anterolateral corners of the shelf preventing is very shallow in lateral view. The lateral surface is the lateral shields to reach the anterior margin of smooth, showing only two mental foramina. the bone (Villa & Delfino 2019a). This appears not Tooth-bearing bones and isolated teeth. to be true for at least some Ophisaurus species (see BRS4?/H/4, composed by two separated por- e.g., Ophisaurus holeci in Klembara 2015). It seems tions, was part of a large-sized tooth-bearing possible, therefore, to assign the herein described bone. it shows four tooth positions, bearing sub- parietals to Ophisaurus. The frontal from brS 25 pleurodont teeth. Two of the four teeth are well is also not attributable to Pseudopus because of the preserved, whereas only the base remains of the completely smooth lateral margin (Klembara 2015; other two. The well-preserved teeth are morpho- Klembara et al. 2017; Villa & Delfino 2019a). This logically similar to those of BRS5tg2/H/7. The is also a feature present in Anguis frontals, but in lateral surface of the tooth-bearing bone appears the latter genus this bone displays a contact be- rough and four foramina are visible on it, roughly tween the articular surfaces for the postfrontal anteroposteriorly aligned. The isolated teeth and and the postorbital and a not clearly distinguisha- the teeth carried by most of the other fragments ble articular surface with the nasal (Villa & Delfino of tooth-bearing bones are similar in morphology 2019a). Given that, the above mentioned frontal to those of BRS4?/H/4 and BRS5tg2/H/7, even is also assigned to Ophisaurus. The trunk vertebrae if BRS25/H/136 differs from them in the absence are attributed to the same genus because of the of the cutting edges. BRS25/H/137, on the oth- subtriangular vertebral centrum with concave lat- er hand, shows teeth similar to those of the den- eral margins and the neural canal higher than the tary BRS25/H/135. In BRS25/H/137, the cutting cotyle in anterior view (Klembara 1981; Čerňanský edges are present, whereas they are not clearly vis- et al., 2019). Most of the Ophisaurus species known ible in BRS25/H/135 because of this specimen from european neogene localities are described being plunged in the matrix. based on parietals, but because of the very frag- Trunk vertebrae. The two vertebrae are medi- mentary nature of the parietals from Monticino, um sized and rather lightly built. The procoelous it is here preferred to identify them (and all other centrum is 4 mm long in BRS28/H/15 (Fig. 14h- Ophisaurus remains) only at generic level. It has to k) and exceeds 6 mm in BRS5bis/H/35; it is dis- be noted, however, that the anterior margin of the tinctly dorsoventrally compressed. in ventral view, interparietal shield not reaching the anterolateral the centrum is subtriangular, with oblique and dis- corner (i.e., the anterior end of the sulcus between tinctly concave lateral margins and a flat ventral it and the lateral shield is located far from the cor- surface (Fig. 14i). There is no precondylar constric- ner) is a feature of O. holeci, in contrast with an in- tion. Only the right synapophysis of BRS28/H/15 terparietal shield more laterally expanded or even is preserved (Fig. 14k): it is clearly dorsoventrally contacting the anterolateral corner in the other elongated. The neural arch is poorly preserved, but species for which the parietal is known (Klembara it appears low. in both specimens, there is one sin- 2015; Klembara & Rummel 2018): Ophisaurus fej- gle zygapophysis preserved, the right prezygapo- fari, Ophisaurus spinari, and Ophisaurus robustus. 318 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. fig. 15 - Pseudopus from Monticino Quarry - Pseudopus sp.: left frontal (BRS25/H/117) in dorsal (a), ventral (b) and lateral (c) views; right maxilla (BRS25/H/182) in medial view (d); fragment of tooth-bearing bone (BRS25/H/122) in lateral (e) and medial (f) views; trunk vertebra (BRS28/H/4) in dorsal (g), ventral (h), right lateral (i), anterior (j) and posterior (k) views. Scale bars = 1 mm. Genus Pseudopus Merrem, 1820 process. The ornamentation reaches the lateral mar- gin and is made up of low ridges and pits. because Pseudopus sp. of the smooth surface of the posterolateral process, fig. 15 the frontoparietal shield cannot be recognized, but the oblique groove between it and the frontal shield referred material: Two frontals (BRS25/H/117-118); is distinguishable. The margin of the frontal shield one fragmentary maxilla (BRS25/H/182); one fragment of tooth- bearing bone (BRS25/H/122); three trunk vertebrae (BRS24/H/9; marked by this groove, its posterolateral margin, BRS25/H/166; BRS28/H/4). is distinctly longer than its posteromedial margin. Moreover, the frontal shield reaches the posterior description margin, excluding therefore the interfrontal shield Frontal. BRS25/H/117 is a well-preserved left from the dermal cover of the frontal. The second frontal (Fig. 15a-c), lacking only the anterior end. It frontal, BRS25/H/118, is more poorly preserved, is roughly 8.5 mm long. in dorsal view, there is no being represented only by a moderately eroded pos- constriction at midlength and both the lateral and terior half. However, its morphology recalls that of medial margins are straight and roughly parallel (Fig. BRS25/H/117, except for the presence of a mod- 15a). In lateral view, the articulation surface with the erately developed dermal cover on the posterolat- prefrontal is moderately wide and long, reaching eral process also. BRS25/H/118 is roughly 5.3 mm midlength, whereas the one with the postfrontal is in length. smaller and shorter (Fig. 15c). The two surfaces are Maxilla. BRS25/H/182 is a fragmentary and not in contact, but they are not far from one anoth- very poorly preserved maxilla (Fig. 15d), represent- er. The posterolateral process is wide, pointed and ing the anterior portion of the bone. The anterior moderately long. A poorly developed lappet is pres- premaxillary process is clearly composed by an an- ent on its ventral surface. A small and triangular ar- terolateral and an anteromedial processes, of which ticulation surface is visible ventrally, medially to the the former is twice as robust as the latter. both pro- lappet. Though broken, the crista cranii is clearly ro- cesses are strongly eroded and the anteromedial one bust and well developed. A strong dermal ornamen- also misses its anterior end. They are separated by a tation covers almost completely the dorsal surface roughly U-shaped notch. The anterior concavity in of the bone (Fig. 15a), except for the posterolateral which the vomeronasal foramen opens is bordered Late Miocene vertebrates from the fissure fillings of Monticino Quarry 319 both laterally and medially by low ridges, but the found in Monticino from the other neogene Pseu- foramen is not clearly distinguishable because it is dopus species. first, both the ornamentation reach- covered by matrix. The medial ridge continues on ing the lateral margin for the entire length and the the dorsal surface of the anteromedial process in very long posterolateral margin of the frontal shield a moderately low lappet. The lateral surface of the compared to its posteromedial margin separate the specimen is smooth, carrying only four ventrolat- frontals BRS25/H/117 and 118 from Pseudopus ah- eral foramina. four tooth positions are visible and nikoviensis and Pseudopus laurillardi, in which the orna- three teeth are still present, though eroded. The mented cover touches the lateral margin only near teeth are subpleurodont, cylindrical and distinctly the anterior end and the posterolateral margin of robust; their crown is blunt and rounded. it seems the frontal shield is either as long as (in P. ahnikovien- that some striae are present on both the lingual and sis) or slightly longer (in P. laurillardi) than the pos- the labial surface of the teeth, but this cannot be teromedial one (Klembara et al. 2010; Klembara ascertained because of the erosion. The size of the 2012; Klembara & Rummel 2018). Pseudopus ah- teeth increases posteriorly. nikoviensis differs from the herein described remains Tooth-bearing bone. BRS25/H/122 is a very also because it does not show striae on the dentition poorly preserved fragment of an indeterminate ro- (Klembara 2012; Klembara & Rummel 2018). The bust and large-sized tooth-bearing bone (Fig. 15e- interfrontal shield, absent in the two frontals from f). The bone carries a single, well-preserved tooth, Monticino, is on the other hand present and dis- which is subpleurodont, robust, cylindrical and tinctly large in all Pseudopus species, except for Pseu- monocuspid. its crown is rather rounded and bears dopus apodus and Pseudopus confertus, in which it can be distinct striae on both the labial and lingual surfaces. either absent or narrow (Klembara et al. 2010, 2017; Trunk vertebrae. BRS24/H/9, BRS25/H/166, klembara 2012, 2015; klembara & rummel 2018; and BRS28/H/4 (Fig. 15g-k) have a medium size Villa & Delfino 2019a). The frontal referred to P. (centrum length goes from 4 mm to 6 mm), a robust- confertus (= Pseudopus rugosus) by Klembara (2015) ly-built aspect, and a dorsoventrally compressed, displays a dermal ornamentation which is distinctly procoelous centrum, provided with a smooth and different from the one of BRS25/H/117 and 118. flattened ventral surface and lacking a precondylar Therefore, the Pseudopus from the late Miocene of constriction. in ventral view, the centrum is sub- Monticino seems to be somehow different from triangular, since its lateral margins converge pos- all other species of the genus found in european teriorly (Fig. 15h). Such margins are straight, with neogene localities. nonetheless, given the scarce the only exception of BRS24/H/9, in which they amount of remains and the absence of other tax- appear slightly concave in ventral view. The neural onomically important skeletal elements (i.e., pariet- arch is low, with a neural canal that is lower in height als), either a possible identification as a new taxon than the anterior cotyle in anterior view (Fig. 15j). or a tentative attribution to one of the known spe- The neural spine was present, but it is broken in all cies are here avoided and the fossils are attributed specimens. when preserved, the zygapophyses are only to Pseudopus sp. suboval in dorsal view and dorsally tilted at an angle of about 45°. non-Anguis Anguinae indet. remarks. These specimens are clearly refer- fig. 16 able to Pseudopus because of the dermal ornamen- referred material: Two fragmentary maxillae tation reaching the lateral margin of the frontals, (BRS5tg2/H/8; BRS24/H/13); two pterygoids (BRS5tg2/H/3, 22); the cylindrical and robust teeth with a blunt crown, one ectopterygoid (BRS5tg2/H/23); six fragments of tooth-bearing the subtriangular vertebral centrum with straight or bones (BRS5tg2/H/5; BRS6/H/2; BRS25/H/140); 65 fragments slightly concave lateral margins, and the neural ca- of trunk vertebrae (BRS4/H/5; BRS5bis/H/2; BRS5tg2/H/20; BRS9/H/2; BRS15/H/5; BRS16/H/2; BRS19/H/7; BRS24/H/11; nal smaller than the cotyle (Klembara 1981, 2015; BRS25/H/1, 45, 111, 170-171; BRSNoNumber/H/16); 58 cau- Klembara et al. 2014, 2017; Čerňanský et al. 2019; dal vertebrae (BRS4/H/6; BRS5bis/H/3; BRS5tg2/H/21; Villa & Delfino 2019a). The specific identification, BRS16/H/3; BRS19/H/8; BRS24/H/12, 37; BRS25/H/46, 112- 114, 127, 139, 168, 172, 270, 285; BRSNoNumber/H/15); 2737 oste- on the other hand, is more complex, because of oderms (BRS1bis/H/1; BRS3/H/1; BRS4/H/1-4; BRS5bis/H/1; a number of features or combination of features BRS5tg2/H/27; BRS6/H/1; BRS8/H/5; BRS9/H/1; BRS15/H/4; that seems to distinguish the Pseudopus remains BRS16/H/1; BRS18/H/1; BRS19/H/6, 9; BRS24/H/8, 23-24; 320 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. presents a dorsomedially expanded posterior por- tion, the lateral surface of which bears a wide and deep articulation surface for the pterygoid flange of the pterygoid. The anterior half of the later- al surface, on the other hand, is covered by the very narrow articulation surface with the posterior process of the maxilla. The two articular surfaces contact each other. A small and anterolaterally di- rected flange marks ventrally the anterior articula- tion surface. The medial surface of the specimen is smooth. Tooth-bearing bones. The fragments of tooth-bearing bones, BRS5tg2/H/8 included, are very poorly preserved. each of them carries sub- pleurodont, large-sized and moderately preserved teeth. The preserved teeth seem to narrow dorsal- ly, suggesting a possible conical shape, but, since they lack the crown, it is not possible to say wheth- er the tip was pointed or not. no striae are visi- ble on the preserved portion of the teeth, but in fig. 16 - osteoderms of non-Anguis Anguinae indet. from Mon- BRS5tg2/H/8 and BRS24/H/13, they show the ticino Quarry - Osteoderm (BRS19/H/9) in external view anterior and posterior cutting edges visible in the (a). Osteoderm (BRS25/H/185) in external view (b). Osteo- above described Ophisaurus sp. BRS5tg2/H/8 is derm (BRS27/H/2) in external view (c). Scale bars = 1 mm. particularly robust and is 8 mm in length. Fragments of trunk vertebrae. Though the spec- BRS25/H/2, 19, 110, 120, 123-126, 167, 185-186, 233, 237, 246, imens are very poorly preserved, the preserved 275, 293; BRS27/H/1-3; BRS28/H/1-2, 16; BRSNoNumber/H/17; BRS?