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Sus strozzii (Suidae, Mammalia) from the historical locality of Quercia (Early Pleistocene, Italy)
Abstract
Some remains of suids were recovered during the second half of the 1800s from Quercia (Tuscany, Italy), at a close distance to, but from a lower stratigraphic position than the better-known mammal fauna of Olivola. Although they were collected a long time ago, Quercia suids are described for the first time in this work. This sample represents one of the earliest occurrences (middle Villafranchian) of Sus strozzii, a large-sized suid present in Europe during the Early Pleistocene, but only abundant ∼2.0–1.8 Ma. A biometric comparison with selected samples of Pliocene to early Middle Pleistocene European suids is carried out, showing some dimensional changes in S. strozzii as well as differences between S. strozzii and other species. Quercia has been somehow eclipsed by other historical samples from Tuscany, namely the extensive collection of the Upper Valdarno and the diverse fauna of Olivola, but it is a different and important palaeontological locality. Apart from S. strozzii, the local fauna of Quercia-Vaccareccia includes Anancus arvernensis, Canis sp., Stephanorhinus etruscus, Leptobos etruscus, ‘Pseudodama’ sp., and Castor sp., representing one of the few mammal assemblages referable to the Faunal Unit of Coste San Giacomo (late middle Villafranchian, MNQ 17b), corresponding to ∼2.2–2.1 Ma.
... Among the many faunal changes (some of which are controversial for taxonomic reasons; Bellucci et al. [111]), we can mention again the disappearance of Bison (Eobison), whose last occurrence is at Pirro and probably also at Apollonia (Sorbelli et al. [117]), as well as the first occurrence of Praemegaceros verticornis and Bison menneri, and perhaps also of Megaloceros savini (Kahlke [110]; Bellucci et al. [111] and references therein). Also worth mentioning here is the appearance of the evolved form of Sus strozzi (Iannucci [74]), which is possibly a separate subspecies (considered by some to be an ancient form of the later Sus scrofa). ...
... ), Coste San Giacomo (2.2 Ma: Florindo et al.[73]) and Quercia (2.2-2.1 Ma: Iannucci[74], Italy), and Senèze, France (its base age is of ca. 2.2 and the youngest fossils ranged in age between 2.10 and 2.08 Ma after a new investigation in. ...
... comm.). It seems that these localities, which record the first secure mass invasion of the genus (Spassov[11,72,75]; Rook, Mart1F31nez-Navarro[4]; Iannucci[74]) have a very similar/close age and should be placed in the C. San Giacomo Unit. As it seems, two species entered practically simultaneously from the East to Europe (C. ...
The Villafranchian stage in the mammal fauna evolution in Eurasia (ca. 3.6/3.4 Ma—ca. 1.2 Ma) is associated with the beginning of the formation of the modern appearance of the mammal megafauna of today’s Palaearctic. The cooling and the aridification starting with the beginning of the Early Pleistocene gradually eliminated the quasi-tropical appearance of the Late Neogene landscapes and fauna of Europe. The time from the Mid-Piacenzian (ca. 3.3–3.0 Ma) to the end of the Early Pleistocene was a time of particularly intense dispersal of species, of faunal exchange between Eurasia and Africa, and of the entry of new mammals into Europe from the East. That is why the correlation of the biochronology of the Villafranchian fauna between Eastern and Western Europe is of particular interest. Accumulated data make possible a more precise correlation of these faunas today. A correlation of selected Eastern European localities with established faunal units and MNQ zones is made in the present work. Usually, the dispersal from Asia or from E. Europe to W. Europe is instantaneous from a geological point of view, but in a number of cases, reaching W. Europe happens later, or some species known to be from Eastern Europe do not reach Western Europe. The main driving forces of the faunal dispersals, which are the key bioevents in the faunal formation, are climate changes, which in turn, affect the environment. We can summarize the following more significant Villafranchian bioevents in Europe: the End Pliocene (Early Villafranchian: MNQ16) turnover related to the first appearance of a number of taxa, for example, felids, canids, proboscideans, and ungulates; the Quaternary beginning turnover. Correlated with this are the beginning of the Middle Villafranchian, which should be placed at about 2.6 Ma; the Coste San Giacomo faunal unit turnover (Senèze and Slivnitsa localities should be included here, and the FU itself, at the very beginning of the late Villafranchian (=MNQ18a)); the Pachycrocuta event at the very beginning of the Olivola FU; and the events related to the Late Villafranchian/Epivillafranchian bounfary.
... Comparing the Peyrolles MT IV with the homologous element in S. scrofa reveals a clear morphological resemblance, but I assign it to Sus sp., considering the limited knowledge on the postcranial anatomy of S. strozzii and the poor preservation of our specimen. In detail, I could not find an isolated and well-preserved MT IV of S. strozzii among the main collections that preserve remains of the species (see the institutions listed by Iannucci [13,68]). The MT IV is present in a mounted subadult skeleton from Senèze preserved in the Naturhistorisches Museum Basel (NMB) [69], which is the main source of information on the postcranial anatomy of S. strozzii and associated paleoecological inference [70]. ...
... Rather, it appears that suids were rare. Remains assigned to S. strozzii are already known from some localities older than 2 Ma, but none of them include more than a handful of specimens [13]-though Senèze yielded an almost complete skeleton [70]. To the list of early occurrences of S. strozzii discussed by Iannucci [13] (Saint Vallier, Valdeganga II, Coste San Giacomo, Quercia, and Vigna Nuova) should be added Pantalla, which has recently been recognized as older than previously assumed, at~2.2 ...
... Remains assigned to S. strozzii are already known from some localities older than 2 Ma, but none of them include more than a handful of specimens [13]-though Senèze yielded an almost complete skeleton [70]. To the list of early occurrences of S. strozzii discussed by Iannucci [13] (Saint Vallier, Valdeganga II, Coste San Giacomo, Quercia, and Vigna Nuova) should be added Pantalla, which has recently been recognized as older than previously assumed, at~2.2 Ma [91]. ...
It has been proposed that suids were absent from Europe during the post-Olduvai to pre-Jaramillo Early Pleistocene (from less than 1.8 to more than 1.2 Ma) and that their “re-appearance” in the late Early Pleistocene would mark the end of the late Villafranchian and the beginning of the Epivillafranchian. Arguments enumerated in favor of this “suid gap” are the lack of suid remains from extensively sampled fossil localities of this age and the high reproductive potential (r-strategy) of suids, which would translate in a high commonness of their remains in the fossil record. However, here it shown that while suids’ reproductive potential is certainly exceptional within artiodactyls, there is no direct relationship between the reproductive strategy and preservation rate of a taxon in the fossil record. In Early Pleistocene localities of Europe and adjoining areas, where suids are present in a fossil assemblage, they are always rare. In terms of number of occurrences (frequency), suids range from being moderately common (~2.0–1.8 Ma) to moderately rare (~1.1–1.0 Ma). Suid material is also described herein from Peyrolles (Issoire, France; reference locality for MNQ 19), a site dated at 1.47 Ma, providing direct evidence for the presence of suids within the purported “suid gap”. The case of suids underlines an important source of caveat in inferring faunal dynamics of the late Early Pleistocene of western Europe—including the dispersal of hominins—i.e., the unequal geographical distribution of the paleontological sites of post-Olduvai to pre-Jaramillo age. Indeed, Peyrolles is the only large mammal site in western Europe located outside the Iberian and Italian Peninsulas reliably dated around 1.5 Ma. In the post-Olduvai to pre-Jaramillo period, there is a paucity of radiometric estimates (or they have too coarse a resolution) and of paleomagnetic excursions detectable in continental deposits. Basically, for this time span, there is a high dependence on biochronological correlations, although, at the same time, these correlations are less reliable—because these are based on a few sites not covering the entire spectrum of environments present in Europe and the sites are not independently dated with methods that outperform biochronology—than those for other periods.
... Indeed, the main argument to reject the hypothesis of a late Villafranchian "suid gap" is the observation that suids are one of the rarest group of artiodactyls in the Pleistocene of Europe, and, hence, their absence at a given site might easily be fortuitous [1]. In the middle Villafranchian of southwestern Europe (Iberian Peninsula, France, and Italy), suids are known from a few localities that are dated or correlated to~2.4-2.1 Ma, namely, Valdeganga II in Spain, Saint Vallier and Senèze in France, and Coste San Giacomo, Pantalla, Quercia, and Vigna Nuova in Italy [1,12]. The most abundant of these samples is represented by a mere total of six isolated teeth recovered from Quercia, recently described [12]. ...
... In the middle Villafranchian of southwestern Europe (Iberian Peninsula, France, and Italy), suids are known from a few localities that are dated or correlated to~2.4-2.1 Ma, namely, Valdeganga II in Spain, Saint Vallier and Senèze in France, and Coste San Giacomo, Pantalla, Quercia, and Vigna Nuova in Italy [1,12]. The most abundant of these samples is represented by a mere total of six isolated teeth recovered from Quercia, recently described [12]. Martínez-Navarro et al. [8] ignored this publication, where the paucity of the other middle Villafranchian samples is also pointed out. ...
