This work describes the Mousterian lithic artifacts found in the Avetrana karst filling (Apulia), among the oldest of southern Italy. A volcanoclastic layer found at the base of Bed 8, in the highest part of the stratigraphic succession, is also described. The compositional characteristics of the glasses of this volcanoclastic layer indicate a possible area of origin in the Mount Vulture volcano, and allow us to suggest that it is an aeolian fallout that occurred during or immediately after a late eruption of the Mount Vulture, probably within the 132 +/- 10 ka time span. The chronological result relating to the volcanoclastic sediments would place the underlying beds in the last Interglacial, MIS 5e, and the overlying Bed 8 at the onset of last Glacial period, MIS 5d. Moreover, it is in agreement with the paleoclimatic considerations based on the ecological and biogeographical features of mammal assemblages and it has a very important significance also for the biochronology of the Pleistocene. In fact, it allows us to confirm that some taxa found in the Avetrana succession, such as Crocidura gr. suaveolens, Arvicola italicus, the evolved form of Microtus (Terricola) gr. savii, and the modern form of Dama dama, occur for first time in Italy at the beginning of Late Pleistocene.

After more than sixty years since its discovery, the fossils from Monte Peglia (late early Biharian, Umbria) are reconsidered in their entirety. The small mammals of Monte Peglia upper and lower levels had been studied in the past, whereas the remains of large mammals of Monte Peglia lower level, stored in several Institutions of central Italy, are described here for the first time. The following taxa have been described: Hystrix refossa, Homotherium latidens, Panthera cf. P. gombaszoegensis, Felis cf. F. lunensis, Canis mosbachensis, Vulpes alopecoides, Meles meles, Pannonictis cf. P. nestii, Mustela palerminea, Ursus cf. U. etruscus; Macaca sylvanus florentinus, Equus altidens, Stephanorhinus cf. S. hundsheimensis, Sus sp., Capreolus sp., Axis eurygonos, Hemitragus cf. H. orientalis, Bison degiulii. Moreover, the list of small mammals of the lower level has been updated with the addition of three new small vertebrate taxa: Rana sp., Myotis sp. (large size), cf. Miniopterus sp. The study of the remains of large mammals of the lower layer indicate the survival of a number of taxa of latest Villafranchian age. If we accept the biochronological correlation of Monte Peglia with the Colle Curti local fauna, its age should be ~1.072 Ma. In this case, it is possible to pinpoint the accumulation of the lower level to the MIS 35/33, as the small mammals confirm the presence of a mixed environment with forested and open spaces and warm temperate climate. The accumulation of the upper level, characterized by taxa typical of open spaces and steppes and a cooler climate, probably occurred during MIS 34/32.

Revisions performed in the last 15 years added remarkable novelties to the taxonomy and biochronology of Italian fossil lagomorphs. Several new taxa have been erected basing on new materials and on the revision of old materials. This paper aims to illustrate the state of the art of such researches. The lagomorph diversity in the Italian Neogene and Pleistocene is quite high, accounting 9 ochotonids, 14 leporids, and 3 stem lagomorphs. Among the lagomorph taxa recorded in Italy, quite a high number are insular or continental endemics. The oldest Italian lagomorphs are the insular endemic Paludotona aff. minor, P. etruria and P. minor from the early-middle Turolian of the Tusco-Sardinia palaeobioprovince, and Prolagus apricenicus and P. imperialis from the late Turolian of the Abruzzi-Apulia palaeobioprovince. In the Italian peninsula, lagomorphs are known since the late Turolian (early Messinian) [...]