/H/8-12). portions of the fragments of trunk vertebrae are morphologically similar to those described above description as pertaining to either Ophisaurus or Pseudopus. Pterygoids. Pterygoids are medium-sized and However, their poor preservation hinders a confi- moderately robust. both specimens are very frag- dent recognition of the diagnostic features. Some mentary, preserving only the base of the quadrate of the fragmentary vertebrae are very large-sized process and part of the anterior half of the bone (e.g., the centrum of BRS25/H/171 is 10 mm (part of the palatine process in BRS5tg2/H/3 and long) and robustly built. part of the pterygoid flange in BRS5tg2/H/22). Caudal vertebrae. The caudal vertebrae present The palatine process shows a large number of a dorsoventrally compressed centrum and the bas- pterygoid teeth on the ventral surface, whereas a es of broken hemapophyses on the ventral surface. well-developed ridge runs along the lateral mar- Some of the best-preserved specimens are robust gin on the ventral surface of the pterygoid flange. and very large (the centrum length reaches 11 mm The preserved portion of the quadrate process in BRS25/H/114 and 172), whereas the other ones presents a wide and flattened basipterygoid fossa, are smaller. when preserved, the neural arch and which is marked ventrally by a distinctly developed the zygapophyses are similar to those of the above and roughly thumb-like basisphenoid process described trunk vertebrae. The autotomy plane is (sensu Klembara et al. 2010). Part of a deep fossa clearly lacking, except for BRS25/H/113. columellae is visible dorsally in BRS5tg2/H/22. Osteoderms. Osteoderms (Fig. 16), the preser- Ectopterygoid. BRS5tg2/H/23 is a fragmen- vational status of which varies from complete to tary, but very robust and large-sized ectopterygoid, very fragmentary, are mainly large- to very-large- which lacks almost completely the anterior end sized and robust, even if there are also smaller and the posteriormost portions of the posterior specimens. They usually have a subrectangular one. The bone has a medially concave shape and shape (Fig. 16a-b), but some more subtrapezoi- Late Miocene vertebrates from the fissure fillings of Monticino Quarry 321 dal or more suboval specimen is also present (e.g., description BRS27/H/2; Fig. 16c). A vermicular ornamen- Teeth. both teeth are medium-sized and lack tation is always present on the external surface their tip (Fig. 17a-d). BRS6/H/7 is also slightly (except for the smooth anterior gliding surface), eroded. They are canine-like in shape and distinctly whereas the inner one is smooth. The rectangular posteriorly curved. The implantation was subpleu- osteoderms bear also a distinct longitudinal keel in rodont and the tooth base shows a labially direct- the middle of the external surface. ed area made of spongy tissue (Fig. 17a, c). BRS- remarks. numerous vertebral fragments 5bis/H/13 is still connected to part of the original and osteoderms are assigned to indeterminate tooth-bearing bone (Fig. 17a). The teeth are medio- non-Anguis Anguinae. As far as the vertebrae are laterally compressed and present sharpened anteri- concerned, this is because of the large size and or and posterior margins, on which a light milling is overall similarity with those assigned to the two still visible in spite of the erosion. both the lingual anguines already identified in the locality. Identi- and the labial surfaces of BRS5bis/H/13 show a fication of the osteoderms, on the other hand, is number of dorsoventrally directed striae near the based on the large size, rectangular shape, and the tooth base (Fig. 17a-b). In BRS6/H/7, such striae external keel (Holman 1998; Delfino et al. 2011). are not visible on the labial surface (Fig.17c), but both the pterygoids and the ectopterygoid display this could be due to the preservational status. anguid features (e.g., osseous expansion marking Trunk vertebrae. The trunk vertebrae (Fig. ventrally the basipterygoid fossa, articular surface 17e-l) are procoelous, robust and large-sized with the maxilla limited to the lateral surface of (the centrum length of the best preserved one, the anterior half of the ectopterygoid; Villa & BRS25/H/138, is 10 mm). BRS25/H/138 (Fig. Delfino 2019a). The large size suggests that they 17g-l) is almost complete and roughly as long as it is can be attributed to a non-Anguis taxon, like the large. it has a wide and dorsoventrally compressed other similar-sized specimens from Monticino, but centrum, provided with a smooth and flattened since these skeletal elements are not clearly known ventral surface, strongly posteriorly convergent lat- for the neogene anguid species, they are not as- eral margins (giving it a triangular shape in ventral signed neither to Pseudopus nor to Ophisaurus. The view) and a distinct precondylar constriction (Fig. dentition of the fragmentary tooth-bearing bones 17k). The neural arch has a subtrapezoidal section is also comparable with that of a non-Anguis an- in anterior view and carries a neural spine, which guine (teeth not as spaced and curved as in Anguis; is, however, broken. dorsally, there is no pseu- Klembara et al. 2014; Villa & Delfino 2019a), but dozygosphene (Fig. 17g-h). Both the zygapophyses due to their poor preservation, it is not possible and the synapophyses are missing. In BRS6/H/4, to confidently assign them to a particular taxon. only the anterior end of the centrum and part of The only exceptions might be BRS5tg2/H/8 and its prezygapophyses are preserved. Size, robustness BRS24/H/13, the teeth of which are more similar and general morphology of this specimen is similar to those of the specimens attributed to Ophisaurus. to the other trunk vertebra. The prezygapophyses are slightly eroded, but they appear dorsoventrally inclined at about 45° in anterior view. The vertebral Family Varanidae Gray, 1827 (sensu Estes et al., fragments are very poorly preserved, but they are 1988) very large and robust. The size and the robustness Genus Varanus Merrem, 1820 of the specimens is comparable with those of the other remains attributable to Varanus, but some of Varanus sp. them are even larger (e.g, BRS25/H/85; Fig. 17e). fig. 17 BRS25/H/85, 129, and BRSNoNumber/H/1 show a wide and dorsoventrally compressed pos- referred material: Two teeth (BRS5bis/H/13; terior condyle, preceded by a strong precondylar BRS6/H/7); two trunk vertebrae (BRS6/H/4; BRS25/H/138); constriction (Fig. 17e-f). two fragments of trunk vertebrae (BRS25/H/85, 129); two cau- dal vertebrae (BRS25/H/130; BRS27/H/4); four fragments of Caudal vertebrae. BRS25/H/130 is the posteri- vertebrae (BRS6/H/5; BRSNoNumber/H/1-2); one humerus or portion of a moderately large-sized caudal ver- (BRS25/H/128). tebra, provided with a dorsoventrally compressed 322 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. fig. 17 - Varanids from Monticino Quarry - Varanus sp.: isolated tooth (BRS5bis/H/13) in lat- eral (a) and medial (b) views; isolated tooth (BRS6/H/7) in lateral (c) and medial (d) views; fragment of trunk vertebra (BRS25/H/85) in ventral view (e); fragment of trunk vertebra (BRS25/H/129) in ventral view (f); trunk verte- bra (BRS25/H/138) in anteri- or (g), dorsal (h), right lateral (i), left lateral (j), ventral (k) and posterior (l) views; right humerus (BRS25/H/128) in anterior view (m). Scale bars = 1 mm. centrum and a wide posterior cotyle. A precondy- caudal vertebrae and teeth can be confidently at- lar constriction is present and the pedestals for the tributed to this taxon too. The humerus is assigned chevron bone are visible on the ventral surface, near to Varanus because of its very large size, which is the cotyle. The postzygapophyses are suboval and comparable only with Varanus among all limbed liz- dorsally tilted at 45°. A broken, but strongly dorsal- ards identified in Monticino. The robustness of this ly projecting neural spine is present. The autotomy bone agrees with the hypothesis of robust-limbed plane is recognizable anteriorly. BRS27/H/4 is less varanids living in europe during the neogene sug- preserved, but has a comparable size and a similar gested by Georgalis et al. (2018), as also supported morphology, in particular for what the pedestals by the only neogene european varanid species for are concerned. This specimen is also split into two which the limbs are known, Varanus marathonensis portions, but it is not clear if the separation plane (based on the almost complete specimen described between them is a real autotomy plane or just the by Villa et al., 2018). Teeth with a milling are not result of breakage. common among european Varanus. They are Humerus. BRS25/H/128 is the poorly pre- known in few middle and late Miocene sites in cen- served distal epiphysis of a very large-sized and tral and eastern Europe (see Georgalis et al. 2019), very robust humerus (Fig. 17m). but not in material referred to Varanus mokrensis remarks. Together, the precondylar con- Ivanov et al., 2018. Contra Villa et al. (2018) and striction and the absence of the pseudozygosphene Georgalis et al. (2019), V. marathonensis indeed has are distinctive features of the trunk vertebrae of serrated teeth, at least in the almost complete indi- Varanus (Hoffstetter 1943; Estes 1983; Bailon vidual described by Villa et al. (2018) from Cerro 1991). Caudal vertebrae and teeth also display vara- de los Batallones (A.V., pers. obs.). The taxonomic nid features, such as the precondylar constriction significance of this feature is still unknown. How- in the former and the presence of long basal striae ever, new observations on the batallones specimen and spongy tissue (suggesting the presence of pli- seem to suggest that it is strongly influenced by cidentine) in the latter (Estes 1983; Bailon 1991; Pi- taphonomic processes and that it could be even anka et al. 2004; Kearney & Rieppel 2006). Because variable in different teeth of a single individual. of the comparable size and the fact that, among Varanidae, only Varanus has been reported from Suborder Amphisbaenia Gray, 1844 the European Neogene so far (Delfino et al. 2013; Amphisbaenia indet. Georgalis et al. 2017a; Villa & Delfino 2019b), both fig. 18 Late Miocene vertebrates from the fissure fillings of Monticino Quarry 323 remarks. Small trunk vertebrae without zygosphene and neural spine and provided with a dorsoventrally compressed centrum with a flattened ventral surface and subparallel lateral margins can be referred to indeterminate amphisbaenians (Estes 1983; Delfino 2003), but a more precise identifica- tion is not possible. Suborder serpentes linnaeus, 1758 Superfamily booidea Gray, 1825 family boidae Gray, 1825 Subfamily erycinae bonaparte, 1831 Genus Eryx daudin, 1803 Eryx cf. jaculus (Linnaeus, 1758) fig. 19 referred material: One cervical vertebra (BRS25/H/153); 35 trunk vertebrae (BRS2/H/4; BRS4/H/33; BRS5bis/H/15; BRS5tg2/H/26; BRS6/H/9; BRS9/H/10; BRS15/H/3; BRS19/H/4; BRS25/H/151, 156, 198; BRS27/H/17); two ar- ticulated trunk vertebrae (BRS25/H/252); 40 caudal vertebrae (BRS4/H/34; BRS4?/H/2; BRS5bis/H/16; BRS5tg2/H/24; Fig. 18 - Amphisbaenians from Monticino Quarry - Amphisbaenia BRS6/H/8, 10-11; BRS9/H/8-9; BRS24/H/1; BRS25/H/131, indet.: trunk vertebra (BRS19/H/1) in anterior (a), posterior 143, 155, 157-159; BRS28/H/3); two articulated caudal vertebrae (b), dorsal (c), ventral (d) and right lateral (e) views; fragment of trunk vertebra (BRS25/H/18) in right lateral view (f). (BRS5tg2/H/25); one fragment of vertebra (BRS25/H/152). Scale bars = 1 mm. description Trunk vertebrae. All trunk vertebrae (Fig. 19a-b) referred material: Four trunk vertebrae (BR19/H/1, 2; BRS25/H/17, 86); one fragment of trunk vertebra (BRS25/H/18). are massively built. The centrum is distinctly wider than long, with a very broad (but indistinct) haemal description. The trunk vertebrae are very keel (Fig. 19b). The prezygapophyseal processes, small, with the centrum length of the best pre- acute in shape, are very short but well visible in dor- served ones that is 2 mm in BR19/H/1, 2.2 mm sal view. The paradiapophyses are weakly divided in BRS25/H/86, and 2.4 mm in BR19/H/2. into para- and diapophyseal portions. The paracoty- BR19/H/1 (Fig. 18a-e) is almost complete, lack- lar foramina are absent. The basic measurements of ing only the posterior portion of the neural arch, the largest vertebra are as following: centrum length whereas the other specimens are more fragmentary. is 2.9 mm, whereas centrum width is 4.2 mm. The vertebral centrum is procoelous, dorsoventrally Caudal vertebrae. Most caudal vertebrae (Fig. compressed and has a flattened ventral surface. Its 19c-h) come from the posterior part of the caudal lateral margins are very slightly concave and roughly portion of the column. The vertebrae are provided parallel in ventral view (Fig. 18d). The synapophy- with a number of additional apophyses, among oth- ses are bulbous and massive (Fig. 18e-f). The prezy- ers pterapophyses, posterior extensions of prezyga- gapophyses are suboval in dorsal view and dorsally pophyses, posterior extensions of pleurapophyses, tilted at roughly 30° in anterior view. In BR19/H/1 and postzygapophyseal wings (the latter structures and BRS25/H/17, the dorsal surface of the neu- are mostly broken off). ral arch is smooth and there is no trace of a neu- remarks. The complex morphology of the ral spine in its preserved portion (Fig. 18c). A very caudal vertebrae is characteristic of the boid sub- small and rounded prezygapophyseal process is family erycinae and is not observed in any other visible in BRS25/H/18 (Fig. 18f) and 86, but it is groups of snakes (Szyndlar 1991a). Trunk vertebrae almost indistinct in other specimens preserving the with a wider than long centrum and short prezygap- prezygapophyses (i.e., BRS19/H/1 and 2). ophyseal processes are also assigned to this subfam- 324 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. tebrae of E. jaculus the postzygapophyseal wings are not fused anteriorly with the prezygapophyses (Szyndlar & Schleich 1994). The latter condition is also well visible in the herein-described vertebrae (despite the lack of distal portions of the wings). Taxon colubroides zaher et al., 2009 family Colubridae oppel, 1811 Genus Platyceps blyth, 1860 Platyceps planicarinatus (Bachmayer & Szyndlar, 1985) fig. 20 referred material: Three trunk vertebrae (BRS?