According to the “suid gap” hypothesis, suids (Suidae, Mammalia) would have been absent from Europe between 1.8 and 1.2 Ma. This hypothesis has been influential owing to its putative implications for biochronology and paleoecology—Sus scrofa (the modern wild boar) would appear 1.2 Ma in a period of climatic and environmental changes, coinciding with the beginning of the Epivillafranchian and the Early–Middle Pleistocene Transition, and hominins—the arrival of Homo in western Europe would precede the “return” of pigs. However, the “suid gap” hypothesis is based on the wrong premises that suids are abundantly represented in the European fossil record before and after the “suid gap”, that this purported abundance is linked to the suid reproductive potential, and that the paleontological sites dated within the 1.8–1.2 Ma interval yielded enough remains to exclude the notion that the absence of suid is merely accidental. In a recent paper, it is shown that all these assumptions are erroneous and suid material is described from Peyrolles (France), which is dated at 1.47 ± 0.01 Ma, hence perfectly “filling the suid gap”. Some proposers of the “suid gap” hypothesis have now provided comments to this recent paper, casting doubt on the age of Peyrolles and reiterating the arbitrary statement that suids were commonly recorded and abundantly represented in the Pleistocene of Europe. There is no valid reason to question the homogeneity of the faunal assemblage of Peyrolles, which is indeed a key locality for the mammal biochronology of Europe, being the reference for MNQ 19. Suids of comparable chronology have also been found in Krimni (Greece). Moreover, the “suid gap” proposers are basically advocating the use of an interval biozone based on the temporary absence of Sus strozzii—a species not common in the Pleistocene of Europe—providing no ecological explanation for this gap, apart from speculating it would be due to competition with Homo. The defense of the “suid gap” seems motivated by its use from the “suid gap” proposers as a biochronological argument to contend that the localities of Orce in Spain (Barranco León D, Fuente Nueva 3, and Venta Micena) are older than 1.2 Ma, when they postulated suids would “reappear” in the fossil record. However, since the “suid gap” hypothesis was primarily proposed based on the absence of suids from the Orce sites (and, secondarily, from other sites biochronologically correlated with the localities of Orce, like Pirro Nord in Italy), this represents an evident example of circular reasoning.
... However, despite the fact that reported specimens of S. arvernensis encompass the circum-Mediterranean area, including not only the European but also the African side, Anatolia, and even stretching to China, these are mainly represented by isolated dental material, an important portion of which is still undescribed (Athanassiou, 2018;Azzaroli, 1954Azzaroli, , 1975Berdondini, 1992;Cherin et al., 2018;Dal Piaz, 1930;Guérin & Tsoukala, 2013;Guérin et al., 1998;Hünermann, 1971Hünermann, , 1975Mazo & Torres, 1990;Montoya et al., 2006;Pickford & Obada, 2016;Samson et al., 1971; Van der Made & Moyà-Solà, 1989;Vekua, 1972). For instance, a comprehensive work on the most abundant sample of S. arvernensis, collected in the 1940s in the RDB Quarry in the area of Villafranca d' Asti (Italy), has only recently been published (Iannucci, 2024b), while only a portion of the craniodental material has already been described (Berdondini, 1992) or was considered as a comparative sample in previous studies (e.g., Cherin et al., 2018;Iannucci, 2023;Pickford & Obada, 2016). ...
... Apart from a tentative report from Bethlehem of Sus cf. strozzii (Hooijer, 1958), which likely dates to the Late Pliocene but is in need of revision (Iannucci, 2023), the earliest presence of the species might be represented by an isolated and fragmented dp4 recovered from Saint Vallier (Faure, 2004), while several findings are known from localities dated to or correlated with the following Coste San Giacomo Faunal Unit or MNQ 18 (or MNQ 17b) (Iannucci, 2024c). Saint Vallier is not directly dated, but based on the biochronological placement of its faunal unit between Roca-Neyra and Chilhac (which are dated), it should fall in the 2.5-2.3 ...
Sus arvernensis is a Pliocene species that occupies a key position in the evolution of suids (Suidae, Artiodactyla, Mammalia) in Eurasia, and besides, it is considered important for biochronological correlations and paleoecological inferences. However, our knowledge on S. arvernensis is largely based on fossil remains from southwestern Europe. Here, we present a revision of the Hungarian fossil record of S. arvernensis . Up to now, the species was known from only two localities of Hungary, Gödöllő (central Hungary) and Süttő (northwest Hungary), and the latter occurrence has even been questioned. After the comparison with other relevant samples of S. arvernensis , of the Early Pleistocene S. strozzii , and of the extant wild boar S. scrofa (motivated by previous attributions and the chronology of the localities), the presence of S. arvernensis from Gödöllő and Süttő is confirmed, and more material of the species is described from Beremend (southern Hungary) and Kisláng (western Hungary). Collectively, the results of the revision carried out herein reveal a relatively widespread distribution of S. arvernensis in Hungary, hence providing an important link from the eastern to western European fossil record of the species. The specimens from Gödöllő and Süttő are slightly larger than the other material of S. arvernensis from France and Italy included in the biometric comparison, although the paucity of the material precludes to evaluate whether these differences are significant and to relate them to a chronological and/or geographical context. The occurrence of S. arvernensis in the Hungarian localities considered in this work is a biochronological indication of an age older than at least 2.6 Ma, since the species is not recorded after the Pliocene–Pleistocene transition. This in agreement with the age estimates available so far for some of the localities or provides new insights. At Süttő, in particular, the identification of S. arvernensis reinforces the view that travertine deposition started already in the Pliocene.
... Villafranca d'Asti yielded the most abundant sample of S. arvernensis, being therefore of critical significance to broaden our knowledge on the species. Despite its importance, however, the Villafranca d'Asti sample has been only partly described in previous studies, or merely summarily considered while comparing other samples (e.g., Berdondini, 1992;Pickford & Obada, 2016;Cherin et al., 2018;Iannucci, 2023a). Berdondini (1992), in particular, carried out the only dedicated study focused on Villafranca d'Asti suids, but describing only part of S. P. I. the craniodental material. ...
... The material of S. arvernensis from Villafranca d'Asti represents the most abundant sample of the species known to date. The specific attribution of Villafranca d'Asti suids has never been questioned in previous studies (e.g., Azzaroli, 1975;Berdondini, 1992;Pickford & Obada, 2016;Cherin et al., 2018;Iannucci, 2023a), save for some ambiguity in the use of S. arvernensis or S. minor as the valid synonym (S. arvernensis has priority), but a comprehensive description of the material was missing. ...
The area of Villafranca d'Asti (Piedmont, northern Italy) is best known among mammalian paleontologists for having given its name to the Villafranchian, a term that today refers to a widely adopted biochronological unit, although it was initially proposed to denote a continental stage. Amongst the numerous remains of Pliocene large mammals recovered from the sedimentary basin of Villafranca d'Asti, those of suids (Artiodactyla: Suidae) represent the most abundant sample of Sus arvernensis. This species is the first member of the genus that includes the Eurasian wild boar, Sus scrofa, and many extinct and extant species of suids. Sus arvernensis is considered ancestral or close to the ancestors of several suid lineages, as well as being of relevance for biochronological correlations and paleoecological considerations. It is, indeed, one of the few species of large mammals apparently capable of taking advantage of the return to more humid conditions after the Miocene-Pliocene transition. Despite their importance, Villafranca d'Asti suids have been only partly described in previous studies, or merely summarily considered while comparing other samples. Here, combining the examination of this collection with the analysis of other material, I present a detailed study of the suid sample recovered from the Pliocene of Villafranca d'Asti, which provides new insights into this important sample and broadens our knowledge on the biometric and morphological variability of Sus arvernensis. A careful evaluation of curatorial and historical evidence supports the view of a coherent provenance of the fauna of Villafranca d'Asti from a single level of the RDB quarry (~3.2 Ma, Triversa Faunal Unit), which, although generally accepted in most previous studies, remained in-need of a proper justification. Moreover, preliminary taphonomic considerations based on the suid sample suggest that nearly entire carcasses were originally deposited.
... At the present time, Canis etruscus remains from CSG are the clear, earliest evidence of wolf-like canid occurrence from the Early Pleistocene in the Italian Peninsula, concurrently with those of Pantalla, now also dated to ca. 2.2 Ma [32], and maybe in Europe as a whole, pending the taxonomical attribution and the chronology of other Gelasian specimens. In Italy, for instance, several findings of Canis sp. are known from localities placed within the CSG Faunal Unit, namely, Fontana Acetosa [33], Montagnola Senese [34], Quercia [35], Torre Picchio [36] and Vigna Nuova [37]. The CSG specimens slightly pre-date the Italian findings from the Olivola (Val di Magra, Tuscany, ca. 2 Ma; Rook and Martínez-Navarro [38]) and Poggio Rosso (Upper Valdarno, Tuscany; ca. ...