El depósito (cavidad kárstica) de La Grave, localidad cercana a la pequeña ciudad de Avetrana (Tarento, Italia meridional), ha dado lugar a numerosos fósiles de vertebrados. Los restos de grandes mamíferos han sido objeto de varios estudios. En este trabajo se examinan los restos de pequeños vertebrados y se identifican cuatro taxones de anfibios (Bufo bufo, Bufotes gr. B. viridis, Hyla gr. H. Arborea and Rana (s.l.) sp.), cuatro de reptiles (Testudo hermanni, Podarcis sp., Zamenis gr. Z. longissimus, Natrix natrix), y nueve de pequeños mamíferos (Erinaceus europaeus, Crocidura suaveolens, Arvicola italicus, Microtus (Terricola) savii, Microtus (Microtus) arva­lis, Apodemus gr. A. sylvaticus - A. flavicollis, Hystrix (Acanthion) vinogradovi, Oryctolagus cuniculus and Lepus corsicanus). Desde un punto de vista biocronológico, los datos sobre los vertebrados pequeños y grandes indican una edad entre el comienzo del Pleistoceno tardío (MIS 5e) y la parte central del MIS 3. Es probable que el estrato fosilífero más reciente (nivel 8) se haya depositado durante un período más frío en comparación con las capas anteriores. Los datos de pequeños vertebrados fósiles combinados con los que proceden de los grandes mamífe­ros y aves evidencian la presencia, cerca de la cavidad kárstica, de espacios abiertos (praderas) con charcos de agua, bordeados por zonas boscosas y, no muy lejos, la presencia de una costa rocosa.

The fossiliferous deposit (karst cavity) in La Grave, a locality near the small town of Avetrana (Taranto, southern Italy), has yielded numerous fossils of vertebrates. The remains of large mammals have been the subject of several studies. This paper examines the remains of small vertebrates and identifies four taxa of amphibians (Bufo bufo, Bufotes gr. B. viridis, Hyla gr. H. arborea and Rana (s.l.) sp.), four taxa of reptiles (Testudo hermanni, Podarcis sp., Zamenis gr. Z. longissimus, Natrix natrix), and nine taxa of small mammals (Erinaceus europaeus, Crocidura suaveolens, Arvicola italicus, Microtus (Terricola) savii, Microtus (Microtus) arvalis, Apodemus gr. A. sylvaticus - A. flavicollis, Hystrix (Acanthion) vinogradovi, Oryctolagus cuniculus and Lepus corsicanus). From a biochronological point of view, the data on small and large vertebrates indicate an age between the beginning of the Late Pleistocene (MIS 5e) and the central part of MIS 3. The most recent fossiliferous layer (bed 8) is likely to have been deposited during a cooler period when compared to the previous layers.The data from small fossil vertebrates combined with those emerging from the large mammals and birds evidence the presence, near the karstic cavity, of open spaces (prairies) with pools of water, bordered by wooded areas and, not far, the presence of a rocky coastline.

Fossil lagomorphs are very useful palaeogeographical indicators. In the last 15 years, several papers centered on fossil lagomorphs contributed to improve the Italian late Miocene-Quaternary palaeogeographical setting, solving palaeobiogeographical enigmas debated for decades, and providing new, challenging palaeogeographic data. The high number of endemic fossil lagomorphs of Italy is due in part to its complex tectonic history (insular endemisms), and in part to the semi-isolation and the physiography of the Peninsula (continental endemisms). In Italian lagomorphs, a direct causal relationship between dispersal and turnovers is not observed, except for the Toringian. Actually, species replacements are customarily due to archipelago effect (late Miocene), phyletic speciation (Pliocene of Sardinia and Italian mainland) or occur after the extinction of older congeneric species (early Pleistocene). (C) 2019 Academie des sciences. Published by Elsevier Masson SAS. All rights reserved.