/H/3, 17- 18). description. These vertebrae are elongat- ed and reach a centrum length of about 4.5 mm in the largest specimen. They are moderately pre- served. The centrum bears a wide and very low he- mal keel (Fig. 20e, g), which is marked laterally by very shallow (almost indistinguishable in the largest vertebrae) subcentral grooves. The grooves are lit- tle more developed in the smallest specimen. The posterior end of the keel is almost indistinct, but it is clearly separated from the posterior condyle. on the lateral surface of the neural arch, the parapo- physes are longer than the diapophyses (Fig. 20b). However, there is no parapophyseal process. In an- terior view, the neural canal is smaller than the cot- yle in the largest specimen, but as large as or even larger than it in smaller ones. when preserved, the zygapophyses go from slightly tilted dorsally to sub- horizontal, with a gradient toward more horizontal facets with increasing size. The prezygapophyseal processes are never preserved. The zygosphene has a straight or almost straight anterior margin, with Fig. 19 - Erycinae from Monticino Quarry - Eryx cf. jaculus: trunk two small lateral lobes (Fig. 20d). The neural spine is vertebra (BRS25/H/156) in left lateral (a) and ventral (b) always broken. in posterior view, the neural arch is views; caudal vertebra (BRS6/H/11) in dorsal (c) and ante- distinctly dorsoventrally depressed (Fig. 20f). rior (d) views; caudal vertebra (BRS25/H/158) in left lateral (e) and anterior (f) views; caudal vertebra (BRS25/H/155) remarks. The morphology of these trunk in right lateral (g) and anterior (h) views. Scale bars = 1 mm vertebrae perfectly recalls that of Nanus planicarina- tus Bachmayer & Szyndlar, 1985 in having (Bach- mayer & Szyndlar 1985; Szyndlar 1991a): small size; ily (Szyndlar 1991a). All these vertebrae, especially wide and low hemal keel; very shallow subcentral those from the caudal portion of the column, are grooves; absence of parapophyseal processes and referrable to the living european snake E. jaculus or thus not strongly developed paradiapophyses; and a closely related extinct species. Contrary to other straight to almost straight anterior margin of the zy- members of the genus, in the posterior caudal ver- gosphene, flanked by two small lateral lobes. They Late Miocene vertebrates from the fissure fillings of Monticino Quarry 325 fig. 20 - Colubrids from Montici- no Quarry - Platyceps plan- icarinatus: trunk vertebra (BRS?/H/17) in anterior (a), right lateral (b), poste- rior (c), dorsal (d) and ven- tral (e) views; trunk vertebra (BRS?/H/18) in posterior (f) and ventral (g) views. Scale bars = 1 mm. are therefore attributed to this species. Nanus plan- 21a). The zygosphene is well preserved. In dorsal icarinatus was later moved to Coluber by bachmayer view, it is wavy, with a well-developed median lobe & Szyndlar (1987) following the referral of a ba- and two narrower lateral lobes (Fig. 21c). The rest siparasphenoid from the type locality, which was of the vertebra is too damaged to be satisfactorily virtually identical to the same bone in snakes of the Coluber najadum/rubriceps group. In modern taxon- omy, however, the genus Coluber is restricted to a single species living in Central and north America, the eastern racer Coluber constrictor linnaeus, 1758, and the Coluber najadum/rubriceps group is now part of the genus Platyceps. Considering this, and pending a real revision of the extinct species also including a vertebral comparison with other colubroid snakes, we here follow Wallach et al. (2014) in referring it to Platyceps as P. planicarinatus in order to reflect Bach- mayer & Szyndlar’s (1987) original intentions. family dipsadidae bonaparte, 1838 Genus Paleoheterodon Holman, 1964 cf. Paleoheterodon arcuatus Rage & Holman, 1984 fig. 21 referred material: One trunk vertebra (BRS25/H/9); one fragment of trunk vertebra (BRS25/H/191). description. BRS25/H/9 is an elongated trunk vertebra, which is rather poorly preserved. it has a procoelous centrum provided with a very robust hemal keel (Fig. 21d). The keel is wide in ventral view and even slightly wider in its posteri- or half. The same portion is also ventrally flat. The posterior tip of the keel is somehow pointed and it is located distinctly away from the posterior con- dyle of the vertebra. Shallow but distinct subcen- tral grooves mark the keel laterally. low subcentral Fig. 21 - Dipsadids from Monticino Quarry - cf. Paleoheterodon arcu- atus: trunk vertebra (BRS25/H/9) in anterior (a), posterior ridges are also visible. in anterior view, the neural (b), dorsal (c), ventral (d) and left lateral (e) views. Scale bar canal appears not much wider than the cotyle (Fig. = 1 mm. 326 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. described, but the neural arch appears moderately depressed in posterior view in spite of the bad pres- ervation status (Fig. 21b). The centrum length is 3 mm. one other fragment of vertebra shares with BRS25/H/9 a similar hemal keel. remarks. This small snake displays a rather peculiar vertebral morphology, which appears rath- er similar to the one of the holotype trunk vertebra of P. arcuatus from the Miocene of France (Rage & Holman 1984; Szyndlar 2012). Significant affinities between the italian and the french fossils are found in: the small size; the elongated centrum; the wide hemal keel flanked by shallow subcentral grooves; the low subcentral ridges; the trilobed zygosphene; and the moderate depression of the neural arch in posterior view. Based on Rage & Holman’s (1984: fig. 3c) original figure, the holotype of P. arcuatus also shows a light widening of the posterior half of the keel and a separation between the tip of the latter and the condyle. These are features present in the italian material as well, thus possibly support- ing our attribution. nevertheless, we here consid- ered this identification as tentative due to the few remains available. family natricidae bonaparte, 1838 Genus Natrix laurenti, 1768 Natrix sp. fig. 22 referred material: One trunk vertebra (BRS15/H/1). description. This very small and rather damaged trunk vertebra is procoelous, anteropos- teriorly elongated, and lightly built. The centrum is subcylindrical, with an anterior cotyle which ap- pears slightly wider than the neural canal in anterior view (Fig. 22a). A well-developed hypapophysis is present on the ventral surface, but its distal portion is broken off. it does not develop a keel anteriorly. Fig. 22 - Natricines from Monticino Quarry - Natrix sp.: trunk ver- The preserved portion of the neural arch does not tebra (BRS15/H/1) in anterior (a), posterior (b), dorsal (c), appear compressed in posterior view. The neural ventral (d) and left lateral (e) views. Scale bar = 1 mm. spine is well preserved, being moderately high and overhanging both anteriorly and posteriorly (Fig. ence of hypapophysis, neural arch not compressed 22e). Other important morphological structures are posteriorly, and neural spine overhanging anteriorly; either missing or poorly preserved. Szyndlar 1984, 1991b). Based on the shape of its remarks. The vertebra, although strongly neural spine, which is rather high and provided with damaged, displays features characteristic of natricid prominent anterior and posterior overhangs, the (Natricinae sensu Szyndlar 1991b) snakes (i.e., pres- vertebra is referrable to the modern genus Natrix Late Miocene vertebrates from the fissure fillings of Monticino Quarry 327 fig. 23 - Psammophiids from Mon- ticino Quarry - Malpolon sp.: anterior trunk vertebra (BRS19/H/18) in dorsal (a), anterior (b), posterior (c), ventral (d), right lateral (e), and left lateral (f) views; anterior trunk vertebra (BRS25/H/194) in ven- tral (g) and right lateral (h) views; anterior trunk verte- bra (BRS25/H/188) in ven- tral view (i); anterior trunk vertebra (BRS25/H/193) in anterior (j), right lateral (k), left lateral (l), posterior (m), dorsal (n), and ventral (o) views; posterior trunk verte- bra (BRS25/H/189) in ven- tral (p) and right lateral (q) views; posterior trunk verte- bra (BRS4/H/35) in left lat- eral (r) and ventral (s) views; anterior trunk vertebra (BRS5bis/H/18) in ventral (t) and left lateral (u) views; fragment of trunk vertebra (BRS5bis/H/21) in dorsal view (v); fragment of trunk vertebra (BRS5bis/H/19) in dorsal view (w); fragment of trunk vertebra (BRS- 5bis/H/20) in dorsal view (x). Scale bars = 1 mm. (Szyndlar 1991b; Szyndlar & Schleich 1993; Augé & served specimens, most of which preserving only Rage 2000). A specific identification is not attempt- the centrum. both anterior and posterior trunk ed here due to the poor overall preservation of the vertebrae are represented. The largest ones reach specimen. a centrum length of about 7.5 mm. These verte- brae have a distinct ventral keel, the morphology family Psammophiidae dowling, 1967 of which is variable probably in relation with the Genus Malpolon fitzinger, 1826 position of the vertebra along the column. in those that can be interpreted as anterior trunk vertebrae Malpolon sp. (Fig. 23a-o, t-u), the keel is narrow, tall and sharp. It fig. 23 expands to various degrees at the posterior end. In the posterior portion of the trunk (Fig. 23p-s), on referred material: one hundred forty anterior trunk ver- the other hand, the keel is rather wide and distinct- tebrae (BRS1bis/H/8; BRS3/H/10; BRS4/H/32; BRS5bis/H/17, ly flattened, with a rather constant width. Among 31; BRS5tg2/H/28, 35; BRS6/H/12-13, 17; BRS9/H/11, 15; BRS15/H/14; BRS16/H/8; BRS18/H/13; BRS19/H/17-18, 21; vertebrae interpreted as coming from this portion BRS25/H/52, 88, 144, 150, 188, 192-194, 196, 222, 229, 231, 288; of the column, only BRS4/H/35 possesses a keel BRS28/H/6; BRS28?/H/6; BRSNoNumber/H/6, 20); nine poste- that expands towards the posterior end (Fig. 23s). rior trunk vertebrae (BRS4/H/35; BRS6/H/18; BRS25/H/29, 189, 195, 197; BRS28/H/19); five fragments of indeterminate trunk ver- The distance between the posterior end of the keel tebra (BRS5bis/H/18-21; BRS25/H/291). and the condyle is also subject to a certain varia- tion. Some vertebrae possess very large subcotylar description. Most of these vertebrae are tubercles, which are poorly developed in others represented by fragments or incompletely pre- (e.g., BRS25/H/222). Subcentral grooves are poor- 328 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. ly marked in anterior trunk vertebrae (Fig. 23d, g, i, family Viperidae oppel, 1811 o, t), but deeper in posterior ones (Fig. 23p, s). Some Genus Vipera laurenti, 1768 specimens preserve the paradiapophyses. These are well developed and the parapophyseal portion is Vipera s.l. (“Oriental vipers” group) roughly similar to the diapophyseal portion in size. fig. 24a-d There is no parapophyseal process. Prezygapophy- seal processes are preserved only in few fragments referred material: Thirty-nine trunk vertebrae (BRS5bis/H/14; BRS6/H/19; BRS9/H/16; BRS15/H/2; (e.g., BRS5bis/H/19-20; Fig. 23w-x) represented by BRS18/H/10; BRS19/H/3, 22; BRS25/H/220-221, 228, 249); eight the prezygapophyseal area, with BRS19/H/18 as fragments of trunk vertebra (BRS25/H/50, 76, 97, 284). the only exception preserving their base (Fig. 23a- d). When complete the processes are slender, very description. BRS19/H/3 is a very large ver- long (approximately as long as the facet) and point- tebra (Fig. 24a-b), with a centrum length of about ed. BRS19/H/18 preserves a strongly vaulted neu- 9.5 mm. it is procoelous, elongated but rather short, ral arch in posterior view (Fig. 23c). and robustly built. on the ventral surface of the remarks. These vertebrae are herein re- centrum, a distinct hypapophysis is present. only ferred to Malpolon because of the large size and the its basis is preserved. Anteriorly, the hypapophy- long, slender and pointed prezygapophyseal pro- sis is represented by a sharp keel, flanked by very cesses (Szyndlar 1984, 1991a). A thin and sharp (i.e., shallow subcentral grooves. The subcentral ridges unflattened) hemal keel was also considered diag- are low and robust. The right side of the anterior nostic for the genus by Szyndlar (1984, 1991a), but portion of the vertebra is broken off and so only our own observations on comparative material at the small left paracotylar foramen is still visible (Fig. our disposal confirmed that variation is present in 24b). Part of the anterior cotyle is also missing, but this feature, with vertebrae coming from the pos- it was clearly larger than the neural canal in origin. terior portion of the trunk that carry a flatter and on the left side, para- and diapophysis have roughly wider keel. The extant Malpolon monspessulanus (Her- the same size (Fig. 24a). A short and slender parap- mann, 1804) seems not to show the strongly devel- ophyseal process is present. both pre- and postzy- oped subcotylar tubercles visible in at least some gapophyses are missing. even though the right of the vertebrae from Monticino Quarry. This sug- portion is missing, the overall morphology of the gests that the late Miocene taxon might belong to zygosphene is still recognisable. it has a very slightly a different species. Vertebrae of the extinct Malpo- wavy anterior margin, with a very small central lobe. lon laurenti (Depéret, 1890) are very similar, or even on the left side, a small lateral lobe is visible. The identical, to those of M. monspessulanus according to neural spine is broken, as well as the entire posteri- Hoffstetter (1939), thus hinting at a similarly differ- or portion of the neural arch. The other vertebrae ent taxonomic identity. However, the largest verte- are less preserved, but share with BRS19/H/3 sim- bra described and figured by Depéret (1890), which ilar morphology, robustness, and size range. Some is from the cloacal area, has a shorter appearance fragmentary vertebrae preserve a larger portion of than the Malpolon from Monticino, even though we the hypapophysis (Fig. 24c-d), which is robust and only have trunk