Coste San Giacomo (CSG) represents a significant paleontological site to investigate the faunal and environmental changes that occurred in Mediterranean Europe during the Early Pleistocene. In this work, we described for the first time the Carnivoran assemblage. We ascribed the fossil remains to the following taxa: Ursus sp., Homotherium latidens, Canis etruscus, Pliocrocuta perrieri, Martellictis ardea and Vulpes alopecoides. Considering the value of the carnivoran taxa here identified, we discuss their particular biochronological significance, since the CSG site records the last occurrence of P. perrieri and the first occurrences of H. latidens, C. etruscus, M. ardea and V. alopecoides for the Italian Peninsula. These results will allow us to improve the data of the biochronological scheme of the Villafranchian European Land Mammal Age, recognizing the earliest dispersals and latest occurrences across Europe.
... It is also worth underlining that historical collections such as those housed at MUST constitute a mine of "dark" data (being every piece of information, from those directly associated with the fossils, such as their taxonomic identification, to those indirectly related to them, e.g., the chronology of the fossil-bearing deposits), especially significant when undertaking new fieldworks and excavations are not practicable or feasible, as in the urban area of Rome. Unfortunately, efforts aimed at revising historical collections are mainly limited to accompanying renewed research in old localities, aside from rediscoveries of specimens of peculiar importance (e.g., Bona, 2021;Fabbi et al., 2021;Mecozzi et al., 2022;see Iannucci et al., 2022b andIannucci, 2023, for discussion). To facilitate and promote such revisions, appropriate collection management is of paramount importance. ...
The urban area of Rome and its surroundings (the Roman Campaign, "Campagna Romana") yielded an exceptional amount of fossil remains of terrestrial large mammals, which are of prime relevance for biochronological correlations and for reconstructing the paleoenvironmental conditions in the Italian Peninsula of the Middle Pleistocene. Apart from the mere quantity, the scientific importance of the Roman fossil record rests on the geochronological constraints that allow to date or correlate several findings, as well as on the dense chronological cover of continental deposits, for the last ~0.8 Ma. On the other hand, the case of Rome is emblematic of the challenges posed by investigating historical fossil collections, for instance, the necessity of a thoughtful integration of historical documentation (e.g., geological maps, museum labels) and indirect geological information (e.g., borehole lithology, the correlation between obliterated fossil sites and extant outcrops), owing to the intense urbanization occurred especially since the 1800s. Fossil-rich deposits of Rome have supplied collections housed in the city's major museums, geosites, and universities, creating a valuable and unique paleontological heritage. Synthesis and perspective on these entwined aspects are provided herein, offering a geological and historical background alongside an overview of Middle Pleistocene mammal faunas of the area of Rome, with special emphasis on recent results that offer examples of-and how to deal with-different kinds of recoveries (from sporadic finds to systematic excavations), and ongoing work on the collection of the University Museum of Earth Sciences of Sapienza University of Rome (MUST). Reviewing the history of the MUST collection underlines the profound link between the history of the research on large mammal faunas of Rome and the history of the collection itself. The management of the paleontological heritage of Rome, consisting of thousands of remains spanning from isolated teeth to complete skeletons, is a crucial task for providing new data and support for research and dissemination, both of which are carried out at MUST accompanying traditional and yet fundamental efforts, such as cataloging and restoration, with the digital enhancement of the collection.
... Isolated dental remains of large-sized suids, which are generally referred to S. strozzii, are known from several middle Villafranchian sites, including Saint Vallier, Valdeganga II, Coste San Giacomo, Quercia, and Vigna Nuova, among others, and the locality of Senèze yielded an almost complete skeleton (Schaub, 1943;Mein et al., 1978;Azzaroli et al., 1988;Faure, 2004;Cherin et al., 2018;Iannucci et al., 2020;Azzarà et al., 2022;Iannucci, 2023). ...
The “Wolf event” is a prominent concept in large mammal biochronology of western Europe. It was defined in the 1980s as an intercontinental “dispersal event”, best represented by the arrival of a “primitive wolf”, Canis etruscus, but also involving other species. The Wolf event basically denoted the late Villafranchian faunal turnover, first expressed in Italy in the Olivola Faunal Unit. This event was also considered approximately coincident with the “old” Pliocene-Pleistocene boundary (~1.8 Ma, Gelasian-Calabrian transition), hence indicating important environmental changes and representing a relevant tool for correlation. Whilst it became soon clear that sporadic finds of modern canids (and, to some extent, other species) pre-dated the age assumed for the Wolf event, several authors continued to use the term and to associate it to the late Villafranchian, referring to the “massive expansion” of the species involved, rather than their first appearance in the European fossil record. Several bioevents traditionally included in the Wolf event and others that have been considered to occur later are today already documented in middle Villafranchian faunas. If a single species has to be taken as representative of the late Villafranchian faunal turnover, the best candidate based on the current evidence is the giant hyena Pachycrocuta brevirostris. The “Pachycrocuta brevirostris event” (as a replacement term for the Wolf event) can be characterized by the arrival in Europe of the giant hyena and Panthera gombaszoegensis, and the increase in the documentation of other species traditionally included in the Wolf event. However, this does not correspond to a sharp faunal turnover as traditionally envisioned for the Wolf event and it is possibly heralded in faunas slightly older than Olivola at ~2.0 Ma.
No remains of Sus strozzii were found during the new excavations in Senèze, but the nearly complete skeleton of a young male was found before World War II and is preserved in the Basel Natural History Museum. It was briefly studied and well illustrated by Azzaroli in 1954. The species is a large one, anatomically closer to the recent Southeast Asian long-snouted Sus verrucosus and S. barbatus than to the recent Eurasian Sus scrofa. Its cheek teeth are wider and lower, its molars less complicated and covered with thicker enamel than those of S. scrofa. Its limb bones are stockier than those of S. scrofa. S. strozzii is known in Western and Southeastern Europe and in the Middle East from the Early Villafranchian (biozone MNQ 16) onwards up to the early Middle Pleistocene (biozone MNQ 20). The presence of Sus strozzii is a clear indication of a rather forested area under a wet and probably warm climate.
The “Wolf event” is a prominent concept in large mammal biochronology of western Europe. It was defined in the 1980s as an intercontinental “dispersal event”, best represented by the arrival of a “primitive wolf”, Canis etruscus, but also involving other species. The Wolf event basically denoted the late Villafranchian faunal turnover, first expressed in Italy in the Olivola Faunal Unit. This event was also considered approximately coincident with the “old” Pliocene-Pleistocene boundary (~1.8 Ma, Gelasian-Calabrian transition), hence indicating important environmental changes and representing a relevant tool for correlation. Whilst it became soon clear that sporadic finds of modern canids (and, to some extent, other species) pre-dated the age assumed for the Wolf event, several authors continued to use the term and to associate it to the late Villafranchian, referring to the “massive expansion” of the species involved, rather than their first appearance in the European fossil record. Several bioevents traditionally included in the Wolf event and others that have been considered to occur later are today already documented in middle Villafranchian faunas. If a single species has to be taken as representative of the late Villafranchian faunal turnover, the best candidate based on the current evidence is the giant hyena Pachycrocuta brevirostris. The “Pachycrocuta brevirostris event” (as a replacement term for the Wolf event) can be characterized by the arrival in Europe of the giant hyena and Panthera gombaszoegensis, and the increase in the documentation of other species traditionally included in the Wolf event. However, this does not correspond to a sharp faunal turnover as traditionally envisioned for the Wolf event and it is possibly heralded in faunas slightly older than Olivola at ~2.0 Ma.