Following a recent chronostratigraphic revision of 17 fossiliferous sites hosting assemblages constituting local faunas of the Aurelian Mammal Age for peninsular Italy, we provide a re-structured biochronological framework and discuss the current validity and significance of the middle Pleistocene Faunal Units (FU) for this region. Contrasting with the previous model of a wide faunal renewal during Marine Isotope Stage (MIS) 9 (similar to 330 ka), the First Occurrences (FO) of several species of the Torre in Pietra FU are significantly backdated and referred to the Fontana Ranuccio FU (530-400 ka). We show that the faunal renewal was more gradual and occurred earlier than previously assumed. Many taxa that are typical of the late Pleistocene register their FO in the Fontana Ranuccio FU, latest Galerian, which is characterized by the almost total disappearance of Villafranchian taxa and by the persistence of typical Galerian taxa such as Dama clactoniana, Bison schoetensacki and Ursus deningeri, and by the FO of Stephanorhinus kirchbergensis, S. hemitoechus, Hippopotamus amphibius, Cervus elaphus eostepahnoceros, Ursus spelaeus, Canis lupus, and Vulpes vulpes. The next Torre in Pietra FU is characterized only by the FO of Megaloceros giganteus and Mustela putorius. However, we observe that MIS 9 marks the actual moment when the faunal assemblages of this region are represented only by those taxa characterizing the late middle Pleistocene and late Pleistocene. For this reason, we propose to still consider the Torre in Pietra (lower levels) local fauna as a conventional boundary for the Galerian-Aurelian transition. Finally, we remark that the strong faunal renewal in MIS 13, with five FOs, coincides with the temperate climatic conditions due to the absence of marked glacial periods that could have favored the FO and the subsequent spread of these taxa. (C) 2019 Elsevier Masson SAS. All rights reserved.

The extreme rareness of Sardinian fossil sites older than Middle and Late Pleistocene makes the Monte Tuttavista karst complex (E Sardinia, Italy) very important. Remarkable lagomorph material, recovered from several fissure infillings of Monte Tuttavista referable to the Capo Figari/Orosei 1 and Orosei 2 faunal sub-complexes (early Pleistocene, similar to 2.1/1.9-1.1 Ma), allowed us to describe a new endemic insular leporid, Sardolagus obscurus n. gen. n. sp. The new taxon is characterized by a peculiar combination of an advanced p3 (Lepus-type) and a primitive P2 lacking deep flexa. The origin of such discrepancy, unprecedented among continental and insular endemic European leporids, is unclear. It could be the result of: (1) an independent evolution of p3 from an ancestor bearing the primitive P2/p3 (e.g., Alilepus, Hypolagus), or (2) a selective reversal morphocline from an Oryctolagus/Lepus-Yike leporine. The lack of data about the phylogenetic origin of the new taxon makes any inference about its possible arrival to Sardinia problematic. Crossing the European leporid records and evidence of migrations to Sardinia, we hypothesize three possible ages in which the ancestor of Sardolagus obscurus could have arrived in Sardinia, restricted to the late Miocene-early/late Pliocene (similar to 8-3.6 Ma). The phylogenetic relationship between Sardolagus obscurus n. gen. n. sp. and the oldest Sardinian leporid, recorded from Capo Mannu D1 and dated at the early/late Pliocene boundary (similar to 3.6 Ma), is unclear at present, however it is quite likely that they pertain to the same lineage.

Palaeoenvironmental information on Marine Isotope Stage 3 (MIS 3) coastal Latium is sparse, mainly based on studies of isolated faunal assemblages or long pollen records from lake sediments, often of insufficient resolution to aid in palaeoenvironmental reconstruction. This study describes in detail the Late Pleistocene faunal assemblage from layers SU11 and SU12 of Cava Muracci (Cisterna di Latina, central Italy), the first of which is a partially-preserved hyena den. The first multi-disciplinary palaeoenvironmental reconstruction of coastal Latium between 34-44 ka BP, a critical time span for the presence of the latest Neanderthals and the arrival of Anatomically Modern Humans (AMH), is provided combining palaeoecological inferences from a previous pollen study of hyena coprolites with the palaeontological study described here. The results indicate a temperate climate and a landscape characterised by the coexistence of at least three habitats within a short distance between the coastline and the inland mountains, suitable for a wide variety of species. (C) 2018 Academie des sciences. Published by Elsevier Masson SAS. All rights reserved.