vertebrae at our disposal. Moreo- appears straight and well developed. ver, the french species seems to have shorter prezy- remarks. The presence of a possibly straight gapophyseal processes and a lower hemal keel ac- hypapophysis, rather short centrum, overall robust cording to Depéret’s (1890) descriptions. Malpolon morphology and very large size, and large cotyle mlynarskii Szyndlar, 1988 may possess developed and condyle allow attribution of these vertebrae to subcotylar tubercles (see Szyndlar 1988: fig. 2.3), a member of the oriental vipers species complex but the neural arch has straight sides in posterior (Szyndlar 1991b; Szyndlar & Rage 1999; Bailon et view, not as convex as in the Monticino vertebra for al. 2010). which this feature is known. The specific identity of the Malpolon from Monticino Quarry is therefore Vipera sp. 2 uncertain and we here refrain from identify it more Fig. 24e-j precisely than at genus level, also considering the general poor preservation of most of the vertebrae. referred material: One trunk vertebra (BRS25/H/232). Late Miocene vertebrates from the fissure fillings of Monticino Quarry 329 fig. 24 - Viperids from Monticino Quarry - Vipera s.l. (“Ori- ental vipers” group): trunk vertebra (BRS19/H/3) in left lateral (a) and anterior (b) views; fragment of trunk vertebra (BRS25/H/50) in left lateral view (c); frag- ment of trunk vertebra (BRS25/H/97) in left lateral view (d). Vipera sp. 2: trunk vertebra (BRS25/H/232) in anterior (e), left lateral (f), posterior (g), dorsal (h), right lateral (i) and ventral (j) views. Scale bars = 1 mm. description. BRS25/H/232 is a poorly arch (Szyndlar 1984, 1991b). The small size and the preserved, small trunk vertebra, missing most of low robustness hints for attribution to a small spe- the anterior end of the bone (except for a small cies and not to the large and robust oriental vipers. part of the anterior cotyle and the right parapo- distinction between small species of european physeal process), most of the hypapophysis, the viperids (namely, the Vipera aspis and Vipera berus neural spine and the left posterolateral corner of complexes; Szyndlar 1991b; Bailon et al. 2010) is, the neural arch. The overall shape of the vertebra however, difficult based on such a few and poorly is anteroposteriorly elongated, with a lightly-built preserved record. because of this, this specimen is construction. Ventrally, the base of the hypapoph- here attributed only to a second, smaller viper. ysis is preserved and it is possible to infer that the latter was well developed. laterally, the subcentral colubroides indet. grooves are shallow and the subcentral ridges are fig. 25 lowly prominent. based on what remains of the anterior end of the vertebra, it is possible to state referred material: Seven trunk vertebrae that the neural canal was either as large as or maybe (BRS25/H/133-134, 289; BRS27/H/11). even narrower than the anterior cotyle. The pre- served parapophyseal process is moderately slen- description. These few trunk vertebrae are der and pointed (Fig.24i). The interzygapophyseal too poorly preserved to be significantly described. ridges are well developed. The single zygapophysis They are elongated and provided with an hemal preserved, the right postzygapophysis, is subhori- keel. Some more information could be provided zontal. in posterior view, the neural arch is distinct- for BRS25/H/134 (Fig. 25). This very large and ro- ly roof-shaped (Fig. 24e, g). bustly-built trunk vertebra is moderately preserved, remarks. A second viperid species is repre- but it displays a strong degree of wearing caused by sented by this small and lightly built vertebra, pro- an indeterminate agent. This hinders a confident vided with hypapophysis and roof-shaped neural recognition of the original morphology of most 330 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Fig. 26 - Possible indeterminate snake from Monticino Quarry - ?Serpentes indet.: toot-bearing bone (BRS9/H/17) in medi- al (a) and lateral (b) views. Scale bar = 1 mm. ticino, suggest that another large colubroid snake was present in the assemblage, even though it is currently not possible to identify it. Fig. 25 - Indeterminate Colubroides from Monticino Quarry - Col- ubroides indet.: trunk vertebra (BRS25/H/134) in anterior Serpentes indet. (a), posterior (b), dorsal (c), ventral (d), left lateral (e) and right lateral (f) views. Scale bar = 1 mm. remarks. See the Supplementary informa- tion for a list of the remains identified only as Ser- pentes indet. of its structures, because it is not clear how much ?Serpentes indet. the preserved morphology is influenced by the fig. 26 wearing. nevertheless, it clearly has an elongated procoelous centrum, with a wide hemal keel (Fig. referred material: One tooth-bearing bone (BRS9/H/17). 25d). The hemal keel appears moderately distinct and flanked by rather shallow subcentral grooves, remarks. A small fragment of tooth-bear- even though, as mentioned before, this might be a ing bone could pertain to a snake. it is elongated result of the wearing as well. and carries three partial teeth, plus an empty space remarks. elongated trunk vertebrae with a that is as wide as a tooth and could therefore have hemal keel are a feature of “Colubrines” according hosted a fourth one even though no sign of such to Szyndlar (1984, 1991a), but this taxon includ- a tooth is present. Teeth were most likely conical, ed snakes that are currently referred to different but the tip is always missing and thus this cannot be families. because of their poor preservation, these confirmed. They are fused with the tooth-bearing vertebrae cannot be identified more precisely than bone in a subpleurodont way. as indeterminate Colubroides, even though the ab- sence of an hypapophysis prevents assignment to (Non-snake) Squamata indet. both natricids and viperids. The very large and ro- bust morphology of BRS25/H/134 and its wide remarks. See the Supplementary informa- hemal keel, a combination of feature that is not tion for a list of the remains identified only as in- seen in any other snake already recognised in Mon- determinate lizards. Late Miocene vertebrates from the fissure fillings of Monticino Quarry 331 Class Aves linnaeus 1758 order otidiformes wagler 1830 order galliformes Temminck, 1820 Family Otididae Rafinesque, 1815 family Phasianidae Vigors, 1825 otididae indet. Phasianidae indet. fig. 27c fig. 27a-b referred material: one distal fragment of a left tibiotarsus referred material: One proximal fragment of left scapula (BRS5b/Av/1). (BRS24/Av/3); one proximal left coracoid (BRS24/Av/1); one shaft of right coracoid (BRS25/Av/6); one distal right humerus (BRS19/ description. BRS5b/Av/1 is a portion of Av/2); one distal left ulna (BRS86/Av/1); one proximal right ulna a distal tibiotarsus with the condylus medialis, sul- (BRS24/Av/2); one fragment of proximal left carpometacarpus (BRS4/Av/1); one proximal right carpometacarpus (BRS15/Av/1); cus extensorius, pons supratendineus, and a short two complete left wing phalanges (BRS4/Av/2; BRS5b/Av/2). portion of shaft well preserved. The condylus lat- eralis is missing and the incisura intercondylaris is description. All the remains are extremely preserved only in its medial part. A tuberculum on fragmented, with the exception of the distal right the lateral distal side of the pons supratendineus is humerus BRS19/Av/1 (Fig. 27b) and the proximal clearly visible and well preserved. The distal edge left coracoid BRS24/Av/1 (Fig 27a). Despite the of the pons supratendineus is characterized by a preservation status, in all the remains it is possible ridge that runs for the abovementioned tubercu- to detect at least one morphological character of lum to the proximal side of the condylus medialis. the Phasianidae, as described by Göhlich & Mour- remarks. The tibiotarsus, even if incom- er-Chauviré (2005). plete, shows the general features of otididae, es- remarks. The fossil remains from Monticino pecially the presence of the tuberculum and the belong to a middle species of Phasianidae, similar in ridge on the distal edge of the pons supratendi- size to the extinct Palaeortyx gallica Milne-edwards, neus, which allow to recognize them from the oth- 1869 or Palaeocryptonyx donnezani depéret, 1892. The er Gruiformes. first species has been reported from several Oligo- This incomplete tibiotarsus belongs to a cene and Miocene European localities (Göhlich & small species of otididae, similar in size to the re- Mourer-Chauviré 2005); moreover, it has been also cent Tetrax tetrax linneaus, 1758. The fragmentary tentatively identified from the almost coeval site of status of the specimen does not allow to further Moncucco Torinese (Colombero et al. 2017). Pal- clarify its taxonomic attribution, even at a tentative aeocryptonyx donnezani is commonly found in Plio- level. nevertheless, the presence of a shaft por- cene and early Pleistocene sites in Mediterranean tion allows to exclude its attribution to the sim- Europe (Pavia et al. 2012; Bedetti & Pavia 2013). ilar-in-size genus Gryzaja zubareva, 1939, which The fragmentary status of the specimens described shows a very peculiar blade-shaped shaft of the herein prevents any further taxonomical attribution, tibiotarsus (Olson 1985a; Mlíkovský 2002; Mayr since the osteological differences among the various 2017). middle-sized Phasianidae can be detected, in some bustards have not been recorded before the cases with uncertainty, only with complete bones, as Miocene and the late Miocene fossils are quite pointed out by Göhlich & Mourer-Chauviré (2005) fragmentary and of doubtful affinities (Boev et and Louchart (2011). al. 2013; Mayr 2017). The earliest true Otididae The Phasianidae are among the most com- dates back to the Pliocene of eurasia and north mon avian taxa in the Paleogene and Neogene fossil Africa (Mlíkovský 2002; Mourer-Chauviré & Ger- record, with a large number of described fossil taxa, aads 2010), thereby implying that the record from the relationships among which are not always clar- Monticino Quarry represents the oldest Otididae ified (Mlíkovský 2002, Göhlich & Mourer-Chau- known so far. viré 2005). The oldest representatives of the crown group Phasianidae dates back to the early oligocene order strigiformes wagler, 1830 of France (Mayr 2017). After the early Oligocene, strigiformes indet. they became much more common in the old world fossil record (Mayr 2017), sometimes representing referred material: One ungueal phalanx with broken tip the dominant group in the bird assemblages. (BRS25/Av/2). 332 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. Fig. 27 - Birds from Monticino Quar- ry - Phasianidae indet.: prox- imal left coracoid (BRS24/ Av/1) in dorsal view (a); dis- tal right humerus (BRS19/ Av/2) in cranial view (b). otididae indet.: distal left tibiotarsus (BRS5b/Av/1) in cranial view (c). Accipitri- dae indet.: complete ungueal phalanx (BRS5/Av/1), side view (d). Scale bars = 2 mm. description. The ungueal phalanx shows a remarks. The characters described above rounded shape of the claw and a rounded and not allow to refer these ungueal phalanges to the Ac- well-protruding shape of the processus flexorius. cipitridae. The two phalanges from brS 25 and remarks. The abovementioned characters brS 85 belong to a small-sized form, comparable allow to refer this phalanx to the Strigiformes rather in size to the recent Accipiter nisus (Linneaus, 1758), than to the Accipitridae or to any other bird of prey. whereas that from brS 5 belongs to a middle-sized The inadequate preservation of the specimen and form, such as the recent Buteo buteo linneaus, 1758. the extreme similarities between the Tytonidae and The fossil record of Accipitridae dates back to the the Strigidae prevent a more detailed identification. oligocene, although some middle eocene speci- The fossil record of Strigiformes dates back mens have been tentatively referred to this family. to the Paleocene (Mayr 2017) and became very rich Unfortunately, the scarce number and fragmentary from the Eocene onward, with the extant genera status of the remains and their scattered distribu- well established during the late Cenozoic. tion made the phylogenetic affinities of the various fossil forms and their relationships with the crown order Accipitriformes Vieilliot 1816 group poorly constrained. family Accipitridae Vigors, 1824 Accipitridae indet. order Passeriformes linnaeus 1758 fig. 27d Passeriformes indet. referred material: One complete ungueal phalanx (BRS5/ referred material: One distal left humerus (BRS85/Av/2); Av/1); two ungueal phalanges with broken tip (BRS25/Av/1; two proximal left ulnae (BRS25/Av/3; BRS25/Av/4); one proximal BRS85/Av/1). right carpometacarpus (BRS25/Av/5); one distal right tarsometatar- sus (BRS9/Av/1). description. The three ungueal phalanges referred to the Accipitridae show a not-round- description and remarks. Although the ed shape of the claw and an edged and plantarely specimens show all the morphological characters of well-protruding processus flexorius. the Passeriformes and allow to exclude other small- Late Miocene vertebrates from the fissure fillings of Monticino Quarry 333 sized bird groups commonly reported in Miocene hyaena well represented during the late Miocene deposits, such as the Coliidae (Colombero et al. across the eurasiatic continent from Pakistan to 2017), their poor preservation prevent any further Greece (Rook et al. 1991). The most abundant car- considerations on their relationships within the pas- nivore at Monticino is the small hyaenid Plioviverrops serines. during the late Miocene, the Passeriformes faventinus (Torre 1989) (Fig. 2). Plioviverrops is a mon- already showed a considerable diversity and wide- goose-like, carnivorous/insectivorous/omnivorous spread global distribution, after their first appear- type (Werdelin & Solounias 1996), with a special- ance