AMQ29(2) 4 Palombo 143-168NAG.pub Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 29 (2), 2016, 143 - 168 LARGE MAMMALS FAUNAL DYNAMICS IN SOUTHWESTERN EUROPE DURING THE LATE EARLY PLEISTOCENE: IMPLICATIONS FOR THE BIOCHRONOLOGICAL ASSESSMENT AND CORRELATION OF MAMMALIAN FAUNAS Maria Rita Palombo Dipartimento di Scienze della Terra, Sapienza Università di Roma, Rome, Italy Corresponding author: M.R. Palombo ABSTRACT: This research aims to investigate the large mammal faunal dynamics in SW Europe during the late Early Pleistocene. At that time, the climate forcing known as Mid-Pleistocene Revolution (MPR) induced deep, more or less gradual alterations and latitudinal dis- placements in European terrestrial biomes and exerted great influence on dispersal and dispersion of mammalian species. Large mammals did not generally move in multi-species waves of dispersal, rather each species changed its range depending on the suitability of environ- mental conditions in respect to its own environmental tolerances and ecological flexibility. Factors driving the remodelling of the range of a taxon, and time and mode of its dispersal and diffusion into SW Europe differed from species to species as from one territory to another, leading to diachronicity/asynchronicity in local first appearances/lowest local stratigraphical occurrences. As a result, correlations and bio- chronological assessments of local faunal assemblages may be difficult especially when firm chronological constraints are unavailable. Whether the peculiar composition of the late Early Pleistocene fauna, characterised by the persistence of some Villafranchian species and discrete new appearances of some taxa that will persiste during the Middle Pleistocene, may be indicative of any high rank biochronologi- cal unit is discussed. Evidence from SW Europe suggests that the chronological range of the so-called Epivillafranchian biochronological unit (whatever its biochronological rank could be) may span from about 1.5 (i.e. Lowest local Stratigraphical Occurrence, LlSO datum, of, among others, Homo, Xenocyon lycaonoides, Canis ex gr. C. mosbachensis, Megantereon whitei, advanced stenonoid horses, Praemegaceros, Bison) and about 0.85 Ma (i.e. Highest local Stratigraphical Occurrence, HlSO datum, of X. lycaonoides, M. whitei, P. pardoides). The biochronological unit would tentatively correspond from a biostratigraphical point of view to a virtual “range-zone”, i.e. the hypothetical body of strata representing the known stratigraphical occurrence in W Europe of X. lycaonoides-M. whitei. Nonetheless, sev- eral factors (e.g. the confusing taxonomic treatment of some taxa, the heterogeneous consistency of the fossil record in space and time - particularly as regard the number and richness of local faunal assemblages -, taphonomical biases, and the apparent dyachronicity asyn- chronicity of LlSO of some species that dispersed toward Western Europe during the post-Olduvai Early Villafranchian) make it difficult any attempt to “formalise” the “Epivillafranchian biochronological unit" as a European Land Mammal Age (ELMA). All in all, several lines of reasoning suggest an informal use of the term Epivillafranchian, pending a complete revision of the Villafranchian ELMA, its subdivisions and of the rank that each subdivision may actually have. Keywords: Biochronology, Bioevents, Epivillafranchian, Large mammals, late Early Pleistocene, SW Europe 1. INTRODUCTION Disentangling issues dealing with climate changes, mammal faunal dynamics and its implications for long distant correlations in the continental realm, is a chal- lenging task due to a number of hotly debated, unan- swered questions behind them (e.g. Walsh, 1998; Barnoski et al., 2003; Gingerich, 2003; van Dam, 2003; Lindsay, 2003; Prothero, 2004, 2012; Barnosky, 2005; Barnosky & Kraatz, 2007; Prideaux et al 2007; Lawler et al., 2009; Stewart, 2009;Davies et al., 2011; Badgley & Finarelli, 2013; Blois et al., 2013; Fortelius et al., 2014; Fernández et al., 2015 and references in those papers). During the evolutionary history of our planet, especially during the Cenozoic, climate changes were a recurring phenomenon (Zachos et al., 2001) and a number of sequences of ecological and evolutionary events devel- oped on a backdrop of climatic change, both warming and cooling phases (e.g. the Oligo-Miocene so-called Grande Coupure by Stehlin, 1910), although cooling episodes generally had a major impact at the Boreal middle latitudes. Some organisms, however, seem to have re- mained “the same” over thousands to perhaps millions of years in the face of environmental perturbations, high- lighting that the biotic response of individuals, species and communities to climate warming and cooling events is a highly complex phenomenon. Some other organ- isms, especially small mammals, rapidly evolved, thus their fossil record could be an important base for the biochronological divisions and regional correlations be- tween distant stratigraphic sequences/successions (see among others e.g. Chaline, 1972, 1987; van der Meulen, 1973; Agustí, 1986; Maul & Markova, 2007; Cuenca Bescós et al., 2010; Maul & Parfitt. S. A., 2010; Minwer- Barakat et al., 2011; Lozano-Fernández et al., 2013; Mayhew, 2013; Palmqvist et al., 2014). The synergistic action of climate and environmental changes, biotic interactions and feedback processes undoubtedly had some effects on the physiology of or- ganisms as well as on changes in the distribution of species and shifts of the limits of their range. During the Quaternary, in keeping with the vegetational cover and latitudinal displacement of biomes, large mammals, for Work presented during the AIQUA scientific conferences "Waiting for the Nagoya INQUA XIX Congress", held in Florence, June 18 - 19, 2015 144 M.R. Palombo instance, more frequently reacted to climate changes by varying the limits of their range than by originating new species in loco (see e.g. Magri & Palombo, 2013; Palombo, 2015a and references therein). As a result, discrete dispersal bioevents, introducing alien species into previously existing large mammal communities, changed their structure, giving rise to new internal dy- namics that led to a progressive reorganisation of mam- malian faunal complexes. This is the reason why dispersal is not only one among the fundamental processes in biogeography (crucial for understanding the evolutionary dynamics of organism distribution throughout time and across space) but also a factor to carefully consider for making feasible correlations among local faunal assemblages and fos- siliferous distant stratigraphic sequences/successions. Reconstructing and correlating chronological se- quences by means of large mammal fossil record is a problematic task because the continental sedimentary record is highly discontinuous, distribution of sites in time and space is not homogenous, rare species are sparsely represented, and ghost lineages are not so uncommon due to ecological factors and/or taphonomi- cal and sampling biases. In addition, continuous sedi- mentary sequences are rare and a number of local fau- nal assemblages lack sound chronological constraints. As a result, the known stratigraphic lowest/ highest oc- currence of a taxon within a geographical area does not necessarily reflect the timing of its actual first/last ap- pearances in time (e.g. Lindsay, 2003; Palombo, 2009 and references therein) and the diachronous appear- ance of a taxon may not necessarily depict its actual dispersal trajectory. Understanding time and mode of biological dispersals and their consequences is there- fore of crucial interest for any chronological assessment of local faunal assemblages, as well as for correlations and identification/characterisation of biochronological units. This research aims to investigate whether the pe- culiar composition of the late Early Pleistocene fauna (characterised by the persistence of some Villafranchian species and discrete new appearances of some taxa that will have persisted during the Middle Pleistocene) may be indicative or not of any high rank biochronologi- cal unit (i.e. European Land Mammal Age, ELAM). 2. MATERIAL AND METHODS The database consists of taxonomically revised lists of large mammal species (with a body weight of at least 10 kg) from selected local faunal assemblages (LFAs) mainly from SW Europe. A few Western Euro- pean LFAs, having a particular relevance for a better understanding of changes in the fauna structure in the post-Olduvai Early Pleistocene, were added to the analysis. The analysed LFAs range in age from about 1.6 to 0.8 Ma (V5a, V5b and G1 faunal complexes in Palombo, 2014). Besides to LFAs with a sound strati- graphic control, also isolated finds having particular taxonomical or chronological relevance were added to the analysis. The lists were compiled by revising and updating those resulting from previous studies on the Pleistocene mammals from W Europe (see e.g. Breda et al., 2010; Lister et al., 2010; Kahlke et al., 2011; Palombo, 2014, 2015a). Lutrini were not included in the study because of the scantiness and disproportion of their fossil record in time and space across the studied region. To provide a uniform baseline for the study ma- terial, the identifications of species was based on a taxo- nomical uniform view even for species/specimens whose taxonomy, systematics, and identification are controversial. 3. DISCRETE DISPERSAL EVENTS AND LOWEST LOCAL STRATIGRAPHICAL OCCURRENCES OF LARGE MAMMALS DURING THE POST-OLDUVAI EARLY PLEISTOCENE IN SW EUROPE In the post-Olduvai Early Pleistocene, in SW Europe discrete bioevents led to a progressive reorgani- sation of mammalian faunal complexes that lasted dur- ing the beginning of the Middle Pleistocene. The bio- events mainly consist of dispersals of few carnivores that either disappeared by the end of the Early Pleisto- cene or were still recorded in the Middle Pleistocene, and of large herbivores, most of which persisted throughout the Middle Pleistocene. A few extinctions of some among the most specialized Villafranchian spe- cies, and originations of new taxa in indigenous phyletic lineages are also recorded. Changes in the suitability of environments and the extent of ecological barriers triggered by climate changes affected time and mode of organism dispersal (see e.g. Colbert et al., 2012; Schlosset al, 2012). Dur- ing the active diffusion known as secular dispersal (which may take place so slowly that the diffusing spe- cies can undergo appreciable evolutionary change dur- ing the process), a species range can expand, contract or creep (expand in one direction while contracting in another). Sometimes organisms can move across great distances and a population of the original disperser's descendants may successful establishes at the end- point, whereas no viable populations may be present in the crossed territories. Moreover, barriers can either prevent the range of some taxa to reach some regions or cause long delays in dispersal, slowing or hampering the spread of a taxon in some territories. All considered, occasional jump-dispersals, the slow, progressive shift of the range (including translation) and the filtering ac- tion of barriers synergistically acted during secular- dispersal, leading to the diachronous versus asynchro- nous local appearances/disappearances of dispersing taxa. As a result, species-specific patterns of secular dispersal and dissimilarities in the inter- and intra-guild competition (sometimes leading to speciation and ex- tinction events at a local scale) may cause taxonomical composition of faunal assemblages to differ at a local scale even within the same geographical region. In SW Europe, discrete dispersal bioevents of taxa originally coming from both Asia and Africa led to a com- plex interplay among inter-sub-region dispersals and temporary shifts in the amplitude of the range of some species. The process, merging previously independent taxa in new palaeocommunities, triggered changes in the taxonomic and structural composition of mammalian 145 Late Early Pleistocene mammalian assemblages from SW Europe ............ faunal complexes. Mammals having a broad niche and high ecological tolerance persisted in face of alien spe- cies invasion and environmental changes, whilst the most specialized one disappeared (Palombo, 2014, 2015a, b and references therein). Available data indicate that during the late Early Pleistocene, the local lowest stratigraphical occurrence (LlSO) of a number of large mammal species was asyn- chronous across most of Europe. Only a few diachro- nous LISO permit to follow the displacement/ enlargement of the limit of the species’ range throughout time (Palombo, 2015b and references therein). Several factors (e.g. the different impact of global climate changes in different geographical settings, differences in species resilience and variability in competition and predation patterns within local faunal complexes, het- erogeneity of the fossil record in space and time and taphonomical biases) may have caused asynchronous appearances, though their interplaying is challenging to disentangle. As a result, the correct chronological order of LFAs and the correlation among fossiliferous strati- graphic sequences may be difficult to ascertain. Hence, the successions of SW European FCs here proposed (Fig 1) and the resulting chronological assessment of SW European LFAs can be regarded as the “best-fit” allowed by available data. 3.1. Major bioevents within the large carnivore guild The European carnivore guild was progressively renewed throughout the late Early Pleistocene, though the lowest and highest stratigraphical occurrences of some species were slightly asynchronous across the region (Fig 2). The middle sized canid Canis mosbachensis (whose taxonomy and phylogenetic relationships divide scholars, see e.g. Thenius, 1954; Kurtén, 1968; Bonifay, 1971; Kurtén & Poulianos, 1977; Pons-Moyà, 1987; Palmqvist et al., 1999; Sotnikova, 2001; Olive, 2006; Brugal & Boudadi-Maligne, 2011), was recorded for the first time in the Mediterranean region about 1.5 Ma in the ‘Ubeidiya LFA (Jordan Rift Valley, Israel), and may have dispersed towards SW Europe roughly in the same period. The species, indeed, is recorded at about 1.5-1.4 Ma (V5a FC) in the Iberian (Venta Micena LFA) and Italian (Pirro Nord LFA) peninsulas, together with the larger, hypercarnivous Lycaon-like dog Xenocyon ly- caonoides, which origined by anagenetic evolution from Xenocyon falconeri. In France the two canids are first reported later, shortly before (C. mosbachensis, V5b FC) and during (X. lycaonoides, G1 FG) the Jaramillo palaeomagnetic event respectively (cf. Martínez-Navarro & Rook, 2003; Moullé et al., 2006; Olive, 2006; Brugal & Boudadi-Maligne, 201; Palombo, 2015a for contrasting opinions). The African sabre-toothed cat Megantereon whitei possibly dispersed towards Europe shortly after the Olduvai-Matuyama paleomagnetic inversion. In Spain and Italy the species is recorded roughly at the same time at Venta Micena (Spain), Pirro Nord and Argentario promontory (Italy). M. whitei is reported in Greece (Apollonia 1 LFA, =M. cultridens in Koufos, 2014) and maybe in France (Ceyssaguet “couche 2” =M. cultridens in Tsoukala, 2004; Argant & Bonifay, 2011) for the first time shortly before the Jaramillo palaemagnetic event, Fig. 1 - Biochronological setting and chronological scheme for the post-Olduvai Early Pleistocene mammalian record from SW Europe. Fig. 2 - Biochronological setting of selected post-Olduvai Early Pleistocene large mammals from SW Europe (Spain =orange; France =green, Italy =bue) The data on which this table is based were established by using First/Last Historical Apparence (FHA/LHA) bioevents inferred from the lowest/highest occurrences of fossil remains in fossiliferous levels of key Iberian, French and Italian localities (LSDk/HSDk =Lowest/ Highest known Stratigraphical Datum). As discussed in Palombo (2009), the lowest and highest stratigraphical records of a taxon, respec- tively correspond to the “terminus ante quem” and “terminus post quem” of the actual time of the origination and extinction biovents (FHA/ LHA). Hence, new discoveries may substantially change the chronological range of taxa on which the scheme proposed here is based. RT =Ranging-through taxa ? =doubtful presence because of uncertain taxononomical identification and/or unsure chronological setting of finding sites and at about 1.1 Ma in central Europe (e.g. Untermass- feld, Germany, =M. cultridens adroveri, Hemmer, 2001). The time of dispersal of the leopard in Eurasia is uncertain. The species is not reported thus far in the Early Pleistocene of Asia, while it presence in Europe has been questioned due to the arguable identification of two lower carnassial teeth from Le Vallonnet, tenta- tively identified as P. pardus (De Lumley et al., 1988; Moullé et al., 2005). During the early Middle Pleistocene P. pardus was certainly present in Italy (e.g. Valdemino LFA, Masini & Sala, 2007), while it is doubtfully reported in the Iberian peninsula (Panthera sp., Sima de los Hue- sos LFA, see Gracía et al., 1997 for a discussion), where it was present for sure in the late Middle Pleisto- cene (Cueva del Congosto LFA, Palombo et al., 2009). Remains of cheetah-like cats identified as A. pardinensis were reported from a number of SW Euro- pean late Early Pleistocene LFAs, though they would not represent the same taxon within the A. pardinensis ‘macrospecies’. Their presence may be related to a dispersal event (see Hemmer et al., 2008, 2011 for a discussion). The subspecies A. pardinensis pleistocae- nicus, descendant of the Asian subspecies A. p. linxi- aensis, dispersed towards Europe during the late Early Pleistocene climatic worsening and aridification increas- ing. The LlSO in WEurope (abotut 1.05 Ma at Unter- massfeld, Germany, and Le Vallonnet, France) is slightly younger than that in North China (about 1.25 Ma, Qiu, 2006) (Hemmer, 2001; Hemmer et al., 2011; Kahlke, 2006; Spassov, 2011). The spotted hyaena appeared in Europe later, by the end of the Early Pleistocene. Its LlSO is at Atapu- erca (Trinchera de Ferrocarril, Northern Spain) in the TD4W level of Gran Dolina (García & Arsuaga, 2001), dated to about 0.9 Ma (Berger et al., 2008). In Italy, the spotted hyaena is first recorded about 0.72- 0.68 Ma BP in the Ponte Galeria 2 LFA (Roman Basin) (Palombo & Milli, 2011; Sardella & Petrucci, 2012), and a bit later in France at about 0.6-0.5 Ma BP (Grotte XIV, Cénac-et- Saint-Julien, Langlois, 2002). Some members of the large carnivore guild, first recorded in the post-Olduvai Early Pleistocene, likely originated from lineages already present in SW Europe. For instance, the hypothesis that the remains from Le Vallonnet (1.0 Ma, France), Gran Dolina TDW4 (ca. 0.9 Ma, Northern Spain) and Vallparadís (CGRD5, EVT10, EVT7 and EVT6, ca. 1.0-0.83 Ma, Vallés-Penedés Ba- sin, Spain) may belong to the cave bear lineage (Ursus ex group U. deningeri) cannot be discounted (Dabney et al., 2013; Madurell-Malapeira et al., 2014 and refer- ences therein). Tempo and mode of the appearance of extant European lynxes are debated as well as the taxonomic rank of the Middle Pleistocene South European ones. Lynx issiodorensis, a widespread species recorded in a number of Early Pleistocene sites, is regarded by most authors as the common ancestor of the European (Lynx pardinus and Asian (Lynx lynx) lineages. Lynx spelaeus/ Lynx pardinus spelaeus (see Boscaini, 2014 and refer- ences therein for a discusssion) is first recorded in Italy at Valdemino (? MIS15, Nocchi & Sala, 1997; Ghezzo et al., 2014) and in France at L’Escale (ca. 0.6-0.5 Ma; Bonifay, 1971) and Arago Cave (ca. 0.55 Ma; MIS14; Moigne et al., 2006). The LlSO of the species, however, may date back to the late Early Pleistocene, though the identification of the scanty remains found in the French LFAS of La Sartanette and Le Vallonnet (respectively dated to about 1.3 and 1.0 Ma) is challenging. The disappearance, i.e. the highest local strati- graphical occourence (HlSO), of some large Villafran- chian carnivores was sometimes nearly contemporane- ous, sometimes slightly asynchronous across the stud- ied region. Pachycrocuta brevirostris is last recorded in Spain in the EVT7 layer of Vallparadís Section dated to about 0.86 Ma (Duval et al., 2015), and in Italy in the Slivia LFA, believed to be close in age to the Early to Middle Pleistocene transition (Palombo, 2009 and refer- ences therein). The species seems to have disappeared in France in the post-Jaramillo Early Pleitocene, while is still recorded in central Europe during the early Middle Pleistocene (e.g. Süssenborn LFA, Turner & Anton, 1996). Vallparadís Estació (layer EVT7) records the HlSO of the puma-like cat P. pardoides in Europe (Madurell-Malapeira et al., 2010b). In Italy the species is last recorded in the Pirro Nord LFA (Petrucci et al., 2013) and in France even earlier [but see Hemmer (2001) as regards to the lower carnassial teeth from Le Vallonnet discussed above]. Panthera gombaszoegensis was the Villafranchian felid that lasted for the longest time in Europe. Its disap- pearance was apparently diachronous. The species is last recorded in Italy at the Early to Middle Pleistocene transition (Slivia LFA), in Spain in the earliest Middle Pleistocene (Gran Dolina TD8a LFA, Van der Made, 2013), in France in the early Middle Pleistocene (Grotte XIV LFA Cénac-et-Saint-Julien, ?MIS 15, MIS 13) (see Langlois, 2002), and finally in southern Poland in the late Middle Pleistocene (Biśnik Cave, MIS 9 or MIS 10) (Marciszak, 2014). The species was also recorded at Atapuerca in the Sima de los Huesos LFA (minimum age 427±12 ka, Arnold et al., 2014) by Morales et al. (1987), but the poor diagnostic morphology of some metatarsals, intermediate in dimension between the jaguar like cat and the leopard, prevents any firm spe- cific identification (Gracía et al., 1997). 3.2. Major bioevents within the primary consumer and omnivore guilds Changes in the guild of primary consumers re- corded in the post-Olduvai Early Pleistocene consist of: i) discrete appearances of herbivores mainly dispersing into SW Europe from Asia via East Europe and Africa, ii) some originations of new species within phyletic line- ages already present in SW Europe, and iii) discrete disappearances of the less ecologically flexible taxa in the face of alien species invasion and the climate insta- bility, forerunner of the Mid-Pleistocene Revolution/ Transition (MPR/T) (in its broadest definition between 1.2 and 0.5 Ma, Head & Gibbard, 2005; McClymont et al., 2013) (Fig 2). Among biovents believed to be related to the cli- mate worsening and increasing in aridity, the replace- ment of Leptobos representatives (e.g. Leptobos (Smertiobos) etruscus) by Bison species, more gregari- ous, and adapted to open, somehow arid environment, is regarded as an important event, reflecting the climatic 148 M.R. Palombo and environmental change. The LlSOs of primitive, small Bison in SW Europe are traced between 1.6-1.2 Ma. in Spain (Venta Micena, Fuente Nueva-3, Barranco León-5 and Atapuerca Sima del Elefante TE9), and in Italy (Pirro Nord) (Martinez Navarro et al., 2011; Van der Made, 2013; Masini et al., 2013; Palombo, 2015b). In France a more advanced species, Bison schoetensacki, is present at the time of Jaramillo subchron (Le Vallonnet) (Moullé et al., 2006 and references therein). In the same period a primitive species, Bison menneri, with quite long, slender meta- podials is recorded in central Europe at Untermassfeld (Germany) (Sher, 1997), together with a long-legged large deer Eucladoceros giulii. The latter has been re- garded as the final, large stage of the Eucladoceros lineage in W Europe, and one among the epitomes of the “Epivillafranchian” biochronological unit (see inter alios Kahlke H-D., 1997, 2001; Croitor & Kostopoulos, 2004; Kahlke R-D., 2009). The replacement of Eucladoceros representatives by giant deer belonging to the Asian tribe Megacerini has been