The adaptation to a particular function could directly influence the morphological evolution of an anatomical structure as well as its rates. The humeral morphology of moles (subfamily Talpinae) is highly modified in response to intense burrowing and fully fossorial lifestyle. However, little is known of the evolutionary pathways that marked its diversification in the two highly fossorial moles tribes Talpini and Scalopini. We used two-dimensional landmark-based geometric morphometrics and comparative methods to understand which factors influenced the rates and patterns of the morphological evolution of the humerus in 53 extant and extinct species of the Talpini (22 extant plus 12 extinct) and Scalopini (six extant plus 13 extinct) tribes, for a total of 623 humeri. We first built a synthetic phylogeny of extinct and extant taxa of the subfamily Talpinae based on all the available information from known phylogenies, molecular data, and age ranges of fossil records. We tested for evolutionary allometry by means of multivariate regression of shape on size variables. Evolutionary allometric trajectories exhibited convergence of humeral shape between the two tribes, even when controlling for phylogeny, though a significant differences in the evolutionary rates was found between the two tribes. Talpini, unlike Scalopini, seem to have reached a robust fossorial morphology early during their evolution, and their shape disparity did not change, if it did not decrease, through time. Furthermore, the basal Geotrypus spp. clearly set apart from the other highly fossorial moles, exhibiting a significant acceleration of evolutionary shifts toward higher degree of fossorial adaptation. Our observations support the hypothesis that the evolution of allometry may reflect a biological demand (in this case functional) that constrains the rates of evolution of anatomical structures.

Mediterranean marginal basins recorded the Messinian salinity crisis (MSC) in various structural settings, including syn-rift, thrust-top, foredeep, and foreland basins. During the MSC, the Apennines were one of the mobile belts of the peri-Mediterranean chain. Starting from more hinterland areas, allochthonous units of both the northern and southern Apennines migrated toward the Adriatic foreland, developing allochthonous-top sedimentary basins. One of these basins, on top of the Molise allochthonous units (southern Apennines), recorded almost all of the main steps of the MSC. In a gypsum quarry district, the occurrence of conduits yielding flow-mobilized sediments, which cross-cut the Lower Evaporites, testifies to a fluid-migration event responsible for the formation of a brecciated mudgrade limestone buildup. This event can be connected with the Mediterranean drawdown responsible for the Messinian erosional surface (MES). The post-evaporitic marly succession that unconformably overlies the Lower Evaporites and the “Brecciated limestones” is characterized by the presence of Paratethyan molluscs and ostracods (Loxoconcha muelleri and Loxocorniculina djafarovi zones), and small mammals, among which is recorded the occurrence of Stephanomys debruijni. A disconformity within the post-evaporitic succession divides it into lower (p-ev1) and upper (p-ev2) Lago-Mare deposits. A further erosional surface separates the Messinian Lago-Mare sediments from fully marine lower Zanclean (MPl2 Zone) deposits. Whilst the unconformity separating the Lower Evaporites and the Lago-Mare deposits (although enhanced by a tectonic event) could be related to a late Messinian base level drop (MES1), as well as the disconformity between p-ev1 and p-ev2 (MES2), the younger angular unconformities affecting both the Lago-Mare deposits (p-ev2) and the fully marine Pliocene sediments might be related to two different phases of orogenic transport. Those events affected the Molise allochthonous units during their Late Miocene-Early Pliocene forelandward migration.