outside Australia during the Oligocene (Mayr ized dentition characterized by the increasing cusps 2017). height in molars and premolars. Plioviverrops faventinus is one of the most advanced and specialized species within the evolutionary lineage of the genus. The Class MAMMAliA linnaeus, 1758 carnivore guild is completed by the occurrence of a small felidae belonging to “Felis” ex gr. attica-christo- The Monticino Quarry site yielded a diverse li. This group of felids is documented in a number mammalian assemblage (39 species are represent- of localities across the Mio-Pliocene boundary in ed at the site). Most of the mammal fauna was de- the Old World (Rook et al. 1991). One of the rarest scribed in detail in a series of papers and, for this mammals documented within the rich mammals as- reason, herein we provide a synthesis of the mam- semblage is the aardvark (Tubulidentata) Orycteropus malian taxa identified to date at the site. cf. gaudryi. Albeit relatively uncommon, the genus Primates are represented by the colobine Orycteropus is documented across a wide geographic monkey Mesopithecus cf. pentelicus, documented by a range during Mio-Pliocene time at Mediterranean very limited number of remains. Mesopithecus is rel- latitudes. The Monticino fossils represent the west- atively common in the late Miocene to Pliocene of ernmost occurrence of Orycteropus in the Miocene eurasia, and, albeit rare, occurs in a number of latest of Eurasia (Rook & Masini 1994). Perissodactyla Miocene sites in Italy (Rook 1999; Alba et al. 2014). are represented by few remains of rhinoceros and Two fragmentary enamel fragments (a splinter of hipparionines. rhinoceros specimens have been as- a tusk and a molar fragment) are the only remains signed to “Dihoplus” megarhinus, a species tradition- documenting proboscidean occurrence. Albeit the ally considered as a typical Pliocene taxon but re- impossibility to attribute them to a specific taxon, cently recorded in several late Miocene (MN 12 and the molar fragment would suggest the presence of a MN 13) mammal assemblages of eastern Europe gomphotherid mastodont (Ferretti 2008). At Mon- (Pandolfi et al. 2015). The occurrence of “D.” me- ticino, Carnivora are recorded by five species, be- garhinus in the latest Messinian of Italy (in addition longing to four families. Eucyon monticinensis (Rook to Monticino, the species is recorded also at bac- 1992, 2009) is a small primitive dog, well represent- cinello V3 and Moncucco Torinese) confirms the ed at Monticino, among the oldest european repre- presence of this species during the late Miocene in sentatives of the family Canidae. The genus Eucyon europe and suggests a dispersal of eastern europe- is the earliest advanced member of the tribe Canini, an taxa in western Europe and Italy during the latest first occurring in the Clarendonian (late Miocene; Miocene (Pandolfi & Rook 2017). The small horse 12-9 Ma) of North America and expanding its geo- sample is attributable to a single species of hippar- graphic range into the old world by the end of the ionine horse (Rook & Bernor 2013) and is referred Miocene (Rook 2009). A single mandibular ramus to Hippotherium malpassii, a species named based on is assigned to Mellivora benfieldi (Rook et al. 1991); the rich hipparionine remains from the late Messin- this large mustelid represents the only occurrence ian of Baccinello V3 in southern Tuscany (Bernor of the taxon outside the African continent, where it et al. 2011). Arctiodactyla (like Perissodactyla) are is a relatively common carnivore in late Miocene to also underrepresented at Monticino with respect to Pliocene sites (Haile Selassie & Howell 2009). The coeval european sites. The best represented bovid Hyaenidae family is represented by two different from the site, and among the most common taxa species. A fragmentary skull and a few dental and in the assemblage, is the caprine antelope Oioceros postcranial remains represent Thalassyctis (Lycyaena) occidentalis (Pandolfi et al. 2021c). The species stands ex gr. chaerethis-macrostoma, a large bone-cracking as the recentmost representative of an evolutionary 334 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. lineage that originated in late Vallesian and Turolian sp. and cf. Miosorex pusilliformis (De Giuli 1989). Fi- of Greece and Turkey (Masini & Thomas 1989). nally, Chiroptera are relatively diverse in the assem- The species has been recently recorded in other late blage, with six different species (Kotsakis & Masini Miocene Italian localities (Pandolfi et al. 2021c) such 1989): the megadermid Megaderma cf. mediterraneum; as Verduno in Piedmont (Colombero et al. 2014) the two rhinolophids Rhinolophus cf. kowalskii and and Borro Strolla in Tuscany (Abbazzi et al., 2008a). Rhinolophus sp.; the vespertilionid Myotis cf. boyeri; Other scanty Bovidae remains allow the identifica- and the two hipposiderids Hipposideros (Syndesmotis) tion of four other bovids at Monticino: Antilopinae cf. vetus and Asellia cf. mariaetheresae. indet. (medium size), Bovidae indet. (small-size), cf. boselaphini, and cf. Parabos (Pandolfi et al. 2021c). A large sized Suidae is represented in the assemblage dIscussIon and referred to as Propotamochoerus provincialis, a spe- cies with a wide distribution in the late Miocene of diversity and palaeoenvironmental impli- Europe (Gallai & Rook 2011). Finally, a small num- cations of the Monticino non-Mammal verte- ber of brachydont teeth as well as a horn fragment brates testify the occurrence of small deers belonging to A remarkable diversity for the Monticino fis- Pliocervus sp. and to Procapreolus cf. loczyi, two quite sures palaeofauna was already pointed out for its widespread taxa in late Miocene sites across western mammalian component (Rook et al. 2015; see also europe, including other latest Miocene italian sites above). Our detailed study of the non-mammali- (Pandolfi et al. 2021b). an remains highlighted a high diversity also in the Small mammals are actually the best repre- palaeoherpetological assemblage, with at least 26 sented taxa at Monticino, due to the concentration (27 if the two fossils attributed to either indeter- of mostly dental remains resulting from tapho- minate scincids or possible scincids indeed belong nomic concentration. rodents represent the most to different species) taxa identified (Tab. 1). Birds abundant taxa among the assemblage (12 species and especially fish are, on the other hand, much documented by thousands of dental remains). Mu- less diverse, with just five and three taxa recognised rids are the most abundant family, represented by respectively. in the Monticino herpetofauna, rep- five species (De Giuli 1989): Stephanomys debruijni (a tiles are far more diversified than amphibians: 18 species belonging to a genus that spread in the late (or 19) taxa versus only eight. This might be related Miocene and early Pliocene); Centralomys benericettii, to a certain degree of aridity that characterized the a second murid described as a new species on the palaeoenvironment in the area when the sediments basis of material from Monticino (De Giuli 1989) accumulated in the fissures. This somehow dry en- and later on identified also in other Italian sites such vironment is suggested by the presence of reptile as Verduno in Piedmont (Colombero et al. 2014); taxa such as the agamines, eremiadine lacertids, Paraethomys anomalus; Occitanomys sp.; and Apodemus and sand boas of the genus Eryx (Speybroeck et al. cf. gudrunae. Two species belong to the family Cri- 2016), but also by otidid and phasianid birds. Water cetidae (Cricetus cf. barrierei and Ruscinomys cf. lasal- bodies were nevertheless present, as documented lei), two to Sciuridae (Atlantoxerus cf. rhodius and Hy- by fishes and supported by the fact that all amphib- lopetes sp.), and two to the Gliridae (Myomimus sp. ians and some of the identified reptiles (Mauremys, and Muscardinus sp.). The largest rodent present in the crocodylians, Natrix) are strictly linked to the the fauna is the porcupine (Masini & Rook 1993) availability of water. Remains of green toads (Bu- identified as Hystrix depereti (Weers & Rook 2003). fotes gr. viridis) suggest that at least some of these lagomorphs are relatively abundant, with the lep- water bodies might have been temporary. The other orid Trischizolagus sp. (Angelone et al. 2020) and the information given by the Monticino Quarry fossil ochotonid Prolagus sorbinii (Masini 1989; Angelone fauna hints for a complex environment surround- et al. 2020). Among insectivorans (Eulipotyphla), ing the fissures (most of the ecological data for the relatively common is the large-sized erinaceid Paras- taxa reported in the following part of the text are orex depereti (Masini et al. 2019), whereas other insec- from Rook et al., 2015, and Speybroeck et al., 2016). tivorans occurring in the assemblage are the small Pelobates, eremiadine lacertids, amphisbaenians, and erinaceid Mioechinus sp. and the soricids Neomyosorex Eryx indicate loose/sandy soils, but the same Eryx Late Miocene vertebrates from the fissure fillings of Monticino Quarry 335 1bis 1tris 2 3 4 4? 5 5bis 5tg2 6 8 9 15 16 18 19 24 25 27 28 28? 85 86 No Number ? Fish Aphanius crassicaudus X Cichlidae indet. ? Sciaenidae indet ? Amphibians Lissotriton sp. X Urodela indet. X X X Latonia sp. ? ? Discoglossoidea indet. ? Bufo gr. bufo cf. cf. Bufotes gr. viridis X X X X X X X X X Bufonidae indet. X X X X X X Hyla sp. cf. Pelobates sp. X X X Pelophylax sp. X X Anura indet. X X X X X X X X X X X X X X X X X Reptiles Testudo s.l. X X X Mauremys sp. X Testudines indet. X X X X X X X X X Crocodylia indet. X X Agaminae indet. X Eremiadini indet. X X X X X X X X Lacertidae indet. X X X X X X X X X X X X X X Scincidae indet X ? Ophisaurus sp. X X X X X X X X Pseudopus sp. X X X non-Anguis Anguinae indet. X X X X X X X X X X X X X X X X X X Varanus sp. X X X X X Amphisbaenia indet. X X Eryx cf. jaculus X X X X X X X X X X X X X Platyceps planicarinatus X cf. Paleoheterodon arcuatus X Natrix sp. X Malpolon sp. X X X X X X X X X X X X X X X Vipera s.l. (“Oriental vipers” group) X X X X X X X Vipera sp. 2 X Colubroides indet. X X Serpentes indet. X X X X X X X X X X X X X X X X X X X X X (non-snake) Squamata indet. X X X X X X X X X X X X X X X X Birds Phasianidae indet. X X X X X X X Otididae indet. X Strigiformes indet. X Accipitridae indet. X X X Passeriformes indet. X X X Tab. 1 - Distribution of the identified taxa in the different Monticino Quarry fissures. and the agamines support the presence of rocky primate remains found in the site (Rook et al. 2015). habitats as well. Similarly, taxa related to open hab- A more vegetated (and more humid) environment itats were found (Eremiadini, Eryx, otididae, Pha- dominated by ferns was also suggested by paly- sianidae, and Orycteropus among the mammals), but nological data (Bertolani Marchetti & Marzi 1988; the tentative identification of a tree frog, if correct, Rook et al. 2015), based on sediments that are how- may suggest that bushy areas were not completely ever slightly younger than those in which the fossil absent from the surroundings. Presence of woody faunas were collected. Adding to this, crocodylians environments could be supported by the very few and Varanus support a warm climate, which was al- 336 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. ready reported for Monticino based on the recovery ment was therefore similar in the Austrian locality of bats of the genus Megaderma (Rook et al. 2015). and Monticino Quarry, even though the latter could Together with the previously mentioned taxa have been less vegetated. Closer affinities to dry with specific ecological requirements and other taxa woodlands could maybe explain the higher abun- the ecology of which is too wide to be significant dance of P. planicarinatus remains in Kohfidisch. As for the palaeoenvironmental reconstruction (either far as P. arcuatus is concerned, this other species is because they are generalist like Bufo gr. bufo or Pe- reported from la Grive-Saint-Alban and Sansan lophylax or because the identification was possible (Rage & Holman 1984), adding to the new occur- only at a high taxonomic level), there are a few ex- rence in Monticino. based on the avian assemblage, tinct taxa in the Monticino non-mammalian fossil Cheneval (2000) reconstructed a wooded environ- assemblage that are worth discussing. first of all, ment for Sansan, an hypothesis later supported also the Monticino Malpolon sp. displays features that are by Pavia (2014). Woods were likely present at La not known in the extant Malpolon (i.e., the developed Grive as well, given that the locality yielded fossils subcotylar tubercles in at least some vertebrae) and of primates (Mein & Ginsburg 2002). Therefore, might therefore represent an extinct species of this P. arcuatus might have been either more linked to genus. living european representatives of Malpolon wooded environments (thus possibly explaining the are found in Mediterranean habitats, with a complex low number of remains identified in Monticino) or of different environmental features such as maquis, less specialized in terms of ecological requirements. sand dunes, humid areas, and rocky outcrops (Spey- Aphanius crassicaudus, in the end, lived in a broeck et al. 2016). The Afroasiatic Malpolon moilen- wide spectrum of different waters, from brackish sis (Reuss, 1834), on the other hand, seems to be to hyperaline (Gaudant 1979; Gaudant et al. 1988; related to even more arid habitats (Trape & Mané Caputo et al. 2009). Together with the rest of the 2006; Oudjani et al. 2016). Extinct members of the ichthyofauna, this indicates that estuarine or la- genus may have had different ecological require- goonal conditions were represented not far from ments of course, but its occurrence in Monticino the area where the Monticino karstic fissures orig- Quarry may suggest