The dispersal of primitive elephantines and monodactyl equids in Eurasia has long been regarded as representative of a substantial turnover in mammal faunas, denoting the spread of open environments linked to the onset of cold and dry conditions in the Northern Hemisphere. During the 1980s, this event was named the “Elephant-Equus event” and it was correlated with the Gauss-Matuyama reversal, today corresponding to the Pliocene-Pleistocene transition and the beginning of the Quaternary, dated at ~2.6 Ma. Therefore, the Elephant-Equus event became a concept of prominent biochronological and paleoecological significance, especially in western Europe. Yet, uncertainties surrounding the taxonomy and chronology of early “elephant” and “Equus”, as well as conceptual differences in adopting (or understanding) the Elephant-Equus event as an intercontinental dispersal event or as a stratigraphic datum, engendered ambiguity and debate. Here, we provide a succinct review of the Elephant-Equus event, considering separately the available evidence on the “elephant” and the “Equus”. Elephantines dispersed out of Africa during the Pliocene (Piacenzian). Their earliest calibrated occurrences from eastern Europe date at ~3.2 Ma and they are usually referred to Mammuthus rumanus, although the allocation of several samples to this species is tentative. Available dating constraints for other localities do not resolve whether the dispersal of Mammuthus was synchronous across Eurasia, but this possibility cannot be ruled out. The spread of Mammuthus was part of an intercontinental faunal exchange between Africa and Eurasia that occurred during the Piacenzian, but in this scenario, Mammuthus is quite unique in being the only genus of African origin dispersing to western Eurasia. The arrival of monodactyl equids from North America coincides with the Pliocene-Pleistocene transition, with several occurrences dated or calibrated at ~2.6 Ma and no compelling evidence prior to this age. In Europe, early monodactyl equids are often aligned to Equus livenzovensis, but the material from the type locality of this species is chronologically time-averaged and taxonomically heterogeneous, and western European samples are seldom abundant or informative. Regardless, this does not diminish the biochronological significance of the “Equus event”. Indeed, while the term “Elephant-Equus event” should no longer be used, as the appearance of elephantines in the European fossil record markedly precedes that of monodactyl equids, we endorse the use of the “Equus event” as a valid alternative to refer to the intercontinental dispersal event that characterizes the middle Villafranchian faunal turnover, epitomized by but not limited to monodactyl equids.
Although the suid assemblages from the Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula) are reasonably well known, taxonomic studies devoted to them have lagged behind in recent decades. We describe the unpublished suid dentognathic remains from the earliest Vallesian (MN9) of Creu de Conill 20 (CCN20; 11.18 Ma), which represents the First Appearance Datum of hipparionin equids in western Europe. The sample includes 118 specimens, mostly isolated teeth, and a few maxillary and mandibular fragments. More than three-quarters of the specimens are assigned to the suine Propotamochoerus palaeochoerus, which is characteristic of MN9, albeit the described remains are slightly larger than average for the species. The rest of the sample belongs to a large tetraconodontine that is assigned to Parachleuastochoerus valentini, recorded elsewhere from MN7+8 to MN9, except for two specimens attributed to the small suid cf. Albanohyus sp. Our results support a synchronous dispersal of Hippotherium and P. palaeochoerus into Western Europe at ~ 11.2 Ma, suggesting that the latter is a suitable biochronological marker of the Vallesian. In turn, the remains of Pa. valentini refine our knowledge of the dental morphology of this species and strengthen the view that this species (unlike Conohyus doati and Conohyus melendezi) is not a junior synonym of Conohyus simorrensis. The lack of Listriodon splendens and Versoporcus sp. from CCN20, together with the scarcity of Albanohyus, contrasts with their abundance in the roughly coeval site of Castell de Barberà, hinting at local paleoenvironmental differences.
We redescribe and revise the taxonomic attribution of a lost hyena mandible recovered from Paciano (Umbria, Italy), originally reported in the early 1900s, by comparing it with relevant samples of Pliocene, Pleistocene, and extant species. The mandible of the Paciano hyena was initially attributed to Hyaena striata (= Hyaena hyaena) and subsequently listed as a record of the giant hyena Pachycrocuta brevirostris, but is here assigned to another “bone-cracking” hyena, the Pliocene-Early Pleistocene Pliocrocuta perrieri. The Paciano hyena contributes to the discussion on the relationships and turnover between Pl. perrieri and P. brevirostris. On the one hand, the two species are very similar in craniodental morphology; their isolated remains are often separated by size; and P. brevirostris is thought to derive from a large-sized population of Pl. perrieri. On the other, a larger size is not an aspect to disregard in hyenas as it usually correlates with the acquisition or better development of “bone-cracking” features; remains attesting the co-occurrence of Pl. perrieri and P. brevirostris are known from some sites dated at around 2.0–1.8 Ma; and the observed size differences between the two species exceed those recorded between extant and (at times) sympatric species. Therefore, taken alone the competition with P. brevirostris does not explain the extinction of Pl. perrieri, but considering it together with the concurrent spread and resulting peak of carnivoran diversity in Late Villafranchian faunas might. Pliocrocuta perrieri was outcompeted by its larger descendent in scavenging carcasses, while other carnivorans limited its options to deviate to other resources or adopt a different feeding behavior (e.g., active hunting). The case of Paciano is also significant in that it offers an example of the importance and feasibility to reconsider historical collections, even when the original material is lost entirely. In general, the seek for “novelty” that permeates current scientific literature ensues in few descriptions or reconsiderations of historical samples, especially if the new examinations confirm old results, but we emphasize the significance of such efforts in making old data truly available for the scientific community. In addition, the hyena from Paciano has a biochronological and stratigraphic value, confirming the occurrence in western Umbria of a depositional cycle older than the late Early Pleistocene one (well represented by the rich assemblages from Pietrafitta and Selvella; Farneta Faunal Unit).
The region of Umbria (Central Italy) represents one of the most interesting areas for the study of Mediterranean Plio-Pleistocene mammal faunas due to the occurrence of numerous paleontological deposits. Most of the fossils have been discovered within the Tiberino Basin, which extends across the centre of the territory for ca. 1800 km 2 , and the small Tavernelle-Pietrafitta Basin, south of Lake Trasimeno. Herein, we provide an additional report on an assemblage within continental deposits from the locality of Vigna Nuova (Piegaro, Perugia) in the Valdichiana Basin, which are observed to crop out in only a few areas of western Umbria. Taphonomic and geological evidence, coupled with rare excavation notes, have allowed the sample to be divided into two assemblages. The younger was collected from the conglomerates (here called "upper layer") which are still visible on the outcrop and is composed only of remains of cf. Leptobos. The data obtained from this fragmentary sample together with some sedimentological-stratigraphic considerations suggest that it can be correlated with other well-known Valdichiana assemblages (e.g., Selvella, Farneta), dated at ca. 1.5 Ma. The older sub-sample comes from a putative paleosol (here called "lower layer") no longer visible in the field. It includes cf. Megantereon, Canis sp., Mustelidae indet., Proboscidea indet., Equus cf. senezensis, cf. Leptobos, Croizetoceros ramosus, Pseudodama sp., Cervidae indet. (large size), and Sus cf. strozzii. This assemblage can be referred to the Coste San Giacomo Faunal Unit (late Middle Villafranchian; ca. 2.2-2.1 Ma). It represents the first well-preserved sample of this age in Umbria and one of the few recorded in Italy, opening new interesting research perspectives on the distribution of mammals within the Peninsula and also on the stratigraphy and evolution of the Valdichiana Basin. "Aggiungeva che per scavare le poche ossa che mi avrebbe subito inviate aveva dovuto faticare enormemente e lottare per due giorni contro ogni sorta di difficoltà. Ricevuto il graditissimo invio mi affrettai a consolidare e restaurare con ogni cura quelle primizie ed oggi sono lieto di poter annunziare, in così solenne adunanza, la interessante scoperta…" ("He added that in order to excavate the few bones that he would immediately send me he had had to work enormously and struggle for two days against all sorts of difficulties. Having received the very welcome shipment,
Here, we report on the exceptionally well-preserved deer record from the locality of Pantalla (central Italy), dated in the Early Pleistocene (ca. 2.1–2.0 million years ago). The fossils show a combination of characters that allows an unambiguous attribution to ‘Pseudodama’ nestii, of which they represent one of the most informative collections to date. Our comparisons—also conducted through CT-based methods on endocranial structures—reveal that the cranial and postcranial skeleton of ‘P.’ nestii displays a mosaic of intermediate characters between extant Dama and Cervus, but also that the affinities with Dama are prevalent. Some Cervus-like features especially in cranial morphology, can be interpreted as plesiomorphic characters supporting a basal position of ‘Pseudodama’ among the Cervini. Interestingly, three bone anomalies are described in the two male crania of ‘P.’ nestii from Pantalla and are interpreted as palaeotraumatological evidence resulting from different injuries suffered by the deer during their life. This allows opening a treasure trove of information on paleobiological aspects, including ontogeny and antler cycle and function.
The Listriodontinae were a common and widespread group of Suidae (pigs) that lived in an area extending from Portugal to China and to southern Africa. Here, we describe the new species Listriodon dukkar from Pasuda (Gujarat, India). It shares features with Li. pentapotamiae, evolved from it, and is the last representative of this lineage. The Listriodontinae flourished for about 10 million years, reached their maximum diversity and geographic extension during the Mid-Miocene Climatic Optimum (about 17–13.6 Ma), and their last records are close in age and date to ~ 9.8 Ma in the Indian Subcontinent, 9.78 Ma in Europe, and ~ 10 Ma in Africa. We review the environments in which the last listriodont lineages lived and went extinct. Their extinctions occurred against a background of increasing seasonality, vegetation change, a rise in bovid diversity and abundance, and local events, such as the European Vallesian Crisis and a dramatic drop in tragulid abundance in the Siwaliks. However, changes in the atmospheric pCO 2 may have contributed to their decline and extinction in all their geographic distribution. Decreasing pCO 2 is expected to have decreased sugar content and increased protein content of leaves and fruit. Hindgut fermenting Suoidea have higher protein requirements, while foregut fermenting Suoidea are more efficient in digesting sugars. Listriodontinae were probably foregut fermenters and were less well adapted in a low pCO 2 world.