The systematics of Geotrypus is among the most debated within Talpidae, but the recent development of quantitative methods for shape analyses allows us to provide a thorough reconsideration of Geotrypus spp. In the present study, we perform a systematic revision of the species Geotrypus minor from the early Oligocene of Germany using two-dimensional geometric morphometrics on the humerus, and cladistic analyses using two different character matrices. Our results suggest a distinct generic allocation for this species based on its unique humeral shape. Cladistic analyses reveal that G. minor has closer phylogenetic relationships with urotrichine shrew-moles than with other Geotrypus species or highly fossorial moles. Quantitative methods applied in this study support qualitative observations and fully justify a new generic allocation. In light of these results, Tegulariscaptor gen. nov. is proposed to encompass the material previously assigned to G. minor. http://zoobank.org/urn:lsid:zoobank.org:pub:8A839F1E-0EC8-4799-B3AE-1A4E54A95F0E

Ceratopsians were herbivorous dinosaurs that dominated many of the terrestrial ecosystems in Asia and North America during the Cretaceous. The bizarre variety of skulls and lower jaw morphologies as well as the inferred ecological abundance of many species in this Glade indicate that Ceratopsia was a successful group. Here we analyzed 126 lower jaws from 50 ceratopsian species, using two-dimensional geometric morphometrics and finite element analysis to investigate differences in shape and structural performance of this part of the feeding apparatus across Ceratopsia. Morphological differences in lower jaws across ceratopsian clades are said to originate from feeding adaptation. Our results show that the stress (physical loadings modeled in response to biting) in lower jaws was quite similar between "basal" and "derived" taxa, whereas major differences among clades occur for stress values associated with the coronoid process. The basal ceratopsians Hualianceratops and Yinlong had a highly stressed and primitive lower jaw, indicating that those animals may have fed on relatively soft foliage and fruits. A similar condition was found for basal neoceratopsians and protoceratopsids. Psittacosaurids possessed a well-integrated and compact lower jaw able to withstand high stress, at the cost of having a highly stressed coronoid process. Leptoceratopsids were characterized by the opposite condition. Taxa such as Leptoceratops, Prenoceratops, Zhuchengceratops and Cerasinops appear to have had a comparatively efficient feeding apparatus. Ceratopsidae represents the Glade with the most efficient masticatory apparatus within Ceratopsia, even if the horizontal ramus of the lower jaw appears less able to withstand high levels of stress as compared with other ceratopsians. Additionally, we found the dentary and surangular angular complex co-evolved to generate a masticatory apparatus able to withstand high stress, particularly in Protoceratopsidae and Triceratopsini. The major phenotypic evolutionary rate and morphological changes occurred during the mid- to Late Cretaceous, when intense climate change and angiosperm diversification could have affected the evolution of ecological diversity and feeding biomechanics in Ceratopsia. (C) 2017 Elsevier Ltd. All rights reserved.

The lower molar of a large rodent collected near Terranuova Bracciolini in the Upper Valdarno Basin (Tuscany, Italy) and identified by Bosco (1899, 1900) as the beaver Trogontherium cuvieri Fischer, instead belongs to the porcupine Hystrix refossa Gervais. To date there is no evidence of the occurrence of the large beaver Trogontherium south of the Alps.