regarded as an important component of the the post-Olduvai Early Pleistocene faunal turnover. Eu- cladoceros and large deer belonging to Praemegaceros and Megaloceros groups (or even Avernoceros as claimed by some authors), however, seem to have co- existed in Europe during the post-Olduvai Early Pleisto- cene (see inter alios Kahlke H-D., 1997, 2001; Croitor & Kostopoulos, 2004; Croitor, 2009; Van der Made & Dimitrijević, 2015 and references in those papers). Au- thors, for instance, are divided as the identification of the giant deer recorded at about 1.5 Ma in the Venta Micena LFA (V5a FC) [Praemegaceros solilhacus in Menéndez (1987) and Vislobokova, (2013); Euclado- ceros giulii in Van der Made (1999); Praemegaceros verticornis in Espigares (2010)], though the hypothesis that the giant deer from Venta Micena may belong to the genus Praemegaceros seems to be the most parsimoni- ous. At about that time, Praemegaceros obscurus is recorded in Italy in LFAs dating from about 1.5 to 1.3 Ma (e.g. Val di Chiana, Selvella, Pietrafitta, and Pirro Nord LFAs of V5a FC) (Napoleone et al., 2003; Abbazzi, 2004; Croitor, 2006; Petronio & Marcolini, 2013 and references therein). In France the LlSO of P. obscurus is slightly younger (about 1.2 Ma, Ceyssaguet LFA) (Croitor & Bonifay, 2001 and reference therein), while the presence of the species in Spain is questionable. A part for the disputed presence at Venta Micena (see above), the appearance of P. verticornis, generally con- sidered as more advanced than P. obscurus, was slightly asynchronous across the studied region. The species is reported shortly before the Jaramillo palaeo- magnetic event (about 1.4-1.3 Ma BP) in Spain (Barranco Léon 5 and Fuente Nueva 3, V5b FC) (Abbazzi, 2010), and during the Jaramillo submagneto- chron in Italy (Colle Curti, Coltorti et al., 1998) and pos- sibly in France (Saint Prest, Guérin et al., 2003; Vislo- bokova, 2013, but see Bonifay, 1981; Geraads, 1990; Lister, 1993 for contrasting opinions). Giant deer of the Megaloceros lineage appeared shortly after the Praemegaceros ones. A giant deer showing some simi- larities with Megaloceros savini may be present about 1.2 Ma in Greece (Libakos LFA) and in Northern Spain (Sima del Elefante LFA, Atapuerca) (cf. Van der Made & Tong, 2008). The species is identified roughly at the same time at (Cal Guardiola Layer CGRD2(Terrassa, Catalonia) (Madurell-Malapeira et al., 2015). Specimens identifiable as or closely relate to M. savini (i.e. in Spain Megaloceros novocarthaginiensis from Cueva Victoria, Van der Made, 2015) were present from the Jaramillo time to the transition to the Middle Pleistocene in Spain, France and Italy, while the species became more com- mon during the early Middle Pleistocene (see e.g. Vislo- bokova 2011, 2013 and references therein). A further evidence of the progressive, environ- mental related modifications of the first consumer guild across SW Europe is given by the roughly synchronous appearance of the slender, middle-sized Equus altidens (whatever related to in loco origination or a dispersal from Asia, see Alberdi and Palombo, 2013 and van del Made, 2013 for a discussion), and the slightly asynchro- nous LlSO of the larger Equus suessenbornensis. E. altidens is recorded, for instance, at about 1.5-1.4 Ma in Spain (Venta Micena LFA) and Italy (Selvella and Pirro Nord LFAs). The species was possibly present in France (Lézignan-le-Cèbe LFA) slightly later, between 1.3 and 1.1 Ma (Lézignan-le-Cèbe LFA) (Bourguignon et al., 2014, 2015). E. suessenbornensis is recorded firstly in Italy (Selvella and Pirro Nord), then in Spain (Barranco León 5 and Fuente Nueva 3) and maybe even later in France (Lézignan-le-Cèbe and possibly Le Vallonnet) (Alberdi, 2010; Alberdi and Palombo, 2013; Bourguignon et al., 2015). During the Jaramillo submagnetochron, a horse, Equus wuesti, slender but quite larger than E. altidens, is recorded in Germany in the Untermassfeld LFA (Musil, 2001). A horse, similar in size to E. altidens, but more robust, with wider distal articular surface and stronger keels (Equus apolloninsis) was present roughly at the same time in Greece at Apollonia (1.2 to 0.9 Ma) (Koufos et al., 1997; Spassov, 2003), and possibly in the slightly older site of Tsiotra Vryssi (Konidaris et al., 2015). The exact taxonomical position of E. wuesti and E. apolloniensis, the later believed to show some ass characters (Eisenmann & Kuznetsova, 2004), is contro- versial, although the hypothesis that they represent dif- ferent local ecomorphotypes (maybe species?) of the same lineage seems quite reasonable. During the MPR, and consistently with the spread of open environments in SW Europe, a number of Ca- prini representatives (subtribe Caprina and Ovibovina sensu Ropiquet & Hassanin, 2005; Hassanin, & Ropiquet, 2009) were recorded, whatever their appear- ance was related to an intraregional dispersal or they came from Eastern Europe. Among others, Praeovibos, already reported at the latest Gelasian-Early Calabrian transition from the Fonelas LFA (Guadix- Baza basin, Spain) (Arribas et al., 2009) and in the Caucasus at Dmanisi (Crégut-Bonnoure, 2007) (=Ovibovini indet. in Buksianidze, 2005), is recorded shortly after the Oldu- vai/Matuyama palaeomagnetic inversion first in Italy (Casa Frata LFA, V4 FC) (Masini el al., 2013) and then in Spain (Venta Micena and Barranco de los Conejos, V5a FC) (Crégut-Bonnoure, 2007). P. mediterraneus is reported around the time of Jaramillo palaeomagnetic event also in France (e.g. Le Vallonnet) and Greece (Apollonia 1) (Crégut-Bonnoure, 2002, 2007). 149 Late Early Pleistocene mammalian assemblages from SW Europe ............ The “steppe goat” Soergelia minor is recorded for the first time in Spain (Venta Micena) and Italy (Monte Argentario) (Martinez Navarro et al., 2012). Whether the geographical distribution range of the more advanced species Soergelia brigittae included or not the most of the Mediterranean region (at least from France to Greece, where the species is recorded in the Apollonia 1 LFA, Kostopoulos, 1997) is a matter of debate, due to the contrasting identification of the Caprini remains found at Le Vallonnet and Tour de Grimaldi (see Crégut- Bonnoure 2005; 2007; Crégut-Bonnoure & Dimitrijevic, 2006; Van der Made, 2013 for a discussion). The re- mains were ascribed by Moullé et al. (2004) to a new species, Ammotragus europeus, which was also identi- fied in Spain, at Fuente Nueva 3 (Guadiz-Baza Basin). The presence in Spain in the pre-Jaramillo Early Pleis- tocene of a bovid closely related to the extant Barbary sheep Ammotragus levia poses a question about the alleged dispersal from Africa to Europe of the Barbary sheep as well as the origin of the extant species, seen as endemic of Africa (cf. Geraads, 2010). Authors also disagree on the identification of Ovi- bovina remains from Pirro Nord, identified as “Ovibovini” indet. by De Giuli et al. (1987) and as Megalovis cf. M. balcanicus by Crégut-Bonnoure & Dimitrijevic (2006). Although some affinities with Megalovis cannot be dis- carded, the orientation of horn cores of an unpublished, badly preserved skull and the rather low hypsodonty index of molars make problematic any firm identification of the Ovibovina remains from Pirro Nord (cf. Masini et al., 2013 for a discussion). There are also some divergences about the identi- fication of a number of European Early Pleistocene Ca- prina remains. In particular scholars disagree about the taxonomy and systematic of thar/goat remains found in the post-Olduvai LFAs from SW European. A primitive Capra, may be descending from Capra dalii recorded at Dmanisi (Caucasus) (Bukhsianidze & Vekua, 2006), is recorded in the late Gelasian fauna of Fonelas 1 (Spain) (=Capra betica n. sp., Arribas and Garrido, 2008b). Dur- ing the post-Olduvai Early Pleistocene, primitive Caprina remains are reported in a number of Spanish LFAs, ranging in age from about 1.5 to 0.9 Ma (V5a to G1 FC) (Venta Micena, Barranco Léon, Fuente Nueva, Quibas, Huéscar 1), but their systematic, taxonomy and phylog- eny are controversial. Some authors consider them as belonging to a goat-like taxon “Capra alba”, may be descending from the Fonelas 1 goat, for others Spanish remains would belong to the genus Hemitragus (H. al- bus), or eventually they might belong to a clade related to Pseudovis or Ammotragus (cf. Crégut-Bonnoure, 2007 and Van der Made et al., 2008 for a discussion). These uncertainties make it difficult to ascertain whether the appearance of Capra? alba/Hemitragus albus re- lated to anagenetic evolution within an endemic lineage or to any dispersal event. During the post-Olduvai Early Pleistocene Hemitragus is confidently recorded at Le Vallonnet (=Hemitragus bonali in Moullé et al., 2006; =H. cf. H. orientalis in Crégut-Bonnoure, 2007). Among taxa believed to inhabit open environ- ments, Theropithecus, a herbivore primate assumed to inhabit rocky environments as its extant relative T. ge- lada, is sparsely reported from few Eurasian sites. The African baboon appeared in Asia (T. oswaldi delsoni) at about 1 Ma BP (Lower Boulder Conglomerate, Mir- zapur, Punjab, India) (Delson 1993 and references therein), and roughly at the same time in SW Europe, where it is only recorded in the Cueva Victoria LFA (Murgia, Spain) (Gibert et al., 1995), a site dated to about 0.85 Ma (from 0.98 to 0.78 Ma) (Gibert et al., 2015). Theropithecus sp. has claimed to be present at ‘Ubeidiya (Israel) about 1.5 Ma (Belmaker, 2002). As- suming the identification of remains from ‘Ubeidiya as correct, it seems rational to suppose that the African Pleistocene baboon reached SW Asia about 1.5 Ma BP, and then the dispersal of small populations (maybe across poorly suitable territories) enlarged the species range towards Eurasia. Tempo and mode of such a dispersal are, however, difficult to ascertain (cf. Hughes et al., 2008). On the one hand, the alleged dispersal towards Europe by crossing the Strait of Gibraltar (cf. Ferràndez-Cañadell et al., 2014) is hardly probable if not impossible (see O’Regan, 2008 for a discussion). On the other, the hypothesis that the species followed a Levantine dispersal route and would be unknown in the European intermediate areas because sub-optimal climatic and environmental conditions caused low population densities and a concomitantly extremely sparse fossil record, needs to be fully substantiated. It is worth noting, however, that some other species are recorded in one or just a couple of SW European sites [i.e. among others, the small sized buffalo from Venta Micena (Spain), which shows some similarities with the Chinese species Hemibos gracilis, (Martinez-Navarro et al., 2012), and the problematic remains of “Ammotragus/Soergelia” mentioned above]. Therefore, it is conceivable that the scattered distribution of some species depends on taphonomic factors and on the disparity of the fossil record during time and across space in the studied region. The long-term cooling trend and the environ- mental instability that intensified from about 1.2 Ma possibly facilitated the expansion towards SW Europe of the limits of the range of some generalist, ecologi- cally flexible species. Among others, suids belonging to the so-called “scrofic group” dispersed towards W Europe shortly before the Jaramillo subchron. Suids likely different from the Villafranchian Sus strozzi [a species reported for the last time at about 1.5 Ma in Val di Chiana (Italy) (Palombo et al. 2003; Napoleone et al., 2003)], is recorded at about 1.2 Ma in the layer TE9 of Sima del Elefante (Atapuerca, Northern Spain) (=Sus sp. in Carbonell et al., 2008; Sus gr. scrofa in Martinez-Navarro et al., 2015). Primitive, large wild boars were present shortly later (1.1 to 1.0 Ma) in Ger- many (Untermassfeld) (Sus scrofa priscus, Guerin & Faure, 1997), in France (Le Vallonnet) (Sus sp. in Moullé et al., 2006) and Spain (Vallparadís Estació layer EVT12 =Sus scrofa priscus, Madurell-Malapeira et al., 2010; Gran Dolina TD6B =Sus scrofa, Van der Made, 1998) (cf. Palombo, 2015a