that at least the taxon inhab- inated. Still, the fact that only very few remains of iting this area during the Miocene shared a similar fish were found in the fissures suggests that main environment with living european congenerics. it water bodies were not in direct contact with them. has to be noted, however, that this may not hold fish remains may have been brought into the Mon- true for other extinct Malpolon as well; for example, ticino Quarry fissures by flooding events, as is also the palaeoenvironment of layna, the Pliocene type the case for some remains of large mammals show- locality of M. mlynarskii, is thought to have been ing erosion signs due to water transportation (Rook more wooded, with only occasional savanna and et al. 2015). The same floods could also have buried desertic areas (Domingo et al. 2013). Definition of animals that either used the fissures as dens or were specific ecological requirements is even more diffi- trapped in them. Articulated remains of animals cult for P. planicarinatus and P. arcuatus. The latter has that died in the fissures and were not subsequently no living close relatives to which its environmental disturbed by scavengers or other factors are indeed preferences could be correlated, whereas affinities present (Rook et al. 2015). On the other hand, the of the former with the genus Platyceps still has to accumulation of the very abundant small vertebrate be thoroughly confirmed through a real phyloge- remains, particularly those of the small mammals, netic analysis. Thus, possible ecological data can be has been interpreted as the result of the accumu- only presumed for both based on the palaeoenvi- lation of prey remains by birds of prey (De Giu- ronment reconstructed for the localities where they li et al., 1988; Rook et al., 2015). In contrast with were found, which involve just Monticino Quarry, what stated in previous studies (De Giuli et al. 1988; their respective type localities, and, only for P. arcu- Rook et al. 2015), the presence of mostly isolated atus, an additional locality. According to bachmayer mammal teeth and heavily fragmented postcranial & Szyndlar (1985) and Tempfer (2005), P. planicar- elements, both with extensive digestion tracks and inatus lived in a warm, Mediterranean climate with polished surfaces, suggests that the main accumula- relatively dry conditions, but with available woods tion agent was not a bird of prey, either nocturnal or and water bodies in Kohfidisch. The palaeoenviron- diurnal, but more likely a mammalian predator (An- Late Miocene vertebrates from the fissure fillings of Monticino Quarry 337 drews 1990) of small or middle size, extremely spe- went extirpated and reappeared in the continent in cialized in small ground vertebrates (thus including the MN 4 (Rage & Roček 2003). Both European amphibians and reptiles as well), rather than a more green toads, which are the first to appear in Miocene generalist species. Such hypothesis is in agreement Europe (MN 4/5 of Spain and France; Alférez Del- with the dietary habit of the most abundant preda- gado & Brea López 1981; Bailon & Hossini 1990; tor of the Monticino assemblage, the small hyaenid Rage & Roček 2003), and common toads may have Plioviverrops (cfr. Werdelin & Solounias 1996) as well an Asian origin (Stöck et al. 2006; Garcia-Porta et al. as with the recent dietary evaluation of the small ca- 2012; Syromyatnikova 2015). Bufonids were already nid Eucyon (Bartolini Lucenti & Rook 2020). Herpe- known in the european Messinian, even from local- tofaunal remains with possible signs of digestions ities very far from each other such as in Spain and are also present. This hypothesis is also supported Hungary. In Italy, late Messinian fossil green toads by the virtual absence of bird remains, less than 20 were recently reported from Piedmont (Colombero specimens among the several thousands of bones et al. 2014, 2017), but the common toad remains and teeth of the other taxa, which is at odds with from Monticino Quarry represent the oldest known other fissure-filling sites with bones accumulated occurrence of this bufonid group in the Apennine by birds of prey (Ballmann 1969, 1973, 1976; An- Peninsula. Sanchiz & Roček (1996) considered Hyla drews 1990; Abbazzi et al. 2004; Pavia 2004; Pavia as a possible Miocene Asian immigrant as well, & Mourer-Chauviré 2011; Bedetti & Pavia 2013), even though at least some of the extant Europe- where birds are always present, sometimes in large an populations are the result of more recent dis- numbers, including the predators responsible of the persal events from Africa likely mediated at least in bone accumulation. part by human introduction (Recuero et al. 2007). Other anurans found at Monticino Quarry such as Palaeobiogeographical insights Latonia, Pelobates, and Pelophylax were persistent in- Miocene european continental faunas were habitant of Europe since Paleogene times (Roček strongly affected by immigration from both Asia 1994; Rage & Roček 2003; Vasilyan 2018). At the and Africa, with the latter which came in contact moment, it is impossible to hypothesize the palae- with Europe as soon as the early Miocene (MN 3; obiogeographical affinities for the indeterminate Rögl 1999; Koufos et al. 2005). During the whole possible discoglossoid found in fissure BRS 25. The epoch, new animals entered the continent follow- urodele Lissotriton is widely distributed in europe ing different dispersal waves (among others, Rage & during the Miocene, following a first appearance in Roček 2003; Koufos et al. 2005; Rage 2013; Geor- the late Oligocene (Böhme 2008). This first occur- galis et al. 2016a, and reference therein). For am- rence is referred to the species Lissotriton rohrsi (Trit- phibians and reptiles, this lead to a modernization urus roehrsi in the original identification by Böhme, of the fauna, following the extinction of peculiar 2008), which was referred to Ommatotriton by Martín Paleogene taxa at the Oligocene/Miocene bounda- et al. (2012) but recently considered as better allo- ry or shortly after (e.g., amblyodont lacertid lizards; cated to Lissotriton by Georgalis et al. (2019). In any Villa & Delfino 2019b) and with the appearance of case, clear geographical affinities for this late Pale- close relatives of modern groups (e.g., relatives of ogene/early Neogene Lissotriton populations in eu- extant urodeles and anurans, green lizards; Bailon et rope are not well understood. al. 1988; Čerňanský 2010; Villa & Delfino 2019b). Among the reptiles, one of the most inter- It is therefore not surprising to find taxa with Af- esting taxa in palaeobiogeographical terms is the ro-Asian affinities in the late Miocene fauna from member of the eremiadini tribe. Currently, repre- the Monticino Quarry fissures, as well as taxa linked sentatives of this lacertid tribe are largely distribut- with extant groups in its herpetological compo- ed in arid habitats in Africa, but they are also found nent. This is the case, for example, for the toads, as in Asia and even in Europe (Sindaco & Jeremčen- both the green toads and, even if only tentatively ko 2008; Sillero et al. 2014; Speybroeck et al. 2016; identified at Monticino Quarry, the common toads Uetz et al. 2019). According to Arnold et al. (2007), belong to groups that are still present in europe. eremiadini originated probably in the middle Mio- bufonids were reported in the european Paleogene cene (around 16 Ma), in association with coloniza- herpetofauna preceding the eocene, but then they tion of Africa by lacertids passing through Arabia. 338 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. A similar, but slightly younger divergence time for to the presence of the Mid-Aegean Trench (Kyriazi the split between eremiadini and lacertini was also et al. 2008). Geographical barriers and recent colo- found by Guo et al. (2011). In contrast with this, nization appear therefore to exclude relationships Hipsley et al. (2009) obtained an Eocene split for between the Monticino Eremiadini and both extant the two tribes, thus pushing back their origin and european Eremias and Ophisops. Affinities with the consequently the possible colonization of Africa. Acanthodactylus lineage that originated the extant Eu- Unfortunately, this controversy cannot be solved by ropean populations of this genus cannot be ruled the fossil record at the moment, given that there are out on the same bases, on the other hand. fonseca no undisputed stem or crown eremiadini remains et al. (2009) gave a confidence interval comprised from either the Paleogene or the first half of the between 8.2 and 2.9 Ma for the split of the euro- Miocene. Succinilacerta succinea (Boulenger, 1917) pean lineage, which is not in contrast with a possi- from the baltic amber was originally referred to the ble diffusion of these lizards towards the Apennine eremiadini genus Nucras, but real affinities with this Peninsula resulting in their presence in Monticino and/or other African lacertids should be thorough- in the late Messinian. nevertheless, this cannot be ly assessed with new analyses of this taxon possi- confirmed at the moment, due to the absence of bly including study of its osteology using CT-scans. other Miocene eremiadini remains in europe and The oldest fossil occurrence of eremiadini is then the impossibility to further identify the fossils stud- that attributed to Eremias sp. by Rage (1976) from ied by us, and therefore other scenarios, such as a the Serravallian (around 13 Ma) of Beni Mellal, in different colonization event still occurring in the Morocco. later, Eremias was also reported from the late Miocene or even earlier from either Asia or Af- Zanclean of Kazakhstan (Vasilyan et al. 2017). Er- rica or the persistence of a still unknown european emias remains from the late Miocene of Mongolia lineage, should be considered equally probable at (Böhme 2007) were recently reallocated to indeter- the moment. Thus, the Monticino eremiadini are minate Eremiadini (Čerňanský & Augé 2019). Not of poor help in assessing the palaeobiogeographical considering the Monticino Quarry occurrence, the affinities of the Monticino fauna. Nevertheless, they oldest (and only pre-Quaternary) Eremiadini report represent the only occurrence of this lacertid tribe in europe is possibly a still unpublished and unver- known so far in the italian territory, both consider- ified Acanthodactylus sp. mentioned by böhme & ilg ing the fossil record (Villa & Delfino 2019b) and the (2003) from the Zanclean of La Gloria 4, in Spain current herpetofauna (Sindaco et al. 2005; Sillero et (Villa & Delfino 2019b). As far as the extant Eu- al. 2014; Speybroeck et al. 2016). They might there- ropean populations are concerned, molecular data fore have been part of a short-living offshoot of hint at a recolonization of the continent either from the tribe in the area. Asia or Africa, rather than at a derivation from an Similarly, a few biogeographical information ancient european stock. As a matter of fact, the can be retrieved from other reptilian taxa recov- european Acanthodactylus erythrurus (Schinz, 1833) ered in Monticino Quarry, such as the crocody- entered europe from Africa and separated from lians, anguids, most of the lizard taxa identified at its African relatives likely around the Miocene/Pli- suprageneric level, and the amphisbaenians. This is ocene boundary (Fonseca et al. 2009). Origin of because of the wide distribution these taxa had in Eremias arguta (Pallas, 1773) is also likely dated to Europe in the Neogene and/or the poorly refined a comparable time (Guo et al. 2011; Poyarkov et taxonomical identification of the herein-studied re- al. 2014), but it probably occurred in western Asia; mains, which hinders a deep understanding of their northwestern populations of this species (i.e., those biogeographical dynamics in the considered times. reaching Europe) show a low genetic differentia- As far as the crocodylians are concerned, a poten- tion, thus suggesting a recent expansion (Poyarkov tial referral to Crocodylus could result in another tax- et al. 2014). Ophisops elegans Ménétries, 1832, in the on with African affinities in the Monticino fauna. end, originated earlier in the late Miocene, probably neogene Crocodylus from europe were considered around the Tortonian/Messinian boundary, and col- African immigrants by Kotsakis et al. (2004), main- onized eastern Aegean islands (the only part of the ly on the basis of the old idea that the origins of European territory where it is found today) from such genus were related to that continent. More Asia, maybe being unable to move further west due recently, Oaks (2011) demonstrated that Crocodylus Late Miocene vertebrates from the fissure fillings of Monticino Quarry 339 originated in Australasia. Nevertheless, Delfino et the Apennine Peninsula is now a fact. Possible af- al. (2007) considered the only confident Crocodylus finities with both the middle Miocene Testudo antiqua occurrence from europe, from late Messinian to bronn, 1831 from Germany and the late Miocene lowermost Pliocene deposits in southern italy, to T. amiatae from Tuscany were highlighted by Cirilli possibly share morphological features with the ly- et al. (2016) for the remains studied by them, even bian Crocodylus checchiai Maccagno, 1947, thus poten- though warning that a reevaluation of the latter spe- tially supporting African relationships for the euro- cies is needed. Anyway, considering these affinities, pean representatives of the genus. in any case, only a possible central european origin for the italian (poorly diagnostic) teeth were found in Monticino Testudo should not be discounted. so far and so the presence of Crocodylus (and thus The Monticino snake fauna is dominated by of another Africa-related taxon) in this locality still descendants of Miocene immigrants. both orien- cannot be ascertained in absence of the discovery tal vipers