We present the study of a composite, yet continuous sedimentary succession covering the time interval spanning 2.6–0.36 Ma in the intramontane basin of Anagni (central Italy) through a dedicated borecore, field surveys, and the review of previous data at the three palaeontological and archaeological sites of Colle Marino, Coste San Giacomo and Fontana Ranuccio. By combining the magneto- and chronostratigraphic data with sedimentologic and biostratigraphic analysis, we describe the palaeogeographic and tectonic evolution of this region during this entire interval. In this time frame, starting from 0.8 Ma, the progressive shallowing and temporary emersion of the large lacustrine basins and alluvial plains created favorable conditions for early hominin occupation of the area, as attested by abundant tool industry occurrences and fossils. This study provides new constraints to better interpret the hominin migratory dynamics and the factors that influenced the location and spatial distribution during the early occupation of this region.
A river otter hemimandible has been rediscovered during the revision of the historical collections of G.A. Blanc from Grotta Romanelli, complementing the ongoing multidisciplinary research fieldwork on the site. The specimen, recovered from the level G (“terre rosse”; early Late Pleistocene or late Middle Pleistocene), is here assigned to Lutra lutra . Indeed, morphological and morphometric comparisons with other Quaternary Lutrinae fossils from Europe allow to exclude an attribution to the relatively widespread and older Lutra simplicidens , characterized by distinctive carnassial proportions. Differences with Cyrnaonyx antiqua , which possessed a more robust, shellfish-feeding dentition, support the view of a successful niche repartition between the two species during the late Middle to Late Pleistocene of Europe. The occurrence of Lutra lutra from the “terre rosse” of Grotta Romanelli suggests deep modifications of the landscapes due to the ecological adaptation of the taxon, and indicates that the Eurasian otter spread into Europe at the Middle–Late Pleistocene transition.
The Miocene-Pliocene (Turolian-Ruscinian) transition represents a fundamental interval in the evolution of Euro-Mediterranean paleocommunities. In fact, the paleoenvironmental changes connected with the end of the Messinian salinity crisis are reflected by a major renewal in mammal faunal assemblages. An important bioevent among terrestrial large mammals is the dispersal of the genus Sus, which replaced all other suid species during the Pliocene. Despite its possible paleoecological and biochronological relevance, correlations based on this bioevent are undermined by the supposed persistence of the late surviving late Miocene Propotamochoerus provincialis. However, a recent revision of the type material of this species revealed an admixture with remains of Sus strozzii, an early Pleistocene (Middle Villafranchian to Epivillafranchian) suid, questioning both the diagnosis and chronological range of P. provincialis. Here we review the late Miocene Suidae sample recovered from the Casino Basin (Tuscany, central Italy), whose taxonomic attribution has been controversial over the nearly 150 years since its discovery. Following a comparison with other Miocene, Pliocene, and Pleistocene Eurasian species, the Casino Suidae are assigned to P. provincialis and the species diagnosis is emended. Moreover, it is recognized that all the late Miocene (Turolian) European Propotamochoerus material belongs to P. provincialis and that there is no compelling evidence of the occurrence of this species beyond the Turolian-Ruscinian transition (MN13-MN14).
The extant wild boar Sus scrofa has one of the largest geographical range of all mammals, and from its appearance in the late Early Pleistocene (Epivillafranchian) it is also widely represented in the European fossil record. Early forms of the species were larger than Late Pleistocene ones, but neither the chronology nor the causes of the size reduction have been thoroughly investigated. Here, we considered for the first time a large number of fossils from several late Middle Pleistocene to Early Holocene sites of the Apulian region (Italian Peninsula). In contrast to the supposed existence of a progressive trend towards small dimensions, morphometric comparisons and body mass estimates allow us to recognize several size oscillations during the late Middle Pleistocene-Early Holocene, with large forms occurring during interglacial stages and smaller ones during glacial stages. This suggests that fossil Apulian wild boar did not conform to Bergmann’s rule, that predicts larger size in colder climates due to the selective pressure towards lower surface area to volume ratio. Climate recrudescence may have played an indirect role in reducing the availability of trophic resources and hence promoting the observed pattern.
Herein we describe a fragment of a mandible with a deciduous fourth premolar (dp4) from the Early Pleistocene locality of Dunaalmás, representing the first confirmed report of Sus strozzii from Hungary. The comparison of dp4 measurements supports a statistically significant distinction between S. strozzii and Sus scrofa. The two species overlap in time during the late Early Pleistocene of Europe (Epivillafranchian), but suid remains of this time-span are seldom classified at a species level. The correct taxonomic identification of the Epivillafranchian suids, which are often associated with evidences of hominin presence, is of great palaeoenvironmental value because S. scrofa and S. strozzii possess different ecological requirements.
Introduction As with all members of the mammalian family Suidae, wild pigs belonging to the species Sus scrofa are not native to North America. The presence of these animals on this continent is attributable solely to their introduction by man. Such introductions have been both accidental (e.g. escaped domestic swine that have gone wild/feral) and intentional (e.g. Eurasian wild boar released to provide a new huntable game species) (Mayer & Brisbin 2008, 2009). Despite a prolonged presence on the continent, the wild pigs in North America have undergone a recent dramatic expansion in both their distribution and numbers. Concurrent with this increase has been an upsurge in the amount and diversity of damage being done by these animals to their host environments (Mayer & Brisbin 2009; West et al. 2009; Bevins et al. 2014). Annual estimates of this damage in the USA alone are in the billions of dollars (Pimentel 2007). Because of this recent situation, wild pigs are now typically considered to be an undesirable invasive species in places where they are found in North America. Unfortunately, invasive wild pigs are very difficult to either successfully manage or control. In response to these circumstances, research is being conducted to improve control techniques as well as trying to better understand the biology of these animals as it relates to their management. However, no viable or consistently successful solution to this problem has been identified to date. History of the Introduction Wild pig populations have had a long history in North America dating as far back as the European period of exploration and colonization in the western hemisphere. As such, the introductions of these animals have been both widespread and varied over the centuries. Additional new introductions have continued up through the present. Taxonomically, all of the free-ranging established populations of wild pigs found in North America belong to one species, Sus scrofa. Two types of S. scrofa, domestic swine and Eurasian wild boar, were introduced at various times onto the continent. Because these two types are conspecifics, wherever both of them have been found together in the wild, interbreeding has occurred (Mayer & Brisbin 2008, 2009). As illustrated in Figure 28.1, there are at present three general types of wild S. scrofa found in North America.
Introduction South America is facing a new wave of biological invasion of Sus scrofa and the novelty is related to Eurasian wild boar phenotype instead of the most common invasive variety, the feral pig (Jaksic 1998; Jaksic et al. 2002; Herrero & Luco 2003; GISP 2005; Canevari & Vaccaro 2007; Deberdt & Scherer 2007; Ghione et al. 2008; Novillo & Ojeda 2008; Pescador et al. 2009; MMA/CONABIO 2009; Cuevas et al. 2010; García et al. 2011; Barrios-Garcia & Ballari 2012; Sampaio & Schmidt 2013; Salvador & Fernandez 2014; Pedrosa et al. 2015; Skewes & Jaksic 2015). The current geographic distribution of wild boar is different from that of feral pig and useful to identify the implications of this invasion on political borders, ecosystems and protected areas (Figure 29.1). Sus scrofa in the wild performs many ecological interactions, becoming a new threat for the South American biodiversity as a potential predator, disease reservoir, and competitor (Coblentz & Baber 1987; Sicuro & Oliveira 2002; Pérez Carusi et al. 2009; Desbiez et al. 2012; Keuroghlian et al. 2012). The species is also the reason for many economic and social conflicts with human activities, such as livestock and crop attacks (Barrios-Garcia & Ballari 2012). Potential niche overlap is possible between Sus scrofa and the native pig-like peccaries (Sicuro & Oliveira 2002). Considering competition as one of the threats to biodiversity, the new massive invasion of S. scrofa makes the peccaries the species most threatened by wild boars and feral pigs. If other species and ecosystems coevolved with native pig-like varieties, they could deal better with similar ecological interactions of alien pigs than would do peccaries. For instance, we could assume the peccaries as a reference to assess the relative environmental impact of S. scrofa on the South American ecosystem. Moreover, this threat should take place in most parts of the continent (Figure 29.2). Peccaries can reach elevated biomasses in the Neotropical realm and are engaged in such important ecological interactions that they are often considered engineers or architects of ecosystems (Taber et al. 2008; Keuroghlian & Eaton 2009; Altrichter et al. 2012).