and references therein). Elaphine cervids were likely present around the Jaramillo submagnetochron in France at Saint Prest (Grossouvre et Stehlin, 1912; Guérin et al., 2003) (=Cervus elaphus acoronatus in Di Stefano & Petronio, 150 M.R. Palombo 1992), and in Spain at Barranc de la Boella (about 1.0 Ma) (Mosquera et al., 2015). Red deer lacking at the top of the antler even in adult individuals a multi-pointed “crown”, Cervus elaphus acoronatus, appeared by the end of the Early Pleistocene in Spain (Atapuerca, Gran Dolina TD6 and TDW4) (Van der Made, 1999) and Italy (Slivia) (a LFA estimated to be older than the Matuyama -Brunhes boundary (Bon et al., 1992; Gliozzi et al., 1997; Palombo et al., 2003; Sala & Masini, 2007). Red- deer representatives are also recorded in Germany (Dorn-Du ̈rkheim) (Franzen et al., 2000) and in the Levantine Corridor (Cervus cf. C. elaphus, Gesher Be- not Ya’akov) (Hooijer, 1959). The achievement of a so large geographical range confirms the ecological flexibil- ity of this “generalist” species, a mixed-feeders, inhabit- ing a variety of environments, even though red deer prefer open woodlands. The major bioevent in the primary consumer guild recorded at the transition from the Early to Middle Pleis- tocene is the arrival in SW Europe of two Elephantini (Palaeoloxodon antiquus and Mammuthus trogontherii). An isolated but well-preserved molar found at Slivia (Istria, NE Italy) may document the first appearance of straight-tusked elephants in SW Europe (Palombo, 1995; Palombo & Ferretti, 2005). Palaeoloxodon likely reached Europe from Africa during the latest Early Pleis- tocene, following a dispersal route along the Levantine corridor and maybe the Balkanic coasts (see Palombo 2015a for a discussion). Timing and mode of the re- placement of M. meridionalis by M. trogontherii in Europe during the Early to Middle Pleistocene transition actually depict a complex scenario susceptible to be differently interpreted (e.g. cladogenic evolution, hybridi- zation) (see Lister et al., 2005 for a discussion). Although a few species mostly inhabiting forest also appeared in the late Early Pleistocene [e.g. Capreolus representatives recorded in Germany (Untermassfeld) (Kahlke, 1997), Italy (Castagnone), (Siori and Sala, 2007), and Spain (Villaluenga del Rosario- El Chaparral) (Pacheco et al., 2011)], the pro- gressive taxonomical and functional change of the her- bivore/omnivore guild during the post-Olduvai Early Pleistocene was mainly related to the spread of open landscapes. The synergistic action of cooling, decrease in humidity, the combined effects of forest reduction and grassland extent allowed some large mammals (commonly inhabiting grassland/savannah-like or even mountain environments), to enlarge the limits of their range into the SW European region, though at difference pace and time. 4. THE BIOCHRONOLOGICAL SIGNIFICANCE OF THE PROGRESSIVE MODIFICATION OF THE MAM- MALIAN FAUNA AT THE LATE EARLY TO EARLY MIDDLE PLEISTOCENE TRANSITION IN SW EUROPE The discrete, sometimes diachronous/ asynchronous appearance of a number of large mam- mals recorded in SW Europe during the post-Olduvai Early Pleistocene resultes from the complex interplay among a few anagenetic evolutionary processes within indigenous phyletic lineages, discrete dispersal bio- events of taxa originally coming from Asia and Africa, and shifts in the amplitude of the range of some SW European species that triggered intra-regional dispersals (Palombo, 2015a). The filtering effects of ecological and physical barriers, the structure of palaeocommunities, and the resilience of resident mammals may have played a critical role in regulating the pace of the ap- pearances of each taxon in each area. The resident most flexible taxa persisted in face of alien species inva- sion and environmental changes, while the most special- ized disappeared, some others were apparently present either in only one locality or within a small geographical area. As a result, the competition/coevolution dynamics and the taxonomical and structural composition of mam- malian palaeocommunities changed at a different pace across the studied region. The difference in time and entity of dispersal events and related phenomena (upset of palaeocommunity internal equilibrium, keystone spe- cies extirpation, new inter-guild and intra-guild dynam- ics) at a local scale became more and more evident across the SW European region at the beginning and during the MPR. It is well known that the transition from the Early to Middle Pleistocene marks a fundamental change in the Earth’s climate system (Maslin & Ridgwell, 2005). From about 1.2 to 0.8 Ma, fluctuations in δ18O values significantly increased (Lisiecki & Raymo, 2005) and orbital obliquity at 41-ka cycles was superseded progressively by a rhythm of about 100 ka (Berger & Jansen, 1994). The amplitude of climatic oscillations significantly increased. Glacial-interglacial cycles ac- quired a more and more marked asymmetric structure (slow ice build-up, fast melting) thought to be associated 151 Late Early Pleistocene mammalian assemblages from SW Europe ............ _______________________________________________ >>>>>> Fig. 3 - Chronological assessment and faunal list of selected local faunal assemblages (LFAs) ranging in age from about 1.5 to 0.7 Ma. ? = doubtful presence; (?) = highly questionable identification; X? = doubtful identification; X(?) = identification possibly correct, although based on weak morphological/biometrical characters; X^ = ew bones identified as Hemitragus by Van der Made (2013), whereas according to Crégut-Bonnoure (2007) they belong partim to Hemitragus, partim to Soergelia genus; (?*) and (X?*) =large deer identified as P. verti- cornis and E. giulii by Guérin et al. (2003) and Lister (fide Breda & Marchetti, 2005) respectively. Locality, colour of the name: orange = Spain, green = France, Blue = Italy; black = West Europe. Green square = Lowest local Stratigrafical Occurrence; Gray square = Lowest/Highest local Stratigrafical Occurrence. (1) The group includes Ursus dolinensis, Ursus rodei and other specimens possibly belonging to Ursus deningeri-Ursus spelaeus lineage (see Madurell-Malapeira et al, 2014); (2) The group includes specimens identified as Lynx pardinus spelaeus and Lynx spelaeus; (3) includ- ing specimens identified as M. cultridens adroveri (Hemmer, 2001) and Megantereon “advanced form” (Sardella, 1998); (4) the identification of specimens from Pirro Nord, already identified as “Ovibovini indet.” (De Giuli et al., 1987) has been questioned by Masini et al. (2013). with fourth or fifth processional cycles, which have an important control on seasonality. In the course of the MPR, the response of biota was regionally different and vegetation and mammalian taxa varied with different regional timings and modes. As reported by Magri & Palombo (2013) as regard to SW Europe, from about 1.5 Ma there was a general change of both forest com- position and mammalian community structure: A sub- stantially similar vegetation dynamics were found across the studied region, (e.g. Suc and Popescu, 2005; Tzedakis et al., 2006; Leroy et al., 2011; Sadori et al., 2013; Combourieu-Nebout, et al., 2015), while the change in faunal composition and structure show signifi- cantly different trends at a local scale (Palombo 2014, 2015a, b). The LlSO/HlSO of a number of taxa in selected key Early and early Middle Pleistocene European sites (Fig. 3) on the one hand confirms the diachronicity/ asynchronicity of some local appearance/disappearance biovents, on the other puts out that some taxa were present either in only one locality or in a restricted terri- tory. The discrete appearances of species belonging to different taxonomical groups discounts any “migratory wave” model as causal factor for the faunal reorganisa- tion at the time of the MPR, while the scattered record of some species, especially the rare ones, suggests a possible influence of the heterogeneous consistency of the fossil record and taphonomical factors in altering the original composition of each LFA, as well as the actual chronological order of the first/last local appearances. 4.1 The matter of the question Available evidence indicates that the faunal transi- tion from the Villafranchian to the Galerian European ELMAs (tentatively regarded here as encompassing the time slice from about 1.5 to 0.8 Ma during which V5a,b and G1 FCs are recorded in SW Europe, see Palombo, 2014) was not an “abrupt” phenomenon, but a quite long, progressive transformation phase, which some- times developed in a diachronous way across the Euro- pean region. A common trait of many SW European late Early Pleistocene LFAs (V5a,b and G1 FC) is their pe- culiar taxonomical composition, which includes both long-lasting Villafranchian taxa at their latest evolution- ary stage and primitive representatives of some line- ages, which will characterise the so-called typical Gale- rian fauna (early Middle Pleistocene, G2 FC in Palombo, 2014). The resulting large mammal faunal complex was regarded by some scholars either as formed by a mix of Villafranchian and Galerian taxa, or the manifestation of a quite short (Bonifay, 1978; Azzaroli, 1983), thought composite ‘transitional phase’ [e.g. Azzaroli et al. (1988) stated that the “end-Villafranchian dispersal event”, chronologically close to Jaramillo, from 1.0 to 0.9 Ma, “did not take place at once”], or a much longer period of a progressive fauna modification, lasting about 0.6 Ma (Roebroeks & Kolfschoten, 1995), or even more. Agusti & Moyà-Solà (1998), for instance, discarded the concept of an “end-Villafranchian dispersal event” because in their view the dispersal phase had already started dur- ing the Olduvai submagnetochron with the arrival in Europe of Asian small and large mammals, such as M. (Allophaiomys) pliocaenicus, Lemmus, and “Ovibovini”. The Agusti & Moyà-Solà’s “phase” in it original meaning, however, refers to the “modernisation” shown by the classic late Villafranchian fauna (as epitomized by the Olivola+Tasso Italian Faunal Units, sensu Gliozzi et al., 1997), ranging in age from about 1.9-2.0 to 1.7-1.6 Ma. The faunal transition from the Villafranchian to the Galerian ELMAs was seen by other researchers as a “bioevent” (or a sum of bioevents) marking the end of the Villafranchian ELMA (=end-Villafranchian event, Azzaroli et al., 1988; Azzaroli, 1995; latest Villafran- chian, Koufos, 2001; Final Villafranchian, Spassov 2003), or the beginning of the Galerian ELMA (Gliozzi, 1997), or also an individual high rank biochronological unit (e.g. Protogalerian, Caloi and Palombo, 1995; Epiv- illafranchian sensu Kahlke, 2000, 2007, 2009; Epivilla- franchian sensu Bellucci et al., 2015). The proposed “event/faunal complex/biochron” as defined by scientists differ, however, each other in fauna composition and structure, chronostratigraphical meaning and temporal extent. The “latest Villafranchian”, for instance, would correspond to about the whole Calabrian stage (from ~1.8 to ~0, 78 Ma), the “Final Villafranchian” would ex- tend from about 1.6 to 1.1 Ma (Farneta and Pirro FU, sensu Gliozzi et al., 1997) (Spassov, 2003), while Caloi & Palombo (1995) introduced the term Protogalerian as a potential biochronological unit, in the context of a dis- cussion for a reasonable biochronological arrangement for the transition from the Villafranchian to Galerian ELMA. The Italian authors discussed three hypotheses mainly based on the ltalian mammal fauna: A) the begin- ning of the Galerian corresponds to the climate worsen- ing of MIS 22/24, B) the transition from the Villafranchian to Galerian fauna roughly developed at the time of Matuyama-Jaramillo inversion, C) the so-called “transitional fauna” represents an individual biochro- nological unit, i.e. the Protogalerian, characterised by the “appearance of the "galerian forms", persisting into the early Middle Pleistocene (as such or with their de- scendants)” (Caloi & Palombo, 1995, pag. 395). The Italian authors, in view