Organisms: Ceratopsians were herbivorous, beaked dinosaurs, ranging from 1 m to 9 m in body length, usually four-footed, and with a bony frill that extended backwards from the cranium over the nape of the neck. Known from Asia, Europe, and North America, they appeared in the Late Jurassic and persisted until the end of the Late Cretaceous.Questions: Which evolutionary processes drive the phenotypic evolution of skulls and lower jaws within Ceratopsia? What is the degree of morphological integration between the skull and lower jaw, and between the snout and frill among clades? Finally, are there any morphological evolution rate shifts across the ceratopsian phylogeny?Data: Photographs from 121 ceratopsian skulls and 122 lower jaws in lateral view, both from original photos and published pictures. Fifty-five ceratopsian species are represented in the sample.Methods: We investigated cranial and lower jaw shape changes across ceratopsians applying two-dimensional geometric morphometrics. We also investigated the morphological variation of the snout and the frill. Using phylogenetic generalized least squares regression, we estimated the degree of phylogenetic signal in size and shape data, as well as in the shape-size relationship. We performed phenotypic evolutionary rate analysis on shape data to describe phenotypic shifts across the phylogeny. Using a rarefied version of Escoufier's RV coefficient, we tested morphological integration between skulls and lower jaws, and between snouts and frills. Finally, we explored the potential link between cranial and frill shape evolution in ceratopsians and the radiation of angiosperms using a linear regression model.Results: Skull, snout, and frill shapes differ among clades (with the exception of leptoceratopsids and protoceratopsids). Lower jaws show distinct morphologies among groups. Size and shape changes are phylogenetically structured. The frill drives the morphological variation of the skull, co-varying much more with the lower jaw than with the snout. The frill appears to evolve to co-vary better with the lower jaw in the more morphologically derived clades than in basal ones. A significant linear relationship does exist between cranial shape and angiosperm occurrences, suggesting the hypothesis that the frill evolved in response to changes in dietary compositions associated with the turnover between gymnosperms and angiosperms during the Cretaceous. Significant negative shifts in evolutionary rates characterize skull, snout, frill, and lower jaw shapes, corresponding to nodes where psittacosaurids diverge from other taxa. In contrast, a significant positive shift in skull and snout shape rate of evolution characterizes the clade Ceratopsoidea.Conclusion: The frill is the main driving force in the overall cranial shape within Ceratopsia and evolved secondarily to better co-vary with the lower jaw to produce a more efficient masticatory apparatus. The changes in frill shape are correlated with the angiosperm diversification that occurred in the Cretaceous and thus correlated with changes in diet. Ceratopsians exhibit a slowdown in the phenotypic evolutionary rate in the Early Cretaceous and an acceleration of the phenotypic rate in the Late Cretaceous.

A multiproxy study of the lignite-bearing lacustrine sequence cropping out at Castel San Pietro (CSP) (Rieti, central Italy) provides new insights on the paleoenvironmental evolution of an extensional basin, located on the Tyrrhenian side of the Apennine range. The CSP fossiliferous levels reflect deposition in a stable lacustrine environment. Mammal remains collected from this locality during the past centuries include Stephanorhinus etruscus, Anancus arvernensis and the beaver Castor fiber. The co-existence of the Etruscan rhinoceros and the mastodont suggests a middle Villafranchian age (Montopoli or Coste San Giacomo Faunal Unit) for the lignite beds of CSP. New paleoflora findings are in agreement with an attribution to the Piacenzian or Gelasian age for these deposits whilst the presence of the ostracod Qinghaicypris cf. Q. riojensis suggests a time interval from Zanclean to Gelasian (between ~4.5 and ~1.8 Ma). Accordingly, the lignite beds of CSP most likely were deposited during the Gelasian (from ca. 2.5 to 1.8 Ma; from Montopoli to Coste San Giacomo FUs or possibly to Olivola FU) in forested area and warm and humid conditions.