Introduction Feral pigs (Sus scrofa L.) are widely regarded as one of Australasia’s most destructive pest animals. They inhabit a large proportion of both Australia and neighbouring New Zealand. They can cause severe damage to a wide range of agricultural, biodiversity, and environmental resources. Private landholders, government agencies, and other authorities carry out many feral pig control programmes each year. However, pigs are also highly valued by some sections of the community for hunting or commercial exploitation. Although wild or feral pigs are widely distributed across the globe (Long 2003), the Australian situation is in many ways unique. There has been a long history of active feral pig management in Australia and, in contrast to New Zealand, Europe, and the Americas, wild-living pigs are consistently recognized and treated as a pest in all onshore jurisdictions. Conversely, the most widely used methods and tools for controlling feral pigs in Australia are uncommon elsewhere. This chapter will review the origin and distribution of feral pigs across Australia and New Zealand, their destructive impacts, useful values, and their management and control. It will highlight current and future challenges to the effective mitigation of pig impacts, and propose possible solutions. At a fine scale, some of these challenges may be peculiar to the region, but the broader trends and pressures from which they emerge are common to many regions of the globe where wild-living pig populations conflict with human values. Trends in Abundance and Distribution Notwithstanding speculation of a pre-colonial origin for wild-living pigs in some parts of north-eastern Australia (Baldwin 1983), the first known introduction of pigs onto the Australian continent occurred with the arrival of at least 48 animals with the first European settlers in 1788. While other livestock failed to thrive, free-roaming pigs increased rapidly and soon became such a pest to the fledgling colony’s food supply that orders for their control were issued within seven years of settlement (Robertson 1932). The subsequent declaration of a series of increasingly dire control orders suggests that the first attempts at regulation to manage damage caused by pigs were not very effective.
The extinct bovid Leptobos is one of the most characteristic elements of Eurasian faunal assemblages during most of the Villafranchian Land Mammal Age (i.e., from the late Pliocene to most of the early Pleistocene). Several species of this genus have been established since the end of XIX Century, but their taxonomic status and phylogenetic relationships remain unclear due to the fact that most of them are described on the basis of scanty material. European species are divided into two groups or lineages. The first includes L. stenometopon, L. merlai, and the poorly known L. furtivus, the second L. etruscus and L. vallisarni. While the last two species are well documented in the Italian early Pleistocene fossil record, very little is known on the L. stenometopon-merlai-furtivus group and especially on L. merlai, whose richest sample is from the French locality of Saint Vallier. Here, we describe new material of L. merlai from the early late Villafranchian of Umbria (central Italy), including a nearly complete female cranium and a male neurocranium with horn cores. These remains constitute the best-preserved and complete sample of L. merlai in the Italian Peninsula and bring new consistency to the fossil record of this species. In addition, they serve to confirm the extension of the spatial distribution of L. merlai to the south and of the chronological distribution of this taxon from the middle to the early late Villafranchian. Finally, we offer critical remarks on some not well defined Leptobos species.
The bone-bearing beds of Bethlehem were excavated by Gardner and Bate in the late 1930s, yielding an important Plio-Pleistocene faunal assemblage. In the 1950s, Hooijer revised the fauna and described elephant remains, including a large tusk, a mandible, several molars and some post-cranial elements, identified by him as Archidiskodon cf. planifrons. Recent preparation and Computed Tomography has given new insights into the early elephant remains from Bethlehem e both in terms of their anatomy and their post-depositional deformation. This includes two further mandibles, distorted and almost totally obscured by sediment, whose morphology has been revealed. The elephant material has been studied in detail and compared morphometrically with key taxa including Siwalik Elephas planifrons and European Mammuthus rumanus. The morphology of the molars cannot definitively distinguish between Elephas or Mammuthus, but their evolutionary grade, and the morphology of the mandible, are most conformable with a primitive mammoth intermediate between African M. subplanifrons and European M. rumanus. Conversely a largely complete tusk shows none of the spiral twisting associated with Mammuthus and is more conformable with Elephas. The possibility that two elephantid taxa are represented in the Beth-lehem deposit cannot be discounted, and is consistent with a wide size variation seen among the postcranial bones. These remains, together with some others recently described, represent the most primitive known elephantines out of Africa.
Early Pliocene suids of Europe are generally rare and poorly preserved, but some exceptional fossils have been found at Roussillon, France, Villafranca d’Asti, Italy, and Kvabebi, Georgia. We here describe and interpret an almost complete skull of a small suid from Musaitu and a mandible fragment from Dermenji, Moldova, which add precious information to the data base concerning these small European Pliocene suids, here attributed to Dasychoerus arvernensis (Croizet & Jobert, 1828). The Moldovan fossils provide an important biogeographic link between the Western European and Asian Plio-Pleistocene suines. The Musaitu skull, in particular, shows the elongated snout, enlarged canine flanges, pneumatised zygomatic arch and low slung incisor row characteristic of the genus Dasychoerus Gray, 1873. This species is important because it represents the group from which the African Kolpochoerus Van Hoepen & Van Hoepen, 1932 lineage probably emerged. The latter group is useful for biochronology because, having arrived in Africa, where it has been called Kolpochoerus deheinzelini Brunet & White, 2001 (in fact a junior synonym of Dasychoerus arvernensis) it evolved rapidly in dimensions and dental morphology. During the Pliocene the genus Dasychoerus, already adapted to tropical and sub-tropical climates, was widespread in mid-latitude Eurasia and Africa, but when much of mid-latitude Eurasia became boreal during the Plio-Pleistocene, the range of Dasychoerus shrank equatorwards, giving way to boreally adapted Sus scrofa Linnaeus, 1758 over much of its former territory, leaving a disjunct distribution of its descendants in Africa (Hylochoerus Thomas, 1904, possibly Potamochoerus Gray, 1854) and the tropical islands of the Far East (Dasychoerus). © Publications scientifiques du Muséum national d’Histoire naturelle, Paris.
The Eurasian beaver, Castor fiber is known from the Pleistocene in Europe, although there are references to the presence of the genus Castor since the Miocene, ca. 10 Ma. Beavers are present in the Iberian Peninsula since the Pliocene, though the first appearance of Castor fiber took place in the Early Pleistocene levels of the localities of the Sierra de Atapuerca, where the fossil beaver remains are found in the lower levels of Sima del Elefante and Gran Dolina (Early Pleistocene), Galería de las Estatuas (Late Pleistocene) and Portalón (Holocene). The presence of fossil beavers suggests humid and mild climatic conditions in the Early Pleistocene levels of Gran Dolina and Sima del Elefante. During the Middle Pleistocene, although conditions for beavers seemed favourable, they disappeared from Atapuerca, appearing again during the Late Pleistocene and the Holocene.
This is the author accepted manuscript. It is currently under an indefinite embargo pending publication of the final version.
Palynological investigations were carried out on the basal sediments of the Aulla-Olivola basin, precisely on the Aulla lacustrine deposit, which underlies by about 200 m, both stratigraphically and geometricall", the Olivola village, known for its famous Villafranchian fauna. This research has revealed; a very rich palynoflora and a long, complex, vegetational succession. dominated by arboreal formations, with high abundance of exotic taxa, including also tropical-subtropical elements; a climatic Sequence characterized by oscillations reflecting seasonality, especially in pluvial regime. A most evident climatic change occurs in the middle of the sequence. Moreover, the palynological sequence made it possible: to assign the examined sequence to the Pliocene, because of the abundance of "Tertiary" elements (Taxodium-type, Sciadopitys, Sequoia-type, Nyssa, Palmae, etc.); to-ascertain the rather archaic character of the Aulla pollenflora because of the significant occurrence Of pollen assemblages mainly confined to pre-Pliocene and early Pliocene (Symplocos, Sapotaceae, Clethraceae/Cyrillaceae-type, Magnolia-type, Sapindaceae- type, Sterculiaceae, cfr. Castanopsis, Tricolporopollenites sp. of archaic-type, etc.); to compare the Aulla pollenflora with Other Pliocene pollen microfloras from Italy and Other areas and to evidence affinities and differences. From these comparisons it appears clearly that the Aulla lacustrine episode represents the most ancient continental Pliocene deposit discovered in Italy by means of pollen analyses so far. In fact, the palynological and climatic features indicate that the pollen sequence of Aulla is certainly older than the establishement of glacial/interglacial climatic cycles in the Northern Hemisphere (—2,5 Ma). Also the Villafranca d'Asti deposit (Early Villafranchian in age) is older than 2,5 Ma on the base of the paleomagnetic data. But the pollen floras of Villafranca d'Asti (Fornace R.D.B.) do not include the pre- Pliocene and early Pliocene elements, which characterize the Aulla pollen records. All these considerations lead to Suggest an older chronostratigraphic collocation of the Aulla sequence, most likely in a pre-Vilafranchian interval, i.e., in the Ruscinian stage interval. The large floristic affinities with the pollen zones of the Brunssumian of the Netherlands, entirely included in the Ruscinian stage, and also the pollen affinities with the continental middle-Pliocene western-alpine deposits support the chronostratigraphical assignation of the Aulla deposit to a (probably late) Ruscinian phase. Finally, the pollen record of Aulla represents a particular uniform floristic biozone, which is named here "Macrian" phase, easily distinguished from the contiguous vegetational biozones. The results reported above allow to extend the chronostratigraphic scheme of the paleofloristic evolution in Northern-Central Italy during the Pliocene.