and Eryx appeared in europe in the Mn of further, more taxonomically-significant fossil 4 (Szyndlar & Rage 1999; Rage 2013), at the same material. Asian affinities can, on the other hand, be time of the contact of the latter with Africa and supposed for Varanus, as this is thought to be the Asia. large viperids are known since the early Mi- case for both European extinct species of the ge- ocene in both Asia and Africa (Szyndlar & Rage nus currently considered as valid (Ivanov et al. 2018; 1999, 2000). In Asia, they are reported from the Villa et al. 2018). Mauremys sp. from Monticino has Zaisan Basin in Kazakhstan (Chkhikvadze 1985), an intermediate morphology of the shell elements but these remains were never properly described or between the italian M. campanii and the Greek M. ar- figured and so Szyndlar & Rage (2000) warranted istotelica. These two species were considered as sister caution on them. Rage (2003), on the other hand, taxa by Vlachos et al. (2019), but the Italian species published few vertebrae of a large viperid (“Ori- is found in older deposits: thus, a dispersal from ental viper” or Daboia) from Arrisdrift in Namibia, italy towards the balkans could be a more advisa- the age of which is equivalent with either the Mn ble scenario than the opposite. Chesi et al. (2009) 3 or the MN 4 (Szyndlar & Rage 2000). Based on even supported an eastward dispersal of Mauremys our current knowledge, therefore, it is not possible from central Mediterranean europe to Arabia and to favor either an Asian or an African origin for the Africa, even though Georgalis et al. (2020) recently neogene european large viperids, given that the proposed a possible alternative (or complementary) earliest extra-European remains either have uncer- way for the passage from europe to Africa of Mau- tain taxonomic affinities (the Asian ones) or their remys through the Sicily-Calabria arch. in this con- age is more or less comparable with the earliest eu- text, it is not clear whether the Monticino Quarry ropean fossils (the African ones). The oldest confi- taxon may represent part of a lineage that directly dent occurrences of Eryx from both europe and dispersed towards the Balkans (where then it even- Asia are rather similar in age: these are from the tually gave origin to M. aristotelica) or a descendant of MN 4-5 of France (Vieux-Collonges; Ivanov 2000) an independent italian stock deriving from M. cam- and Kazakhstan (Ayakoz; Malakhov 2005). Possi- panii. Undescribed Mauremys remains from the early ble Eryx are reported from older (MN 2-3) sites Pleistocene of southern Italy (Pirro Nord; Delfino in Spain and Germany (Szyndlar & Böhme 1993; & Bailon 2000; Delfino & Atzori 2013) mentioned Murelaga et al. 2002), thus possibly suggesting an by Georgalis et al. (2019) would be significant in older presence in europe compared to other con- better understanding the taxonomy of Apennine tinents (and maybe also that entrance of this genus Mauremys. in contrast with the pattern showed by in the continent was not related to the connection Mauremys, Chesi et al. (2009) reported Testudo in it- at MN 4). However, further material with a stronger aly only from the Messinian, thus considering it as taxonomical significance is needed before eventually a later immigrant. However, Testudo sp. was recently confirming this hypothesis. Nevertheless, Eryx does reported as part of the oldest known assemblage not appear in the African fossil record prior to the in the late Miocene of Tuscany (Cirilli et al. 2016), Serravallian (Rage 1976; Bailon et al. 2017) and so even older than assemblages with Mauremys from it is possible that this continent was colonized only the same area studied by Chesi et al. (2009), and later by sand boas. At the species level, the Mon- so an earlier appearance of Testudo populations in ticino remains represent the oldest occurrence of a 340 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. form morphologically related to the extant Eryx jac- congeneric with a middle to late Miocene north ulus. The same holds true for the Monticino Quar- American species, Paleoheterodon tiheni Holman, ry Malpolon, a genus that is otherwise only known 1964. Considering their similar appearance in the from Pliocene to Quaternary remains (Rage 1972; fossil record of both europe and north America, Holman 1998; Bailon 2000; Gleed-Owen 2001; Rage & Holman (1984) supposed an Asian ances- Bailon & Aouraghe 2002; Delfino 2004; Bailon try for both species, with the two different lineages & blain 2007; blain et al. 2007, 2008, 2011, 2013; then spreading towards europe and north Ameri- López-García et al. 2008, 2011; Blain 2009, 2015; ca respectively. Platyceps planicarinatus is additionally Jennings et al. 2009; Aouraghe et al. 2010; Pacheco known only from central Europe (Bachmayer & et al. 2011; laplana et al. 2013; daura et al. 2015; Szyndlar 1985, 1987) and the affinities of the in- Bailon et al. 2017; Piñero et al. 2017). Small vipers, determinate large Colubroides represented by the belonging to either the aspis or the berus species trunk vertebra BRS25/H/134 cannot be tracked. complexes, also entered Europe during the Mio- Natrix, on the other hand, is widely distributed in cene, even though in different moments compared the European fossil record, with its first representa- to both oriental vipers and each other. The Vipera tive found in the Oligocene (Natrix mlynarskii rage, aspis group is known in europe since the beginning 1988; see also Rage 2013). This wide distribution, of the neogene, whereas Vipera gr. berus appears both from a geographic and stratigraphic point of in MN 9 (Szyndlar & Rage 1999, 2000; Rage 2013). view, combined with the impossibility to identify Moreover, the latter is apparently confined to east- the vertebra recovered in the Monticino Quarry ern europe until Mn 17, when a westward spread below the genus level, hinders further comments is finally documented by fossils (Szyndlar & Rage on the affinities of the latter. 1999, 2000; Rage 2013). The origins of the Miocene when it comes to the other, non-herpeto- european populations of these two viper groups logical components of the fauna, the widely-stud- are not clear, in a similar manner to the oriental vi- ied Monticino mammals also show the results of per complex. The V. aspis group is known in Africa dispersals from the East (e.g., presence of Eucyon, in the Serravallian (Vipera maghrebiana Rage, 1976), “Dihoplus” megarhinus; Rook 2009; Pandolfi & Rook thus not really being of help for understanding the 2017) that likely occurred in the latest Miocene, origin of the older european occurrences. Chkh- as well as African taxa (M. benfieldi, which is sole- ikvadze (1985) mentioned the V. berus group from ly known in Monticino Quarry outside of Africa; the early Miocene of kazakhstan as well, but, as Rook et al. 1991). Issues related to the low precision for the oriental vipers he reported, there are no de- allowed for taxonomic identification and the wide scriptions or figures and so one should be cautious distribution (and mobility) arise again when consid- with this occurrence. The impossibility of assign- ering fish and birds from the quarry from a palae- ing the Monticino Quarry vertebra to either of the obiogeographic perspective. This holds particularly two small Vipera groups further complicates the true when birds are concerned. As for the fish, A. recognition of biogeographical affinities. Follow- crassicaudus is commonly found in sediments corre- ing a stratigraphic rationale, Vipera gr. aspis could lated with all the phases of the Messinian Salinity be a more reasonable choice, given the apparent Crisis in the Mediterranean area (e.g., Carnevale et absence of the other complex in this part of Eu- al. 2019). Sciaenids exhibit a wide stratigraphic and rope at the time. However, morphology should be geographic distribution during the Neogene (Ban- the first line of evidence when proposing affinities nikov et al. 2018). The putative cichlid recognized of fossils to specific taxa and so we leave the sce- herein based on a single tooth provides evidence nario open here to the alternative possibility of the of the African affinity of part of the fish assem- Monticino Vipera sp. 2 being more related to the V. blage. As discussed by Carnevale et al. (2003), these berus group, which was already present in eastern and other fish taxa of African origin occupied the Europe when the Monticino Quarry palaeofauna northern coasts and rivers of the Mediterranean was alive. The other snakes identified in the Mon- during the Messinian, with the species Oreochromis ticino Quarry assemblage can give even less palaeo- lorenzoi reported from the Messinian infraevaporitic biogeographical information. Paleoheterodon arcuatus shales of Gessoso-solfifera Formation. Even if it is is known from the french middle Miocene and is not possible to clearly correlate the isolated tooth Late Miocene vertebrates from the fissure fillings of Monticino Quarry 341 form brS 19 with O. lorenzoi, this specimen, even mammals evolved during the first part of the late though only tentatively referred to the Cichlidae, Miocene (Rook et al. 2006). Immigration events suggests the possible persistence of these fishes in with european incomers in the ozf are also testi- the same area even in younger times. fied during the period of existence of these island, though (Rook et al. 2006). Later, the endemic Tus- late Miocene terrestrial vertebrate faunas co-Sardinian mammal assemblage was substituted from italy by a non-endemic assemblage with European affin- The continental late Miocene of italy is doc- ities in the Messinian (Hürzeler & Engesser 1976; umented by various sites spread in different parts rook et al. 1999; Abbazzi 2001; benvenuti et al. of the Apennine Peninsula and even the major Ital- 2001; Bernor et al. 2001; Rook et al. 2006), the dis- ian islands. The best record we have is for the Mn persal of which passed through Piedmont and the 13: sites are located mainly in the north or central Monticino Quarry area (Rook et al. 2006; Colombe- north of the Peninsula, but Gravitelli in Sicily is ro et al. 2014). also considered as pertaining to this time span (The But what about non-mammalian vertebrates? NOW Community 2020). Older late Miocene sites do these parallel mammals when it comes to the are known from Tuscany, but also Sardinia (MN recognition of different faunal complexes in the 12) and Calabria (MN 11/12). Age limits from the Italian late Miocene? Amphibian and reptile as- complex of the deposits in Abruzzo and Apulia semblages are described from Piedmont (Cavallo (i.e., localities such as Scontrone and the Gargano et al. 1993; Colombero et al. 2014, 2017), Tuscany karstic complex) areas are less clearly defined, but (Delfino & Rook 2008; Chesi et al. 2009; Cirilli et al. they can be constrained in an interval covering the 2016), Monticino Quarry in Emilia Romagna (Kot- late Miocene and the early Pliocene (Masini et al. sakis 1989; Rook et al. 2015; this paper), Abruzzo 2010). Altogether, these localities record the pres- (Rustioni et al. 1992; Delfino & Rossi 2013), Sar- ence of three different mammal faunal complexes dinia (Abbazzi et al. 2008b; Georgalis et al. 2017b; in Italy during the considered period (Rook et al. Zoboli et al. 2019), and Calabria (Georgalis et al. 2006). First, in Calabria and Sicily there was a mam- 2020). Except for the crocodylian remains (Delfino malofaunal association showing clear affinities with et al. 2007), the palaeoherpetofauna from the Gar- the African continent (Ferretti et al. 2003; Marra et gano “Terre rosse” is still unpublished, but prelim- al. 2011; Marra 2018; Pandolfi et al. 2021a), but also inary comments can be made based on the content some taxa of European origin (Gallai & Rook 2006; of the PhD thesis by Delfino (2002) and our per- Rook et al. 2006). On the other hand, a strongly en- sonal observations. A certain affinity is evident be- demic insular fauna developed in the Apulian Plat- tween sites in Piedmont (Ciabòt Cagna, Moncucco, form (Freudenthal 1971; Rustioni et al. 1992; Rook and Verduno) and Monticino Quarry. If we consid- et al. 2006; Masini et al. 2010; Savorelli & Masini er the Piedmont area as a whole, more than half of 2016; Savorelli et al. 2016; Urciuoli et al. 2016; Pavia the taxa identified there are also present in Montici- et al. 2017). Adding to these, a third faunal complex no. only allocaudates, Chelotriton newts, gekkotans, was present in what is now the central north part green lizards, and scolecophidian snakes are missing of the Apennines, even though involving Sardinia in the latter site, but present in Piedmont. it has as well. This third complex is known as the “Ore- to be noted that the report of a palaeobatrachid in opithecus Zone Faunas” (OZF) after its most iconic Ciabòt Cagna (Cavallo et al. 1993) is likely errone- taxon, the hominoid Oreopithecus bambolii Gervais, ous: it is based on a partially preserved synsacrum, 1872. it is mostly known from Tuscany, in particu- the identification of which is rather doubtful due to lar the baccinello-Cinigiano basin in the province a subcylindrical vertebral centrum that is not dors- of Grosseto (among others, see Bernor et al. 2001; oventrally compressed as in palaeobatrachids (A.V., Rook et al. 2006; Cirilli et al. 2016; Rook 2016), but pers. obs.). Nine taxa (the large discoglossoid, the fiume Santo in Sardinia also yielded mammals in- common toad, the eremiadini, scincids, Pseudopus, dicative of the complex (Cordy & Ginesu 1994; P. planicarinatus, cf. P. arcuatus, Natrix, Malpolon, and Cordy et al. 1996; Abbazzi et al. 2008b; Casano- the small viper) out of 26 (or 27; about one third) vas-Vilar et al. 2011). The OZF record either a large of the taxa identified at Monticino Quarry are not island or a group of smaller islands where endemic known from Piedmont. However, unidentified 