of the first and last historical ap- pearance (FHA-LHA) recorded at that time, proposed for the hypothetical Protogalerian biochronological unit three alternative definitions and temporal extensions. The Protogalerian would correspond to the time slice ranging from about 1.4 (Pietrafitta LFA, characterised by the presence of Microtus ex gr. M. (AIIophaiomys) ruffoi, small-sized rhinoceroses and primitive megacerines) to 8.0 Ma (Slivia LFA, characterised by the presence of a number of new “Galerian” taxa, typically recorded in the early Middle Pleistocene fauna, and a significant reduc- tion of the Villafranchian ones), or alternatively the Slivia LFA would be included in the “true” Galeriant, or even the biochronological unit would have a wider temporal extension and would include the LFAs characterised by the appearance and persistence of voles belonging to the genus Microtus (Allophaiomys) (Caloi & Palombo, 1995, pag. 396, fig. 1). All the terms mentioned above were rarely or not used in literature. Conversely, since the 90s the use of the term Epivillafranchian (regard as a ELMA intermedi- ated in time and composition between the Villafranchian and Galerian ones) has more and more being increasing 154 M.R. Palombo accompanying faunal elements [e.g. Alces carnutorum (=Cervalces carnutorum), Panthera onca gom- baszoegensis (=P. gombazsoegensis), Acinonyx pardinensis pleistocaenicus, Megantereon cultridens adroveri (=M. withei), Puma pardoides, Pachycrocuta brevirostris, Ursus cf. dolinensis (=Ursus ex gr. U. den- ingeri), Canis (Xenocyon) lycaonoides (=X. lycaon- oides), and Canis mosbachensis] found at the site (Fig. 3). Whether the Epivillafranchian as defined by the Kahlke (2007, 2009) satisfies or not the characteristic informally recommended by some researches to indi- viduate/define a LMA (see below) it is a still open issue. The question arises, indeed, as regards to the minimum requirements a biochronological unit, i.e. a LMA, has to meet. 4.2. What is a Land Mammal Age? Loose definition, diverse/ambiguous applications and related issues Biochronology, the prime conceptual method for relating biologic events to the geologic time scale, is an important concept for geochronology, but neither the biochronological principles nor the criteria required to create a biochronological unit have been discussed in any stratigraphic code. Consequently, the biochronologi- cal units, in particular LMA, still are loosely defined and diversely/ambiguously used (see Lindsay, 2003, pag 213, 220). The LMAs’ concept was first introduced in the sci- entific literature about 75 years ago, but no formal defini- tions of a LMA itself has been published to date. It is however clear that bioevents and characteristics of fau- nal assemblages are of primary importance in creating a LMA, while the characteristics of the rocks that produced the fossil record, though to be considered, have a sec- ondary significance. Accordingly, a LMA has to be re- garded as biochronological entities. It is worth noting, however, that the initial attempt to chronologically order the mammalian fauna (i.e. the North American Cenozoic mammals) that led to the development of LMAs (i.e. North American Land Mammal Ages, NALMAs) based on biostratigraphical principles (e.g. Osborn & Matthew, 1909). The dawn of the LMAs’ concept goes back to the first half of the last century, when a seven-member com- mittee of the American Paleontological Society headed by H.E. Wood introduced in a report published in 1941 the “provincial ages” to chronologically order the North American Cenozoic mammal assemblages (Wood et al., 1941). The report illustrated the conclusions reached by the seven members of the committee, who had ap- pointed in 1939 for presenting a terminology to be used by vertebrate palaeontologists, consistently with the recently published North American stratigraphic guide (Ashley et al., 1933). Wood et al. (1941) characterised each “provincial age” (i.e. a faunal association of mam- mals that lived during that age) by listing 1) ‘‘index fos- sils’’ restricted to that age, 2) first and last appearances of fossils appearing /disappearing during that age, and 3) the so-called ‘‘characteristic fossils’’ that, even known from earlier or later ages, were commonly found at the time of the “provincial age”. 19 “provincial ages” were defined for the North American Cenozoic mammalian in literature. At the time when Bourdier (1961) first intro- duced the Epivillafranchian (=Epi-Villafranchién), the meaning was, however, different. Bourdier (1961), ana- lysing the geological and palaeontological context of the Quaternary deposits of the Rhône Basin, informally re- vised the quadripartite subdivision of the Villafranchian as introduced by Viret (1954) for the French faunas (Villafranchien ancien; Villafranchien normal inférieur; Villafranchien normal supérieur and Villafranchien supérieur). The Viret’s subdivision was successively simplified, and the resulting Villafranchian three partition (early, middle and late Villafranchian) is widely used by scientists even today, although meaning and temporal extension attributed by authors to each lower-rank bio- chronological unit differ (e.g. Heintz, 1970 and Azzaroli, 1977). The Bourdier’s work hypothesis was, conversely, the facto ignored. Bourdier (1961) added to Viret’s parti- tion two new subdivisions, the Proto-Villafranchian” for the most archaic faunas (Piacentian in age) and the Epi- Villafranchian for some late Early Pleistocene faunas, such as the Durfort (latest Early Pleistocene, Brugal, 1994; Palombo & Valli, 2004) and Saint-Cosme LFAs. According to Bourdier (1961), the Durfort LFA typi- cally represents the “warm” Epi-Villafranchian (= “Post- Villafranchién inférieur” in Bourdier, 1961, tab. 1), corre- sponding to the “Gu ̈nz-Mindel” interglacial, and regarded by some authors as roughly correlatable with the “Cromerian”, by others with MIS 15. Bourdier (1961) reported from the site Bison (Bos vel Bison), an ad- vanced Mammuthus meridionalis representative “with more tight lamellae, occurrence“, and horses “already similar to Equus caballus“(see Fig 3 for an updated list). The Saint-Cosme LFA was selected to represent the “cold” Epi-Villafranchian (=“Post Villafranchien supérieur” in Bourdier, 1961, tab. 1), corresponding to the ”Mindel glaciation”, by some considered roughly correlatable with MIS 12. The faunal list reporded by Bourdier (1961) includes “Equus stenonis, Rangifer tarandus? and semi-archaic molluscs”. Whether the “cold feature” of the Saint-Cosme LFA would be con- firmed by the attribution of the Saint-Cosme Formation to a glacial phase, although more recent (“recent Riss” in Bonvalot et al., 1984) is uncertain. A reappraisal of the term Epivillafranchian is due to Kahlke (2000, 2006, 2007, 2009), who state that the taxonomical composition of the European LFAs, ranging in age from the Jaramillo submagnetochron to around the end of the Early Pleistocene, cannot be regarded as a mixture of “older” (Villafranchian) and “younger” (Galerian) elements. These LFAs belong to a “a separate chronostratigraphical unit” (Epivillafranchian ELMA), lasting from about 1, 2 to 0, 9 Ma BP. According to the German palaeontologist, the Epiv- illafranchian biochronological unit is epitomized by the Untermassfeld LFA (Germany), the richest and most diversified among the late Early Pleistocene LFAs known in West European at that time (Kahlke, 2007, 2009). Accordingly, the Kahlke’s Epivillafranchian bio- chronological unit is identified by the “Bison menneri- Eucladoceros giulii-assemblage (i.e. the association of Bison menneri, Eucladoceros giulii, Cervus s.l. nestii vallonnetensis (=Dama vallonetensis), Capreolus cu- sanoides, and Stephanorhinus hundsheimensis) and 155 Late Early Pleistocene mammalian assemblages from SW Europe ............ fauna, only one referring to the Pleistocene. This implies a long persistence in time and a wide geographical ex- tent for each unit. A wide temporal and geographical extent also characterises the “mammal horizons”/”life zones” proposed in the conceptual scheme (Life Zone concept) developed by Osborn & Matthew (1909) to chronologically order the NW America Cenozoic mam- malian assemblages. Conversely to “provincial ages”, the “life zones” were based on biostratigraphical more than biochronological principles. Each life zone, indeed, was regarded as a complex of strata containing a par- ticular assemblage of mammalian taxa. Subsequently, the “life zone” concept was differently applied and ac- quired a biochronological meaning (Matthew, 1924). As highlighted by Lindsay (2003, pag. 221) “there was precedent in the historical development of NALMAs for the application of both biostratigraphical (e.g., the life zones of Osborn and Matthew, 1909) and biochronologi- cal (e.g., the faunal zones of Matthew, 1924) concepts”. This semantic and operational confusion, and the di- verse, sometimes ambiguous use of the term “LMA” depend on different concurring factors. On the one hand, the unclear use of terms such as biozone, a entity related to strata, and biochron, a temporal entity origi- nally introduced to indicate the total time represented by a biozone (Williams, 1901), and then sometimes used to indicate the time corresponding to a biochronological unit or the biochronological unit itself. On the other, the fact that the biochronology, in its attempt to define bio- events and chronologically order LFAs actually based on the fossil record available in the continental rocks. To make correlations among LFAs, and relate biochro- nological units to the geologic time scale, indeed, we have to refer to appearance/disappearance bioevents that are recorded in strata by biological crostratigraphi- cal markers (sensu Lindsay, 2003, pag. 215). Biochro- nology, however, differs from biostratigraphy because the evolutionary patterns and biological events, which regulate over time the dynamic evolutionary processes leading to taxonomical and structural changes of mam- malian fauna, are additional important factors to define any biochronological framework. All in all, the appearance and disappearance bio- events and the biological aspect (taxonomical composi- tion and ecological structure) that make a mammalian fauna different from earlier and later faunal complexes recorded in the same geographical area, are the main factors we have to consider to properly define a biocro- nological unit. Following Tedford (1970), for instance, a biochro- nological unit should be regarded as non-overlapping and "ecologically adjusted groups of animals with spe- cif ic geographical l imits and chronologic range" (Tedford, 1970, pag. 602). Lindsay (2003, pag. 222) defined more explicitly a LMA as “a relatively short interval of geologic time that can be recognised and distinguished from earlier and later such units (in a given region or province) by a characterizing assem- blage of mammals” de facto, and correctly, releasing any LMA by the deposits from which its characteristic taxa were derived. LMAs, therefore, have commonly been considered as biological entities (i.e. the natural associations of fossil species), because “the features of life during LMAs are of primary concern; the features of the rocks that produced the record are of secondary concern” (Lindsay, 2003, pag. 221). Although a LMA is, therefore, the period during which a peculiar association of mammalian taxa (usually genera according to most of authors), which differ in composition from the groups of taxa found in any other biochronological unit of the same rank, criteria to create it still remain somehow ambiguous. Moreover, given that the mean duration of a large mammal species is about 3 Ma (i.e. 3.21 Ma for larger North American Cenozoic mammals) (Prothero, 2014), it turns out that shorter the focal time slice and smaller the geographical area are, more difficult is to properly define a LMA. A bioevent, for instance, can only be inferred on the basis of the sum of known, local paleobiological data as