The long-lost mammal fauna from Gravitelli (Messina, Sicily, Italy) represents one of the most important records for investigating faunal dynamics during the Late Miocene of the Mediterranean, although it is unfortunately only known from descriptions carried out in the early 1900s, as the original collection was lost during the Messina Earthquake of 1908. Gravitelli suids have been referred to Propotamochoerus sp. after the redescription of the casts of two specimens that survived to the present day. However, there is further material that has not been considered, which makes that of Gravitelli one of the most abundant samples of Late Miocene suids from Italy, with a minimum number of four individuals represented. A reappraisal of all Gravitelli suids allows to ascribe them to Propotamochoerus provincialis (Suinae, Dicoryphochoerini), following a comparison with related Late Miocene to Pliocene species from Eurasia. Moreover, the re-examination of the geological setting of the locality reveals that the mammal fauna of Gravitelli occurred well below the pre-evaporitic deposits of the Tripoli Formation, whose base is dated in Sicily at ~7 Ma. Therefore, Gravitelli fauna either dates to the late Tortonian or, at most, to the earliest pre-evaporitic Messinian, corresponding to MN 11 or MN 12 in terms of mammal biochronology. This implies that the occurrence of P. provincialis at Gravitelli is the earliest in Italy and that emerged land masses connected Sicily with the European mainland earlier than 7 Ma. Available dates support a diachronous dispersal of Propotamochoerus in western Europe during the Turolian, being first known from the Balkans ~8.3 Ma, then from Gravitelli prior to 7 Ma, and then from the Iberian Peninsula since ~6.2 Ma. A similar pattern is known for Mesopithecus (Cercopithecidae). Although often discussed in light of its potential significance for Afro-Eurasian dispersals, only a fraction of the mammal fauna of Gravitelli has been reconsidered systematically. Notwithstanding the necessity of such dedicated studies, the faunal elements identified so far have an almost entirely European character and no species is shared with Cessaniti (Calabria), despite the two faunas have often been considered part of a paleobioprovince documenting a connection between southern Italy and northern Africa. At Gravitelli, the only species of African origin is the endemic hippo Hexaprotodon? siculus, but the extensive fossil record of insular hippopotamids testifies to their ability to colonize islands even in the absence of land bridges. Gravitelli hippos are nonetheless noteworthy, as the revised age of the site implies that they represent the earliest hippopotamids known outside Africa.
New and previously discussed suid remains recovered from Untermassfeld (Thuringia, Germany) are described. The late Early Pleistocene site of Untermassfeld yielded one of the most abundant samples of Epivillafranchian suids, and yet only a minimum number of 6 individuals has been identified. Untermassfeld suids have been considered the first representatives of the extant wild boar and ascribed to Sus scrofa priscus, then reassigned to Sus sp., and eventually to Sus strozzii. The latter attribution is favoured herein and some morphological differences from typical Villafranchian S. strozzii are noticed, which seem to be consistently present in other Epivillafranchian samples. At the moment it is not possible to evaluate whether these traits emerged abruptly in Epivillafranchian populations or were the result of gradual changes not recorded in the scanty post-Olduvai (c. 1.8 Ma) European suid record.
The Suidae from the late Miocene of Alsótelekes (northeastern Hungary, Borsod-Abaúj-Zemplén county) are described and assigned to Propotamochoerus palaeochoerus (Suinae) and cf. Parachleuastochoerus (Tetraconodontinae). The co-occurrence of these two taxa agrees with a reference to the early Vallesian (MN 9), as previously indicated from biochronological correlation of the small mammal fauna, and suggests the presence of woodland environments, with abundance of below-ground resources and direct access to water. This fits well with the diverse wetlands and riparian forests that characterized Lake Pannon ~10 Ma, as documented in the geographically close site of Rudabánya. The convoluted taxonomy of European Tetraconodontinae is discussed.
The Early Pleistocene site of Palan-Tyukan is located in Transcaucasia, northwestern Azerbaijan. More than 300 mammalian bones were laid close to each other in a 25 m2 lens-like accumulation, in a stratum of normally magnetised (the upper part of the Olduvai subchron) yellowish-grey Lower Apsheron loams. The present study is based on the analysis of the Equidae and Suidae remains. The taxonomic scenario of the Early Pleistocene Equidae is intricate and has been a matter of long debate with a multitude of hypotheses. The small-sized horse remains from the locality are ascribed here to species Equus (Allohippus) senezensis. The species likely used here are mostly open landscapes for their main habitat. The main conclusion about the Suidae remains from Palan-Tyukan was reached after a detailed morphological and metrical analysis. The fossil material is here referred to Sus strozzii. Seemingly, during the initial phase of the Early Pleistocene the species lived in humid subtropical Transcaucasian riverine forests and swamps. An Equidae/Suidae community of the Palan-Tyukan type presents evidence of the wider variety of environments ranging from wooded areas including bodies of water to scrub and even savannas as a landscape in a relatively humid subtropical climate.
A review of the abundant vertebrate fauna found in the Quaternary deposits of the Oricola-Carsoli intermontane Basin is here presented, with a brief excursus on the history of the research in this area and some biochronological considerations on these faunal assemblages. Palaeoloxodon antiquus, Hippopotamus cf. antiquus, and Stephanorhinus sp. remains were found in the 19 th century in the municipalities of Vallinfreda and Riofreddo and only described in pioneering studies by the Italian geologists and palaeontologists Giuseppe Ponzi and Alessandro Portis. After some accidental findings of other bones, never described, a hippopotamus was found in 1980 in Riofreddo and, subsequently, during fieldworks led by Aldo Giacomo Segre of the Italian Institute of Human Palaeontology (IsIPU), a very rich vertebrate fauna was found in the municipality of Oricola (L'Aquila). All the available bone material, most of which never studied, described, or mentioned before, is currently stored in different repositories and here has been re-evaluated and briefly described. Two distinct faunal assemblages have been detected; the older is ascribable to the late Villafranchian ELMA (European Land Mammal Age) and is characterised by Pseudodama cf. nestii, Leptobos aff. furtivus, Equus stenonis, Stephanorhinus etruscus, Mammuthus meridionalis, and other taxa. The younger assemblage is ascribable to the Galerian ELMA and is characterised by Palaeoloxodon antiquus, Hippopotamus cf. antiquus and Stephanorhinus sp.
The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ~2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris, Crocuta crocuta and "Hyaena" prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out.
In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of "H." prisca, occurred at ~0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and "H." prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene.
Seven leopard fossil bones from the important site of Equi (Massa, Italy), stored in the Musei Civici Reggio Emilia (Guido
Manodori Galliani Collection), have been studied. These leopard fossils provide new and important data on this elusive and rare felid. The
described specimens could represent at least two individuals, one female and one male. This note represents an important enhancement of
our knowledge of Late Pleistocene European leopards, especially considering their scarcity in the fossil record.
The Ponte Galeria area within the city of Rome has yielded numerous fossiliferous localities that represent a reference point for the study of the European Middle Pleistocene ecosystems. Within Ponte Galeria a rich collection of fossil mammals has been unearthed from Cava di Breccia – Casal Selce 2 (MIS 15) thus the site represents an optimal laboratory to investigate the palaeoenvironments of a defined territory during the Middle Pleistocene. We investigate the feeding behaviours of the ungulate community of Cava di Breccia – Casal Selce 2 to reconstruct the MIS 15 habitats and also compare the data with those of the nearby site of Fontana Ranuccio (MIS 11) which shares similar faunal composition with Cava di Breccia – Casal Selce 2 to test if ungulates occupied the same niches during two different interglacials. Open habitats with scattered woodlands characterised the Ponte Galeria area during MIS 15, whereas woodlands were more widespread during MIS 11 at Fontana Ranuccio. Ungulates display similar diets in both localities, suggesting that cervids, large bovids and equids adopted the same niche partitioning strategies during both interglacials.