342 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. “colubrine” snakes are known in both Monticino demic nature of this snake, however, as well as a and Verduno, probably even representing more better comprehension of the composition of the than a single species (Colombero et al. 2017). Thus, herpetofauna of the ozf in Tuscany, need to await the purported absence in Piedmont of “colubrines” for more fossil material to be clearly confirmed recovered in Monticino Quarry might also be just a and understood. Similar to what written above for matter of missing identification. Almost all the taxa the geckos, Kotsakis (1989) also mentioned Anguis exclusive of Piedmont, on the other hand, were and Trionyx in his note on Monticino Quarry, but never found in southernmost late Miocene sites in again, our data did not confirm their presence. The the Apennines, thus probably representing groups same holds true for other taxa reported by Kot- the distribution of which did not reach those areas sakis (1989), such as the emydid turtle. From the in this time period. The only exception is represent- Messinian deposits of Tuscany, we only know a ed by geckos, which are present in Gargano with few chelonian taxa (Chesi et al. 2009): M. campanii, a peculiar, maybe endemic, form (Villa & Delfino Testudo, and Trionyx cf. pliopedemontanus. Again, Tes- 2019b). Geckos were reported, without a detailed tudo is present in Piedmont and Monticino Quarry description and figures, from Monticino Quarry by as well, the Mauremys from Monticino Quarry may Kotsakis (1989), but we were unable to confirm this be related to M. campanii, whereas trionychids are and the original material is not available for study currently missing from both Monticino Quarry and anymore. Anyway, an artifactual absence in our the Piedmont localities. Trionychids were also pres- sample of geckos due to a missing preservation of ent in Sardinia during the late Miocene (Georgalis their bones would not be surprising, considering the et al. 2017b), together with tomistomine crocody- overall rarity of fossils of these lizards (Daza et al. lians (Zoboli et al. 2019). Considering all of this, 2014). the Tusco-sardinian palaeoherpetological complex The palaeoherpetofaunal complex charac- could indeed differ from other central north italian terising the Monticino Quarry and Piedmont area assemblages, but additional fossils are needed prior is then somehow different from the pre-Messini- to have a significant confidence on this, especially an one reported from fosso della fittaia by Cirilli for the Messinian levels that are supposedly more et al. (2016) and Tuscany in general by Chesi et al. likely to share affinities with the latter. (2009). In contrast with the younger faunas from As discussed above, only preliminary com- Piedmont and romagna, fosso della fittaia yield- ments can be made about the herpetofauna from ed a small discoglossine (maybe not referable to the the Gargano area, as it is still unpublished and the larger Latonia, even though this hypothesis is just material is still currently under study. our personal based on the size of the few remains found and observations and the preliminary data available at thus should be considered carefully), Anguis, and the moment points out to a fauna composed of both a possible non-Eryx erycine snake, together with elements shared with Piedmont and emilia-romag- Testudo, non-Anguis anguines, indeterminate lizards, na (especially among the anurans, but possibly also and indeterminate “colubrines”. To these, Chesi et some lizards and snakes) and endemic taxa (e.g., the al. (2009) added Trionyx, which is not present in the small gecko briefly mentioned by Villa & Delfino, assemblage from Piedmont and romagna, and M. 2019b). At the current status of knowledge, no campanii, which on the other hand might be some- affinities seem to be shared with Tuscany, on the how related to at least the Mauremys from Monticino other hand. The few remains of crocodylians and (see above). Therefore, almost half of the amphib- chelonians from Scontrone (Rustioni et al. 1992) do ian and reptilian taxa from Tuscany in pre-Messin- not add very much to this picture. Similarly, only a ian late Miocene times are currently unknown in few chelonian taxa are known from Cessaniti in Cal- the Messinian of Piedmont and emilia romagna. abria. in addition to a marine turtle, this site yielded The erycine from fosso della fittaia is even more Trionyx pliocenicus fucini, 1912 and an indeterminate peculiar due to the absence of paracotylar foram- species of Mauremys (Georgalis et al. 2020). In the ina (Cirilli et al. 2016), a feature that distinguishes late Miocene of italy, other trionychid occurrences this taxon from both erycine genera known from are only known from Tuscany, both in the ozf and the Neogene of Europe (i.e., Eryx and Bransateryx; in Messinian levels. Chesi et al. (2009) reported the Szyndlar & Schleich 1993, 1994). The possible en- ozf trionychid as Trionyx unassigned to species and Late Miocene vertebrates from the fissure fillings of Monticino Quarry 343 the Messinian one as Trionyx cf. pliopedemontanus. Tri- Moncucco Torinese and Verduno; Colombero et al. onyx pliopedemontana Sacco, 1889 represents a nomen 2014, 2017) and Monticino Quarry. Phasianids, ac- dubium according to Georgalis & Joyce (2017) and cipitrids, and passeriforms were all shared by both at the moment all the Tuscan late Miocene material areas. Nevertheless, the impossibility to define more can be referred only to interminate trionychines fol- precisely the taxonomy of most of the remains lowing the same authors. nevertheless, the species found hinders a better understanding of these pos- identification of the trionychid from Cessaniti still sible affinities. Otidids, on the other hand, appears highlights affinities with Tuscany, given that the lost only in Monticino, the latter being their oldest oc- holotype of T. pliocenicus (which is the only speci- currence worldwide. A significantly different pic- men known of this species other than the Calabrian ture was represented by Gargano, where migratory remains) came from the Pliocene locality of Mapesi taxa were flanked by a number of endemic forms (or Malpessi), near Poggio alle Monache in Tuscany (Ballman 1973, 1976; Göhlich & Pavia 2008; Pavia (Georgalis & Joyce 2017). Whether the late Miocene 2013; Pavia et al. 2017). occurrences were conspecific with the Pliocene one (and thus with the Cessaniti specimens) or not is impossible to say. differently, Mauremys is present conclusIons in Piedmont, Emilia-Romagna, and Tuscany (both in the OZF and the younger Messinian fauna). It in a similar way to the frame depicted previ- is interesting to note that the chelonian fauna from ously by the study of mammal remains from the Cessaniti seems not to share clear affinities with the site, the Monticino Quarry proved extremely im- Abruzzo-Apulia area. portant also for other vertebrates after a detailed Coastal and freshwater fish assemblages in investigation of the non-mammal component of the post-evaporitic Messinian of italy share nu- the assemblage. This is certainly true for amphib- merous affinities throughout the Italian Peninsula ians and reptiles, given that with 26 (or 27) taxa (Carnevale et al. 2018, 2019). This is particularly ev- identified, it represents one of the most diverse ident between Tuscany and Piedmont, even though herpetofaunas known from the late Miocene. The it could just result from them being better sampled small amount of fossils recovered that can be at- than other sites (i.e., in Emilia Romagna, Marche, tributed to fish or birds hinders a similar achieve- and Abruzzo, for example). Differences are also ment for these two groups (only three and five taxa present, however. Clupeids are known from all identified, respectively), but important findings are above mentioned regions but Piedmont and emilia reported for them as well. As a matter of fact and Romagna, for example, whereas fossil remains of irrespective of the diversity, amphibians, reptiles, atherinids, bleniids, mugilids, and sparids were only and birds from Monticino Quarry include the old- found in Tuscany. Sciaenids, which are only tenta- est known occurrences of some taxa. On global tively identified in the Monticino Quarry fish as- terms, this is the case for the snakes Malpolon and semblage, are known from Piedmont, Tuscany, and Eryx cf. jaculus as well as for otidid birds, whereas Marche, but Piedmont and Tuscany do not share common toads represent the oldest known occur- sciaenid taxa at the generic level and each one of rence from the italian territory and eremiadini liz- the only two taxa identified in Marche is shared with ards are both the oldest finding of these reptiles in only one of the two former regions. Cichlids were europe and the only evidence of their presence, ei- not found in any of the italian late Messinian local- ther past or present, in Italy. None of the few taxa ities, except for (tentatively) the Monticino Quarry. of fish takes part in this list of oldest occurrences, They occupied the area already during the earliest but the possible cichlid tooth adds to only a few part of the Messinian Salinity Crisis event (Carnev- remains of this family in the northern part of the ale et al. 2003). The lack of fossils attributed to this Mediterranean, which furthermore were all found family from other late Miocene sites in italy may be in slightly older levels of few sites in central italy due to some sort of bias (e.g., sampling- or environ- (including Monticino Quarry). mentally-driven). overall, the non-mammal vertebrates from The bird assemblages also show some degree Monticino Quarry picture a complex palaeoenvi- of similarity, between fossil sites in Piedmont (e.g., ronmental scenario, somehow confirming what 344 Villa A., Carnevale G., Pavia M., Rook L., Sami M., Szyndlar Z. & Delfino M. was already suggested by the fossil mammals. Ac- ed by studies on mammals. Uncertainties and gaps cording to the ecological data of the newly-iden- are still present, though, and an increased effort in tified taxa, the Monticino Quarry area was warm the studies of these groups is strongly needed in and rather dry when the fossils deposited in the order to gain a better understanding of the italian fissures, even though with water bodies and a var- non-mammalian vertebrate communities between iegated set of environmental features (such as both the end of the Miocene and the beginning of the sandy soils and rocky outcrops, as well as vegetated Pliocene. areas) in the surroundings. The new data further confirm the hypothesys of a multiple origin of the Acknowledgements: we would like to acknowledge Antonio be- nericetti for the enthusiasm and passion that led him to discover the thanatocoenosis, with flooding events, bones ac- site and recover an amazing amount of fossil remains. Georgios l. cumulations created by predators, and (only ascer- Georgalis (Università degli Studi di Torino, Italy) helped by supplying tained from mammals, at the moment) animals that relevant literature. Fabrizio Coenda (Università degli Studi di Tori- died directly inside the fissures. no, Italy) screenwashed and selected part of the material described in this paper. We further thank Saverio Bartolini Lucenti (Università from a palaeobiogeographic point of view, degli Studi di Firenze, Italy) and Loredana Macaluso (Università degli the Monticino Quarry non-mammal assemblage Studi di Torino, Italy) for useful discussions about palaeoenviron- includes taxa with Afro-Asian affinities that dis- mental implications and the identification of some of the urodele remains respectively, as well as the editor, Silvio renesto, and the persed into europe during different moments in reviewers for useful comments on the manuscript. we also would the Miocene, together with others that already in- like to manifest our gratitude for the work done by the team and the habited the european continent. Considered in the contributors of MorphoSource, an online resource without which context of the Italian late Miocene land vertebrate some of the comparisons we made for this work would have not been possible. This project was supported by grants (ex-60% 2018 assemblages, the Monticino Quarry non-mammal and 2019) from the Università degli Studi di Torino and by Fondi di vertebrates share affinities mostly with Messinian Ateneo dell’Università di Firenze to LR (2017-2019). AV is supported faunas from Piedmont, even though some differ- by a Humboldt Research Fellowship provided by the Alexander von ences are indeed present (such as the presence of Humboldt Foundation. MD acknowledges support from General- itat de Catalunya (CERCA Program) and Spanish Agencia Estatal cichlid fish, some amphibians, some reptiles, and de Investigación (CGL2016-76431-P, AEI/FEDER, EU). This is the otidid birds). In a more general outline, this agrees publication number 355 of the Museo di Geologia e Paleontologia with the presence of a faunal complex involving collections at the Università degli Studi di Torino. the northern and central part of the italian Penin- RefeRences sula already recognised based on fossil mammals in the Messinian. Tuscany seems to be part of this Messinian faunal complex as well (again, as sug- Abbazzi L. (2001) - Cervidae and Moschidae (Mammalia, Ar- gested by mammals) at least as far as fish are con- tiodactyla) from the Baccinello V3 faunal assemblage cerned. The absence of sufficient data for the her- (Late Miocene, Late Turolian, Grosseto, Central Italy). Rivista Italiana di Paleontologia e Stratigrafia, 107: 10-123. petofauna and the avifauna hinders a confirmation Abbazzi l., Angelone C., Arca M., barisone G., bedetti C., of this for these two other group as well. older Delfino M., Kotsakis T., Marcolini F., Palombo M. 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