documented by the fossil record derived from strata (e.g. Walsh, 1998; Palombo, 2009 and references therein). The actual du- ration of a large mammal species at large geographical scale may be obscured by analysing faunal change over geological short time in a small geographical area due to some regionalism, which in turn may depend on the local dispersal dynamics and competition. Additional drawbacks concern the number of LFAs with unclear or imprecise chronostratigraphic constrains, that may be proportionally higher at small than at large geographical scale, and the fact that an analysis performed on short geological time periods sometimes may force to consid- ered taxa at a low taxonomical rank. This increases the problems related to the different taxonomies applied by authors (sometimes irrespectively of the phenotypic plasticity of taxa, according to which a specie/genotype can produce different phenotypes in different environ- ments) that may mask the actual relationships among LFAs. 4.3. The Epivillafranchian as a biochronological unit In the attempt to characterise the Epivillafranchian as a ELMA following Kahlke’s definition (Kahlke, 2007, 2009), some difficulties arise. The main problem relates to the confusing/debated taxonomic treatment of a num- ber of taxa (in particular some among those listed by Kahlke (2006, 2009) as “accompanying faunal ele- ments”), the disagreement about their phylogeny and chronological range, and the exclusive presence at Untermassfeld of a few species that are either only re- corded at the site or also dubitatively reported from few other W European sites, and that may be taxonomically or ecologically vicariant of species commonly recorded in other European LFAs. The biochronological assess- ment of the potential Epivillafranchian LFAs and long distance correlations are therefore problematic, espe- cially if firm chronological constraints are unavailable. In recent times, the discovery of new rich late Early Pleistocene fossiliferous sites has been changing the already proposed chronological scenarios, and the term “Epivillafranchian” seems to have been acquiring a con- notation of “time phase” rather than maintaining the bio- chronological meaning of “a peculiar association of mammalian taxa differing in composition from the groups of taxa found in any other biochronological unit of the same rank”. Bellucci et al. (2015, pag. 87) attempted to formal- 156 M.R. Palombo accompanying the two “index fossils” [see the definition of the Epivillafranchian ELMA proposed by Kahlke (2007, 2009) and emended by Bellucci et al. (2015)] possibly appeared before the Jaramillo submagneto- chron (e.g. scrofic suids reported from Sima del Elefante TE9, and Megaloceros savini recorded in the Cal Guar- diola CGRD2 LFA), most were already present at about 1.5 Ma (e.g. Ursus ex gr. U. deningeri, Canis ex gr. C. mosbachensis, M. whitei, Xenocyon lycaonoides, E. altidens, E. suessenbornensis, maybe Stephanorhinus ex gr. S. hundsheimensis, Praemegaceros spp., the Italian advanced Dama-like deer possibly close to D. vallonetensis, and primitive Bison representatives), and some others, including long-lasting late Villafranchian taxa (i.e. P. gombaszoegensis), were still present during the early Middle Pleistocene (Fig. 2a-b, 3). All things considered, available data suggest that the taxonomical composition and structure of mammal- ian palaeo-communities had already been changing in SW Europe about 1.5-1.3 Ma, roughly in the same time span of the first human dispersal towards Europe. The first appearance of genus Homo, is a bioevent of unde- niable relevance to be considered in proposing any bio- chronological revision of the Villafranchian ELMA. Some other bioevents to take into account are the Highest local Stratigraphical Occurrence (HlSO in the latest Early Pleistocene of some carnivores such as those last recorded in some Spainish LFAs. The dirk-toothed cat Meganteron whitei and the pack-hunting canid Xeno- cyon lycaonoides were reported for the last time in the Cueva Victoria LFA (Guadiz-Baza Basin), dated to about 0.85 (0.98-0.78) Ma (Gibert et al., 2015). The lycaon- lyke canis and and the puma-like cat P. pardoides had their LlSO in the post-Jaramillo Early Pleistocene EVT7 layer of Vallparadis Estació section (Catalonia, Spain) (estimated age ranging from 0.99 to about 0.83 Ma, see Fig. 2b) (Alba et al., 2008; Madurell-Malapeira et al., 2010; Duval et al., 2011b, 2015; Lozano-Fernández et al., 2015). The evidence from SW Europe suggests that the chronological range of the so-called Epivillafranchian biochronological unit (whatever its biochronological rank could be) should span from about 1.5 Ma (LlSO datum of, among others, Homo, X. lycaonoides, C. ex gr. C. mosbachensis, M. whitei, advanced stenonoid horses, Praemegaceros, Bison) to about 0.85 Ma (X. lycaon- oides, M. whitei, and P. pardoides HlSO datum). In this way and from a biostratigraphical point of view the Epiv- illafranchian would tentatively correspond to a virtual “range-zone”, i.e. the complex of bodies of strata re- cording the known lower/higher stratigraphical occur- rence and the presence in W Europe of X. lycaonoides and M. whitei. The appearance/disapprearance of such carnivores would be regarded as biochronological events that acquire the meaning of “biological chronostratigraphical markers”, because “tied to a dis- crete stratigraphic sequence and related to other strati- graphic sequences and/or chronostratigraphic mark- ers” (cf. Lindsay, 2003, pag. 215 for a discussion). Nonetheless, several factors (e.g. the debated taxonomic treatment of some taxa, the heterogeneous consistency of the fossil record in space and time - par- ticularly as regard the number and richness of LFAS -, ize the Epivillafranchian as a biochronological unit (European Land Mammmal Age, ELMA) valid at a Euro- pean scale by revising and integrating the definition proposed di by Kahlke (2006, 2009) “with data from other European sites to better define the faunal turnover occurred at the end of the Jaramillo Subchron”. The Italian authors defined “the Epivillafranchian ELMA” on the basis of appearance bioevents, regarded as valid at a European scale, which would mark respectively the beginning of the Epivillafranchian and Galerian ELMAs. Accordingly, the Epivillafranchian biochronological unit would chronologically extend between about 1.2 and 0.9 Ma, i.e. the time slice encompassed between the ap- pearance of P. verticornis (the Galerian beginning event sensu Gliozzi et al., 1997) and B. menneri and the ap- pearance of C. crocuta (end of Epivillafranchian and beginning of the revised Galerian) (Bellucci et al., 2015, fig. 2, pag. 87). Moreover, the authors stated that “together with the dispersal of P. verticornis and B. men- neri, the beginning of the Epivillafranchian can be placed in correspondence of the diffusion of M. savini and S. scrofa priscus” (Bellucci et al., 2015, pag. 87). There is however some uncertainty related to the chronology of bioevents selected as potential biological chronostratigraphical markes by Bellucci et al. (2015). In particular, P. verticornis, identified at Venta Micena by Espigares (2010), was present in the Barranco Leon 5 and Fuente Nueva 3 LFA. Barranco Leon 5 is dated to about 1.4 -1.3 Ma (Duval et al., 2012; Toro-Moyano et al., 2013), while Fuente Nueva 3 may be even older (1.50 ± 0.31 Ma in Álvarez et al., 2015). Likewise, the spotted hyaena is recorded for the fist time at Atapuerca (North Spain), in the TD3-4 level of the Gran Dolina stratigraphic section, one of the most complete with terrestrial deposits recording the Early to Middle Pleisto- cene transition in Western Europe. The TD3-4 and TD5 levels were dated to about 0.99 and 0.94 Ma (Berger et al., 2008; Moreno-García, 2011). In addition, the actual phylogenetic relationships of B. menneri (well known from the rich sample of Unter- massfeld, but poorly doubtfully recorded in other sites) with the Early Pleistocene Bison relatives reported in Western Europe, and its actual significance as a ‘‘index fossils’’ are unclear. It still is an open question, for in- stance, whether the Untermassfeld bison was a taxon geographically restricted to the Western-central Europe, and the putative ancestor of “Bison voigdstedensis” [a species recorded in the Gran Dolina LFAs and dubiously reported by Cuenca-Bescós & Gracía (2007) in the Sima del Elefante LFA (TE-LRU level, ATAFU1 faunal unit)] as suggested by Van der Made (2013). Furthermore, E. giuli would be already present at about 1.5 in Spain at Venta Micena (Van der Made, 1999) (see discussion above) and the species is tentatively reported for from Sima del Elefante TE9-level (TE-LRU stratigraphic se- quence) (Cuenca-Bescós & Gracía, 2007). The actual taxonomical identity of the deer, the identification of some remains ascribed to the species as well is its chronological range are debated, although the presence of slender giant deer during the late Early Pleistocene is undisputable. Moreover, some Epivillafranchian taxa being part of the “Bison menneri- Eucladoceros giulii-assemblage” or 157 Late Early Pleistocene mammalian assemblages from SW Europe ............ taphonomical biases, and the apparent diachronicity/ asynchronicity of LlSO of some species that dispersed toward Western Europe during the post-Olduvai Early Villafranchian) make it difficult any attempt to firmly de- fine the “Epivillafranchian” as a “formal biochronological unit" (i.e. ELMA). Moreover, it should also be noted that any attempt to assign to the Epivillafranchian a ELMA rank implies a revision of the biochronological meaning and status of the sub-biochrononological units into which the Villafranchian ELMA is currently subdivided. The early Villafranchian, for example, may have a stronger significance as potential ELMA than the Epivil- lafranchian actually has. All in all, several lines of reasoning suggest an informal use of the term Epivillafranchian, pending a complete revision of the Villafranchian ELMA, its subdi- visions and the biochronological rank each subdivision may have. 5. REMARKS The structural and taxonomical changes of large mammalian palaeocommunities that led to the establish- ment of the so-called Epivillafranchian fauna has been regarded as linked to the early phase of the climate forcing known as Mid-Pleistocene Revolution (MIS 36 - MIS15-13). Nonetheless, an initially slight intensification in the amplitude of climatic oscillations is detectable after the quite long MIS 55 interglacial, in particular since about MIS 52, though it progressively increased successively from MIS 36 onward. The astronomically-forced climate changes and the developing of a new global climatic regime promoted alterations and latitudinal displacements in European terrestrial biomes and was a critical factor in driving faunal turnovers, removing keystone species, exerting great influence on dispersal and dispersion of mammal- ian across and between continents, in turn triggering new competition-coevolution dynamics that contributed to either the extirpation of some taxa or to the appear- ances of others by anagenetic evolution in some phyletic lineages. Large mammals did not generally move in multi- species waves of dispersal, rather each species changed the limits of its range depending on the suit- ability of environmental conditions in respect to its own environmental tolerances and ecological flexibility. Therefore, taxa answered to climate stimuli at a dissimi- lar pace and in a different way. Some large mammals, for instance, had already extended the limits of their range to and/or are recorded for the first time in SW Europe at about 1.5 Ma. Some others appeared shortly before or during the early Jaramillo submagnetochron, few just after the Jaramillo-Matuyama palaeomagnetic inversion, some were reported from a number of LFAs, others only from small geographical areas or in a single LFA (Fig. 2, 3). Although few diachronous appearances permit to follow the displacement/enlargement of