The French Massif Central represents one of the rare occurrences in Europe where lower Pleistocene fossil deposits are associated to a substantial volcanic activity. Thanks to the abundance of volcanic zircons in the differentiated products of the Monts-Dore/Guéry stratovolcano, we were able for the first time to use U/Pb geochronology on zircon to precisely date lower Pleistocene fossil deposits. Zircon, a weathering-proof heavy mineral, was dispersed over the landscape through Plinian eruptions, and quickly reworked in fossil-bearing sedimentary deposits. Five Villafranchian fossil sites in the Upper Allier River valley (French Massif Central) have been precisely dated: Chilhac (MNQ17b, 2.285 ± 0.046 Ma), Mont Coupet (2.274 ± 0.032 Ma), Senèze (reference locality for MNQ18, 2.100 ± 0.029 Ma, confirming the previously determined 2.18–2.10 Ma ⁴⁰Ar/³⁹Ar ages), Blassac-La Girondie (MNQ19, 1.946 ± 0.029 Ma), and Vazeilles (1.843 ± 0.028 Ma). These new chronological data constrain the duration of the MNQ18 in central France to about 200 ka. The confirmation of an old age for the Blassac-La Girondie deposit indicates that the MNQ19 biozone started significantly earlier than previously thought.
Suidae remains recovered from the late Pliocene site of Collepardo (Latium, central Italy) are described and assigned to Sus arvernensis, a small-sized Ruscinian to Early Villafranchian (MN14-MN16a) species. In Italy, S. arvernensis only occurs in the Triversa Faunal Unit (MN16a), supporting the recently revised chronology of Collepardo. CT-scan methods are used to virtually extract and analyse a newly discovered neurocranium, providing the content for the first inner cranial description carried out on an extinct Suidae. Our analysis reveals that S. arvernensis has an anteroposteriorly elongated and dorsoventrally flat cerebrum, similar to that of the Asian Babyrousa babyrussa and the African Hylochoerus meinertzhageni. These species substantially differ in size and are representatives of two widely diverging phylogenetic clades, excluding relatively simple evolutionary or allometric explanations for brain morphology in Suidae.
The Vallparadís composite section (VCS) includes the nearby paleontological sites of Cal Guardiola and Vallparadís Estació (Vallès-Penedès Basin, northeastern Iberian Peninsula). The section spans from before the Jaramillo subchron to the early Middle Pleistocene (ca. 1.2-0.6 Ma). In this study, we describe the suid record from VCS and we review those from several other European sites, in order to refine the taxonomic identity and chronostratigraphic range of Quaternary suids in Europe. The VCS sample in- cludes a nearly complete skull, several teeth, and postcranial elements, and stands out as the richest European suid collection from the latest Early Pleistocene. Suid remains have been unearthed from both Cal Guardiola and Vallparadís Estació layers, whose age spans from the Jaramillo subchron (ca. 1.07-0.99 Ma; MIS31) to post-Jaramillo time (ca. 0.86 Ma; MIS21). Several craniomandibular and dental morphological features support an attribution to Sus strozzii. These features include a low and very deep preorbital fossa, a narrow nuchal crest, a well-developed longitudinal swelling in the middle of the mandibular corpus, the presence of styles/stylids in the upper/lower premolars, and especially the “verrucosic” morphology of the lower canine. The attribution to S. strozzii is also sustained by a cladistic analysis. These results provide interesting clues on the chronological occurrence of Quaternary suids. Sus strozzii is relatively common in Europe during the middle and early late Villafranchian (ca. 2.5-1.8 Ma), but it almost completely disappears during the latest Villafranchian (ca. 1.8-1.2 Ma). During and slightly after the Epivillafranchian (ca. 1.2-0.8 Ma), S. strozzii reappears in Europe although with relatively small samples, at VCS and several other sites including Untermassfeld (Germany; ca. 1.0 Ma), Le Vallonnet (France; ca. 1.2-1.1 Ma), Taman Peninsula (Russia; ca. 1.1-0.8 Ma), Arda River (Italy; ca. 0.99 Ma), and Slivia (Italy; ca. 0.8 Ma), among others. Consequently, in contrast to previous knowledge, we conclude that (1) S. strozzii survived in Europe (or returned there with a second dispersal event from Asia during the Epivillafranchian) at least until the end of the Early Pleistocene and (2) the arrival of Sus scrofa into that continent is not older than the Early-Middle Pleistocene boundary.
We investigated the fossil mammal assemblage from the late early Pleistocene (early Galerian) site of the Arda River. Seven taxa were identified: one carnivore (Ursus dolinensis), five artiodactyls (Sus strozzii, Pseudodama farnetesis, Praemegaceros sp., Bison sp., and Hippopotamus sp.) and one perissodactyl (Stephanorhinus hundsheimensis). The Arda River faunal assemblage includes taxa already present in the Villafranchian, such as Pseudodama and Sus strozzii. The bear Ursus dolinensis and the bison, a more evolved form than Bison (Eobison), are taxa typical of the beginning of the Galerian mammal age. Paleontological, sedimentological and paleobotanical data point towards the presence of woodlands settled on an alluvial plain with patches of grassland dated around 1.0 Ma. The continental sediments of the Arda River represent one of the first steps of the forming of the alluvial Po valley.
Suids are found in Europe before and during the Olduvai magnetostratigraphic subchron, including the
Fonelas P-1 site in the Guadix Basin (Andalusia, Southern Spain, ~2.0 Ma), in which the remains have
been ascribed to Potamochoerus magnus, and many other localities that record the presence of Sus strozzii
(e.g., Saint Vallier and Senèze in France). However, there is no pig record in the biochronological range
comprised between the post Tasso Faunal Unit, which marks the base of the Late Villafranchian (~1.8 Ma),
and their arrival in Western Europe at layer TE9 from Sima del Elefante, Atapuerca, Northern Spain
(~1.2 Ma), where pigs are recorded under the name of Sus sp., and at the sites of Untermassfeld (Germany),
Vallonnet (France) and layer EVT12 of Vallparadís Estació (Spain), dated 1.1-1.0 Ma (MIS31),
which suid remains have been ascribed to Sus scrofa priscus in the first site and to Sus sp. in the other two.
Later, the genus Sus is recorded everywhere in Europe as a ubiquitous member of the Epivillafranchian/
Galerian and posterior faunas. When pigs are in an ecosystem, they use to be abundant in the large
mammal community given their opportunistic feeding behavior and high reproductive success. Their
short gestation periods and elevated offspring numbers allow them to colonize new and varied environments
and territories faster than other ungulates, which use to display a K-reproductive strategy, with
a single pup per birth. For this reason, suids are usually preserved in the fossil assemblages after their
dispersal and colonization of a geographic region. The arrival of suids phylogenetically related to Sus gr.
scrofa into Europe marks the end of the Late Villafranchian and the beginning of the Epivillafranchian,
which is approximately dated at ~1.2 Ma. Given that suids are omnivorous, generalist mammals with
bunodont teeth that do not tolerate very low temperatures, this suggests that their colonization of
Europe can be related to a change in the ecosystems and climate. In addition, the arrival of suids
postdates the earliest arrival of hominins into Western Europe, which is documented at the Orce sites of
Barranco León-D and Fuente Nueva-3 (~1.5-1.2 Ma). In these sites, rich faunal assemblages, abundant
lithic artifacts and one human tooth have been unearthed after more than twenty years of excavations,
but no evidence of pigs has yet been detected.
The volcanic system of Senèze (Domeyrat, Haute-Loire) comprises a basanite flow, scoriae, a maar crater and phreatomagmatic products. New field research and detailed cartography clarify their geometry and geological relationships. The maar contains an important palaeontological site of the Early Pleistocene that yielded forty species of mammals attributed to the biozone MNQ 18 (of which it is the reference locality). The excavations undertaken between 2001 and 2006 recovered new fossils and documented their stratigraphic and geodynamic context. The fossiliferous site is located on the shore of the palaeolake and includes several lacustrine and slope deposits linked to the contemporaneous climatic changes which in turn produced a number of locally fossiliferous findspots which appear to be close in age. The discovery of ten tephras emitted by the Mont-Dore strato-volcano, situated 60 km to the NW allowed development of a remarkable tephrochronological framework. Because of the intense weathering of the tephras, their composition is determined by their mineralogical content (feldspars, brown amphiboles, brown and green diopside, titanite, apatite, zircon, biotite-phlogopite, Fe-Ti oxides) and the chemical composition of feldspars (anorthoclase, sodic sanidine and plagioclases). The study of these tephras reveals the importance of the contemporaneous (mainly trachytic) pyroclastic activity and confirms the polyphased deposition of the site. The 40Ar/39Ar laser dating of alkali feldspars from five tephras shows a relatively narrow range of age comprised between 2.09 and 2.21 ± 0.02 Ma (1σ, age relative to ACs-2 standard at 1.201 Ma). Senèze is thus confirmed as a key Early Pleistocene palaeontological site in Europe.