Study of the tectonically active L'Aquila Basin offers new insights into both the creation of the extensional intermontane basins of the central Apennines of Italy and their tectono-sedimentary evolution through time. The combination of large mammal remains, ostracods, molluscs, Mousterian tools, and C-14 dating allows better definition of the onset and stratigraphic evolution of the L'Aquila Basin. Interpretation of a seismic reflection profile and well-log data allow evaluation of the subsurface setting of this sedimentary basin and its tectono-sedimentary evolution. The occurrence of a wedge-shaped seismic unit at the base of the basin sedimentary succession defines the first phase of basin fill during a late -Piacenzian-Gelasian synrift stage. Activity along the main fault of the extensional fault system responsible for the onset and subsequent development of the western sector of the L'Aquila Basin (L'Aquila-Scoppito subbasin) migrated from southwest to northeast, reaching the presently active Mount Pettino normal fault only in the late-Pleistocene-Holocene. The onset of sedimentation in the L'Aquila Basin was synchronous with the onset in the Tiberino Basin, and so the idea that these extensional intermontane basins become progressively younger from the Tyrrhenian toward the Adriatic side of the central Apennines is rejected. In the northern and central Apennines, only two major syndepositional extensional domains can be recognized: a late Miocene rifting area, which includes all the late Miocene extensional basins in Tuscany, and a late Pliocene to earliest Pleistocene rifting area, which possibly includes all the intermontane basins from the Tiberino Basin to the Sulmona Basin. The different time gaps between compressional and extensional deformation at any given locality in the central Apennines could indicate a partial decoupling of processes responsible for the migration of shortening and extension toward the foreland. Diachroneity between the eastward migration of shortening in the foreland and extension in the inner part of the orogen supports the notion that the central Apennines were created as a result of a partially decoupled collision zone. Study of the onset of the central Apennine extensional intermontane basins, together with their seismic activity, indicates that the central Apennine postorogenic extensional domain represents an archive of similar to 3 m.y. of continued crustal extension. These findings help to refine models of the long-term extensional rate of the central Apennines, and they provide a basis for more reliable seismotectonic models for one of the most seismically active sectors of the central Mediterranean area.

The faunal assemblage of Monteviale (Vicenza, northern Italy) represents a rare condition among the earliest Oligocene assemblages of south-eastern Europe at the ‘Grande-Coupure’. The lignitic fossiliferous strata lie above explosive basaltic breccias produced by a volcanic complex raised within a lagoon where the Calcareniti di Castelgomberto Formation (earliest Oligocene in age) was deposited. Systematic revision of the vertebrate remains from Monteviale reveals the presence of 15 taxa belonging to ?Butidae, Palaeobatrachidae, Trionychidae, Geoemydidae, Diplocynodontinae, Dugongidae, ?Pantolesta, Chiroptera, Rhinocerotidae, Anthracotheriidae and Palaeochoeridae. The fossiliferous deposit of Monteviale probably originated in a coastal lagoon characterized by salinity fluctuations, from brackish to fresh water, the latter evidenced by the presence of palaeobatrachid larvae. The terrestrial vertebrate assemblage indicates a humid forest environment with an age close to the Eocene–Oligocene boundary, lowermost Rupelian, MP21. Some of the mammal taxa (e.g. Epiaceratherium, Anthracotherium and ?Propalaeocherus) of Monteviale show a clear affinity with older (late Eocene) southern Asian species, suggesting a dispersal pattern across the several plates of south-eastern Europe and western Asia. By contrast, the herpetofauna (e.g. Trionyx, Bergouniouxchelys and Diplocynodon) suggests a closer relationship to European taxa.

Stephanorhinus etruscus is one of the most abundantly recorded and better known Eurasian Early Pleistocene rhinoceroses. Nevertheless, the first and last appearances of this species, as well as its paleogeographic distribution, are controversial and debated in literature. S. etruscus is documented since the latest Pliocene in Spain (Las Higueruelas), Italy (Montopoli and Castelnuovo di Barardenga), France (Perrier-Les Etouair) and Romania (Iaras-CarieraVeche). During the Early Pleistocene, S. etruscus occurred in several Spanish, French and Italian localities, as well as in The Netherlands (e.g., Tegelen), Germany (e.g., Thiede), Greece (e.g., Aivaliki) and Israel (e.g., Ubeidiya). The last appearance of S. etruscus in Eurasia is debatable. Etruscan rhino populations survived till the Jaramillo subchrone (around 1.1 Ma) in France (Bois-de-Riquet), Romania (Betfia XII) and Hungary (Osztramos 2 and 8), and close to the early-middle Pleistocene transition in Spain (Cueva Victoria, Huescar 1, Atapuerca TD4, TD6 and TD8), and Italy (Monte delle Piche). (C) 2017 Academie des sciences. Published by Elsevier Masson SAS. All rights reserved.