the limit of a species’ range, the majority of LlSO (especially but not only that of rare species) documented in the available fossil record are asynchronous at least across the SW European region. It is a challenging task under- standing which among several factors (e.g. among oth- ers the different impact of global climate changes in different geographical settings, differences in species resilience and variability in competition and predation patterns within local faunal complexes, inadequate avail- ability of long stratigrafical sequences, taphonomical and sampling biases) might have caused the asynchronous appearance of several taxa across W Europe in the post -Olduvai Early Pleistocene. As a result, large mammal assemblages may substantially differ in taxonomical composition at regional and local scale, making correla- tions problematic and the compelling definition of any biochronological unit uncertain. Based on available data it would be suggested, as working hypothesis, that the “Epivillafranchian biochro- nological unit” (whatever its biochronological rank could be) should span from about 1.5 Ma (e.g. Homo, X. ly- caonoides, C. ex gr. C. mosbachensis, M. whitei, ad- vanced stenonoid horses, Praemegaceros, Bison, LlSO datum) and about 0.85 Ma (X, lycaonoides, M. whitei, P. pardoides HlSO datum). Several lines of reasoning, however, recommend a great caution in formalising this biochronological unit. As said above, a number of fac- tors hamper this attempt. To answer the question on the chronological meaning and the temporal range of the “Epivillafranchian” as well as be able to make correla- tions between distant sequences, the central challenges are: i) to remove disagreements about the taxonomy and phylogeny of some “Epivillafranchian taxa”; ii) to scrutinise the factors that cause the restricted geo- graphical distribution of a few others; iii) to appraise the significance of heterogeneous consistency of the fossil record in space and time (e.g. number and richness of LFAs); iv) to remove taphonomical biases that may af- fect the chronological reliability of local first/last appear- ances especially as regard to rare species; v) to improve the chronostratigraphical constrain of various LFA and understand and eliminate some inconsistencies in abso- lute geochronological dating provided for others. More- over, any attempt to properly define the biochronological rank of the “Epivillafranchian biochronological unit” can- not avoid a reappraisal of the Villafranchian ELMA and of the status and significance of its subdivisions. All in all, available evidence seems to be still inade- quate to ascertain whether the late Early Pleistocene LFAs (between 1.5/1.3 to 0.9-0.8 Ma) form or not a well- defined biochronological unit (i.e. an ecologically ad- justed groups of animals with specific geographical limits and chronologic range, Tedford, 1970), and what may be its biochronological rank. Concluding, several lines of reasoning suggest an informal use of the term Epivillafranchian, pending a complete revision of the Villafranchian ELMA, its subdi- visions and the relative rank each subdivision (i.e. the Early Villafranchian comparatively to a hypothetical Epivillafranchian) may have. ACKNOWLEDGEMENTS We wish to thank the two anonymous reviewers for critically reading the previous version of the manuscript. This research was funded by Italian MURST (Sapienza University 2014, project C26A14BNRM led by M.R. Palombo) 158 M.R. Palombo Mammalia) en el registro euroasiático (Fonelas P- 1, Cuenca de Guadix, España). In: Arribas A. (Ed.), Vertebrados del Plioceno superior terminal en el suroeste de Europa: Fonelas P-1 y el Proyecto Fonelas Cuadernos del Museo Geomin- ero, n. 10. Instituto Geológico y Minero de España, Madrid, 2008, 461-473. Arribas A., Garrido G., Viseras C., Soria J.M., Pla S., Solano J.G., Garcés M., Beamud E., Carrión J.S. (2009) - A mammalian lost world in Southwest Europe during the Late Pliocene. PLoS ONE, 4(9), e7127. Ashley G.H., Cheney M.G., Galloway J.J., Gould C.N., Hares C.J., Howell B.F., Levorsen A.I., Miser H.D., Moore R.C., Reeside J.B., Rubey W.W., Stanton T.W., Stose G.W., Twenhofel W.H. (1933) - Clas- sification and nomenclature of rock units. Geologi- cal Society of America Bulletin, 44, 423-459. Azzaroli A. (1977) - The Villafranchian stage in Italy and the Plio-Pleistocene boundary. Giornale Geologia, 41(2), 61-79. Azzaroli A. (1983) - Quaternary mammals and the “end- Villafranchian” dispersal event - a turning point in the history of Eurasia. Palaeogeography, Palaeo- climatology, Palaeoecology, 44, 117-139. Azzaroli A. (1995) - The “Elephant-Equus” and the “End- Villafranchian” Events in Eurasia. In E. S. Vrba, G.H. Denton, T.C. Partridge L.H. Burckle (eds.), Paleoclimate and Evolution, with Emphasis on Human Origins. Yale University Press, New Ha- ven, London, 311-318. Azzaroli A., De Giuli C., Ficcarelli G., Torre, D. (1988) - Late Pliocene to early mid-Pleistocene mammals in Eurasia: faunal succession and dispersal events. Palaeogeography, Palaeoclimatology, Palaeoecology, 66, 77-100. Badgle C., Finarelli J. A. (2013) - Diversity dynamics of mammals in relation to tectonic and climatic his- tory: comparison of three Neogene records from North America. Paleobiology, 39(3), 373-399. Barnosky A.D. (2005) - Effects of Quaternary climatic change on speciation in mammals. Journal of Mammalian Evolution, 12(1-2), 247-264. Barnosky A.D., Hadly E.A., Bell C.J. (2003) - Mammal- ian response to global warming on varied temporal scales. Journal of Mammalogy, 84(2), 354-368. Barnosky A.D., Kraatz B.P. (2007) - The role of climatic change in the evolution of mammals. Bioscience, 57(6), 523-532. Bellucci L., Sardella R., Rook L. (2015) - Large mammal biochronology framework in Europe at Jaramillo: The Epivillafranchian as a formal biochron. Qua- ternary International, 389, 84-89. Belmaker M. (2002) - First evidence of the presence of Theropithecus sp. in the Southern Levant. Israel J. Zool. 48 (2), 165. Berger G.W., Pérez-González A., Carbonell E., Arsuaga J.L., Bermúdez de Castro J.M., Ku T.L. (2008) - Luminescence chronology of cave sediments at the Atapuerca paleoanthropological site, Spain. Journal of Human Evolution, 55 (2), 300-311 Berger W.H., Jansen E. (1994) - Mid-Pleistocene Cli- mate Shift: The Nansen Connection. In Johannes- REFERENCES Abbazzi L. (2010) - La fauna de cérvidos de Barranco León y Fuente Nueva-3. In: Martínez-Navarro B., Agusti J., Toro Mojano I. (Eds), Ocupaciones hu- manas en el Pleistoceno Inferior y Medio de la cuenca de Guadix-Baza. Junta de Andalucía, Con- sejería de Cultura, 273-290. Abbazzi L. (2004) - Remarks on the validity of the ge- neric name Praemegaceros Portis 1920, and an overview on Praemegaceros species in Italy. Ren- diconti Lincei, 15(2), 115-132. Aguilar J.P., Antoine P.O., Bonnet A., Crochet J.Y., Michaux J. (2009) - Compléments à la faune pléis- tocène de Durfort (Gard, Sud de la France). Bulle- tin de la Société d'histoire naturelle de Toulouse, 145, 55-58. Agustí J. (1986) - Synthèse biostratigraphique du Plio- Pléistocène de Guadix-Baza 4 (Province de Gra- nada, sud-est de l’Espagne). Geobios, 19, 505- 510. Agustí J., Moya-Sola S. (1998) - The early Pleistocene mammal turnover in Spain: evidence against an 'End-Villfranchian' event. In: van Kolfschoten T.H., Gibbard P.L. (Eds), The Dawn of the Quaternary. (Proceedings of the SEQS-EuroMam symposium 1996), Mededelingen Nederlands Instituut voor Toegepaste Wetenschappen TNO, 60, 513-520. Alba D.M., Aurell-Garrido J., Madurell J., Gómez M., Moyà-Solà S., Beràstegui X. (2008) - Paleontolo- gia i geologia del jaciment del Pleistocè inferior de Vallparadís (Terrassa, Vallès Occidental). Trib. Arqueol., 2007, 29-44. Alberdi M.T. (2010) - Estudio de los caballos de los ya- cimientos de Fuente Nueva-3 y Barranco León-5 (Granada). In: Toro I., Martínez-Navarro B., Agustí J. (Eds), Ocupaciones humanas en el Pleistoceno inferior y medio de la Cuenca de Guadix-Baza. Arqueología Monografías, Junta de Andalucía, Consejería de Cultura, 291-306. Alberdi M.T., Palombo M.R. (2013) - The late Early to early Middle Pleistocene stenonoid horses from Italy. Quaternary International. 288, 25-44 Álvarez C., Parés J.M., Granger D., Duval M., Sala R., Toro I. (2015) - New magnetostratigraphic and numerical age of the Fuente Nueva-3 site (Guadix- Baza basin, Spain). Quaternary International 389, 224-234. Argant J., Bonifay M. F. (2011) - Les coprolithes de hyène (Pachycrocuta brevirostris) de la couche 2 du site villafranchién de Ceyssaguet (Lavoûte-sur- Loire, Haute-Loire, France): analyse pollinique et indications paléoenvironnementales. Quaternaire. Revue de l'Association française pour l'étude du Quaternaire, 22(1), 3-11. Arnold L. J., Demuro M., Parés J.M., Arsuaga J.L., Aran- buru A., de Castro J.M.B., Carbonell E. (2014) - Luminescence dating and palaeomagnetic age constraint on hominins from Sima de los Huesos, Atapuerca, Spain. Journal of human evolution, 67, 85-107. Arribas A., Garrido G. (2008) - Los representantes más antiguos del género Capra (Bovidae, Artiodactyla, 159 Late Early Pleistocene mammalian assemblages from SW Europe ............ sen et al. (eds.), The polar oceans and their role in shaping the Global Environment, Washinghton DC, AGU Geophysical Monograph, 84, 295-311. Blain H.A., Bailon S., Agustí J. (2008) - Amphibians and squamate reptiles from the latest Early Pleisto- cene of Cueva Victoria (Murcia, southeastern Spain, SW Mediterranean): paleobiogeographical and paleoclimatic implications. Geologica Acta, 6, 345-361. Blasco R., Rosell J., Van der Made J., Rodríguez J., Campeny G., Arsuaga J.L., Bermúdez de Castro J.M., Carbonell E. (2011) - Hiding to eat: the role of carnivores in the early Middle Pleistocene from the TD8 level of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). Journal of Archaeological Sci- ence, 38 (1-2), 3373-3386 Blois J.L., Zarnetske P.L., Fitzpatrick M.C., Finnegan S. (2013) - Climate change and the past, present, and future of biotic interactions. Science, 341 (6145), 499-504. Bon M., Piccoli G., Sala B. (1992) - La fauna pleistoce- nica della breccia di Slivia (Carso Triestino) nella collezione del Museo civico di Storia naturale di Trieste. Atti del Museo civico di Storia naturale di Trieste, 44, 33-51. Bonifay M.F. (1971) - Carnivores quaternaires du sud- est de la France. Mémoire du Muséum national d’Histoire Naturelle Paris. 21, 43-377. Bonifay M.F. (1981) - Les Praemegaceros du Pleisto- cene moyen de la grotte de l'Escale à Saint- Estève-Janson (Bouches-du-Rhône). Bull. Ass. Fr. Et. Quat., 3/4, 109-120. Bonifay M.F. (1991) - Archéologie du comportement: remarques sur l'apport des faunes de grands mammifères au Pléistocène ancien. Actes du Congrès National des Sociétés Savantes. Com- mission de Pré-et Protohistoire, 114, 111-113. Bonifay M.F. (1978) - Faunes de transition du Pléis- tocène moyen de France. Bulletin du Museum d’Anthropologie et Préhistoire de Monaco, 22, 5- 15. Bonvalot J., Courel L., Senac P. (1984) - Etude sèdi- mentologique du remplissage plio-pléistocène de la Bresse. Gèologie de la France, 3, 197-220 Boscaini A (2014) - Plio-Pleistocene lynxes from the Iberian Peninsula. Taxonomy and phylogenetic relationships with the extant and fossil species. Master dissertation, Universitat Autonoma de Bar- celona and Universitat de Barcelona, pp. 106. Boudier F. (1961) - Le Bassin du Rhône au Quaternaire. Géologie et préhistoire. Tome I. Texte. Centre National de la Recherche Scientifique, Paris, 364. Bourguignon L., Crochet J.Y., Capdevila R., Ivorra J., Antoine P.O., Agusti J., Barsky D., Blain H.A., Boulbes N., Claude J., Cochard D., de Weyer L., Filoux A., Firmat C., Lozano-Fernández I., Mag- niez P., Pelletier M., Ríos J., Testu A., Valensi P. (2014) - The Early Pleistocene site of Bois-De- Riquet (Lézignan-la-Cébe, Hérault, France): strati- graphy, dating, fauna, and lithics. Abstracts XVII World UISPP Congress 2014, Burgos, 1-7 Sep- tember, A2a The first peopling of Europe, Burgos, 23-24. Bourguignon L., Crochet J.Y., Capdevila R., Ivorra J., Antoine P.O., Agustí J., Barsky D., Blain H.A., Boulbes N., Bruxelles L., Claude J. (2015) - Bois- de-Riquet (Lézignan-la-Cèbe, Hérault): A late Early Pleistocene archeological occurrence in southern France. Quaternary International, 393 24 -40. Breda, M., Marchetti, M. (2005) - Systematical and bio- chronological review of Plio-Pleistocene Alceini (Cervidae; Mammalia) from Eurasia. Quaternary Science Reviews, 24(5), 775-805. Brugal J.P. (1994) - Le bison (Bovidae, Artiodactyla) du Pléistocène moyen ancien de Durfort (Gard, France). Bulletin du Muséum national d'histoire naturelle. Section C, Sciences de la terre, paléon- tologie, géologie, minéralogie, 16(2-4), 349-381. Brugal J.P., Boudadi-Maligne M. (2011) - Quaternary small to large canids in Europe: Taxonomic status and biochronological contribution. Quaternary International, 243 (1), 171-182. Bukhsianidze M. (2005) - The fossil Bovidae of Dmanisi, PhD, dissertation. The International Doctorate “Environmental, Humans and Compartmental Dy- namics” XVI cycle 2001-2004, University of Ferrara, pp. 192. Bukhsianidze M., Vekua, A. (2006) - Capra dalii nov. sp. (Caprinae, Bovidae, Mammalia) at the limit of Plio- Pleistocene from Dmanisi (Georgia). Courier- Forshungsinstitut Senckenberg, 256, 159. Caloi L., Palombo M.R. (1995) - Late Early Pleistocene mammal faunas of Italy: Biochronological prob- lems. Il Quaternario, 8 (2), 391-402. Carbonell E., Bermúdez de Castro J.M., Parés J.M., Pérez-González A., Cuenca-Bescós G., Ollé A., Mosquera M., Huguet R., Van der Made J., Rosas A., Sala R., Vallverdú J., García N., Granger D.E., Martinón-Torres M., Rodríguez X.P., Stock G.M., Vergès J.M., Allué E., Burjachs F., Cáceres I., Canals A., Benito A., Díez C., Lozano M., Mateos A., Navazo M., Rodríguez J., Rosell J., Arsuaga J.L. (2008) - The first hominin of Europe. Nature, 452, 465-469. Chaline J. (1972) - Les Rongeurs du Pléistocène Moyen et Supérieur de France 3 (Systématique, Biostrati- graphie, Paléoclimatologie). Cahiers de Paléon- tologie, 4 C.N.R.S., Paris, 410, 5 Chaline J. (1987) - Arvicolid data (Arvicolidae, Rodentia) and evolutionary concepts. Evolutionary Biology, 21, 237-310. Clark P., Archer D., Pollard D., Blum J.D., Rial J.A., Brovkin V., Mix A.C., Pisias N.G., Roy M. (2006) - The middle Pleistocene transition: characteristics, mechanisms, and implications for long-term changes in atmospheric pCO2. Quaternary Science Reviews, 25, 3150-3184. Colbert, J., Baguette M., Benton T.G., Bullock J.M. (Eds) (2012) - Dispersal Ecology and Evoution, Oxford University Press, Oxford UK, pp. 462. Collinge S. E., Parfitt S. A., Lister A. M. (2010) - Metric analysis of ungulate mammals in the early Middle Pleistocene of Britain, in relation to taxonomy and biostratigraphy: I: Rhinocerotidae and Bovidae. Quaternary International, 228(1), 136-156. 160 M.R. Palombo 119. Cuenca-Bescós G., Gracía N. (2007) - Biostratigraphic succession of the Early and Middle Pleistocene mammal faunas of the Atapuerca cave sites (Burgos, Spain). Courier-Forschungsinstitut Senckenberg, 259, 99-110. Cuenca-Bescós G., Rofes J., López-García J.M., Blain H-A., Rabal-Garcés R., Sauqué V., Arsuaga J.L., Bermúdez de Castro J-M., Carbonell E. (2011) - The small mammals of Sima del Elefante (Atapuerca, Spain) and the first entrance of Homo in Western Europe Quaternary International, 285 (8), 28-35 Dabney J., Knapp M., Glocke I., Gansauge M.T., Weihmann A., Nickel B., Valdiosera C., García N., Pääbo S., Arsuaga J.L., Meyer M., (2013) - Com- plete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ul- trashort DNA fragments. Proceedings of the Na- tional Academy of Sciences, 110(39), 15758- 15763. Davies T.J., Buckley L.B., Grenyer R., Gittleman J.L. (2011) - The influence of past and present climate on the biogeography of modern mammal diversity. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 366(1577), 2526- 2535. De Giuli C., Masini F., Torre D. (1987) - The latest Villa- franchian faunas in Italy: the Pirro Nord Fauna (Apricena, Gargano). Palaeontographia Italica, 74, 56-62. de Lumley H.D., Kahlke H.D., Moigne A.M., Moulle P.E. (1988) - Les faunes de grands mammifères de la grotte du Vallonnet, Roquebrune-Cap-Martin, Alpes-Maritimes. L'Anthropologie, 92(2), 465-495. Delson E. (1993) - Theropithecus fossils from Africa and India and the taxonomy of the genus. In: Jablonski N.G. (Ed.), Theropithecus: The Rise and Fall of a Primate Genus. Cambridge University Press, Cambridge, 157-189. Di Stefano G., Petronio C. (1992) - Nuove osservazioni su Cervus elaphus acoronatus Beninde del Pleis- tocene europeo. Bollettino della Societa` Paleon- tolologica Italiana, 31(3), 295-315. Duval M., Aubert M., Hellstrom J., Grün R. (2011a) - High resolution LA-ICP-MS mapping of U and Th isotopes in an early Pleistocene equid tooth from Fuente Nueva-3 (Orce, Andalusia, Spain). Quater- nary Geochronology, 6(5), 458-467. Duval M., Falguères C., Bahain J.J., Grün R., Shao Q., Aubert M., Hellstrom, J., Dolo J.M., Agusti J., Martínez-Navarro B., Palmqvist P., Toro-Moyano I. (2011b) - The challenge of dating Early Pleisto- cene fossil teeth by the combined uranium series- electron spin resonance method: the Venta Mi- cena palaeontological site (Orce, Spain). Journal of Quaternary Science, 26(6), 603-615. Duval M., Falguères C., Bahain J.J., Grün R., Shao Q., Aubert M., Dolo J.M., Agusti J., Martínez-Navarro B., Palmqvist P., Toro-Moyano I. (2012) - On the limits of using combined U-series/ESR method to date fossil teeth from two Early Pleistocene ar- chaeological sites of the Orce area (Guadix-Baza Coltorti M., Albianelli A., Bertini A., Ficcarelli G., Laurenzi, M. A., Napoleone G., Torre D. (1998) - The Colle Curti mammal site in the Colfiorito area (Umbria-Marchean Apennine, Italy): geomorphol- ogy, stratigraphy, paleomagnetism and palynol- ogy. Quaternary International, 47, 107-116. Combourieu-Nebout N., Bertini A., Russo-Ermolli E., Peyron O., Klotz S., Montade V., Fauquette S., Allen J.R.M., Fusco F., Goring S., Huntley B. (2015) - Climate changes in the central Mediterra- nean and Italian vegetation dynamics since the Pliocene. Review of Palaeobotany and Palynol- ogy, 2018, 127-147. Crégut-Bonnoure E. (2002) - Les Ovibovini, Caprini et Ovini (Mammalia, Artiodactyla, Bovidae, Caprinae) du Plio-Pléistocène dʼEurope: systématique, évo- lution et biochronologie. Thèse de Doctorat d’Etat és Sciences. Université de Lyon I, pp. 430. Crégut-Bonnoure E. (2005) - Nouvelles données paléogéographiques et chronologiques sur les Caprinae (Mammalia, Bovidae) du Pléistocène moyen et supérieur d’Europe. Munibe Antropol- Arkeol, 57, 205-219. Crégut-Bonnoure E. (2007) - Apport des Caprinae et Antilopinae (Mammalia, Bovidae) à la biostrati- graphie du Pliocène terminal et du Pléistocène d’Europe. Quaternaire. Revue de l'Association française pour l'étude du Quaternaire, 18(1), 73- 97. Crégut-Bonnoure E., Dimitrijevic, V. (2006) - Megalovis balcanicus sp. nov. and Soergelia intermedia sp. nov.(Mammalia, Bovidae, Caprinae), new Ovibo- vini from the Early Pleistocene of Europe. Revue de Paléobiologie, 25(2), 723-773. Crochet J.Y., Welcomme J.L., Ivorra J., Ruffet G., Boul- bes N., Capdevila R., Claude J., Firmat C., Métais G., Michaux J., Pickford M. (2009) - Une nouvelle faune de vertébrés continentaux, associée à des artifacts dans le Pléistocène inférieur de l’Hérault (Sud de la France), ver 1, 57 Ma. Comptes Ren- dus Palevol., 8, 725-736. Croitor R. (2006) - Taxonomy and systematics of large- sized deer of the genus Praemegaceros Portis, 1920 (Cervidae, Mammalia). Courier- Forshungsinstitut Senckenberg, 256, 91-116. Croitor R. (2009) - Systematical Position and Evolution of the Genus Arvernoceros (Cervidae, Mammalia) from Plio-Pleistocene of Eurasia. Muz. Olteniei Craiova. Studii şi comunic ąri. Ştiin ţele Natur, 25, 379-381. Croitor R., Bonifay M.F. (2001) - Étude préliminaire des cerfs du gisement Pleistocène inférieur de Ceys- saguet (Haute-Loire) - Paleo, 13, 129-144. Croitor R., Kostopoulos, D.S. (2004) - On the systematic position of the large-sized deer from Apollonia, Early Pleistocene, Greece. Paläontologische Zeitschrift, 78(1), 137-159. Cuenca-Bescós G., Rofes J., López-García M., Blain H.- A., de Marfà R., Galindo-Pellicena M.A., Bennàsar -Serra M.L., Melero-Rubio M., Arsuaga J.L, Ber- múdez de Castro J.M., Carbonell. E. (2010) - Bio- chronology of Spanish 17 Quaternary small verte- brate faunas. Quaternary International, 212, 109- 161 Late Early Pleistocene mammalian assemblages from SW Europe ............ basin, Spain). Quaternary Research, 77(3), 482- 491. Duval M., Bahain J.J., Falguéres C., Gracía J., Guilarte V., Grün R., Martínez K., Moreno D., Shao, Q., Voinchet P. (2015) - Revisiting the ESR chronol- ogy of the Early Pleistocene hominin occupation at Vallparadís (Barcelona, Spain). Quaternary Inter- national, 389, 213-223. Eisenmann V., Kuznetsova T. (2004) - Early Pleistocene equids (Mammalia, Perissodactyla) of Nalaikha, Mongolia, and the emergence of modern Equus Linnaeus, 1758. Geodiversitas, 26(3), 535-561. Espigares M.P. (2010) - Análisis y modelización del contexto sedimentario y los atributos tafonómicos de los yacimientos pleistocénicos del borde norori- ental de la cuenca de Guadix-Baza. Ph. D. Disser- tation, University of Granada (Spain), Fernández M.H., Cantalapiedra J.L., Cano A.R.G. (2015) - Plio-Pleistocene climatic change had a major impact on the assembly and disassembly processes of Iberian rodent communities. Palaeo- biodiversity and Palaeoenvironments, 95, 387- 404. Ferràndez-Cañadell, C., Ribot F., Gibert L. (2014) - New fossil teeth of Theropithecus oswaldi (Cercopithecoidea) from the Early Pleistocene at Cueva Victoria (SE Spain). Journal of human evo- lution 74, 55-66. Fortelius M., Eronen J.T., Kaya F., Tang H., Raia P., Puolamäki K. (2014) - Evolution of Neogene mam- mals in Eurasia: environmental forcing and biotic interactions. Annual Review of Earth and Plane- tary Sciences, 42, 579-604. Franzen J.L., Gliozzi E., Jellinkek T., Scholger R., Wei- denfeller M. (2000) - Die spätpleistozäne Fossil- lagerstätte Dorn-Dürkheim 3 und ihre Bedeutung für die Rekonstruktion der Entwicklung des rhen- ishen Flusssystems. Senkenbergiana Lethaea, 80 (1), 305-353. Gracía N., Arsuaga, J.L. (2001) - Les carnivores (Mammalia) des sites du Pléistocène ancien et moyen d'Atapuerca (Espagne). L’Anthropologie, 105(1), 83-93. Garcıa N., Arsuaga J.L., Torres T. (1997) - The carni- vore remains from the Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain). Journal of Human Evolution, 33(2), 155-174. Geraads D. (1990) - Contribution des cervidés à la chro- nologie des débuts de l'occupation humaine en Europe occidentale. Quaternaire, 1(3), 167-174. Geraads D. (2010) - Biogeographical relationships of Pliocene and Pleistocene North-western African mammals. Quaternary International, 212(2), 159- 168. Ghezzo E., Boscain, A., Madurell-Malapeira J., Rook L. (2014) - Lynx remains from the Pleistocene of Valdemino cave (Savona, Northwestern Italy), and the oldest occurrence of Lynx spelaeus (Carnivora, Felidae). Rendiconti Lincei, 26, 87-95. Gibert J., Ribot F., Gibert L., Leakey M., Arribas A., Martínez-Navarro B. (1995) - Presence of the cercopithecid genus Theropithecus in Cueva Vic- toria (Murcia, Spain). Journal of Human Evolution, 28, 487-493. Gibert L., Scott G. (2015) - Etad del yacimiento de Cueva Victoria y su relacion con otros yacimientos de la Peninula Ibérica. In Gibert L., Ferrandez- Canadell C., (Eds), Paleontología y Geología de Cueva Victoria, Cartagena, España. Mastia, 11- 13, 85-100 Gingerich P.D. (2003) - Mammalian responses to cli- mate change at the Paleocene-Eocene boundary: Polecat Bench record in the northern Bighorn Ba- sin, Wyoming. Special Papers, Geological Society of America, 463-478. Gliozzi E., Abbazi L., Ambrosetti P., Argenti, P., Azzaroli A., Caloi L., Capasso Barbato L., Di Stefano G., Esu D., Ficcarelli G., Girotti O., Kotsakis T., Masini F., Mazza P., Mezzabotta C., Palombo M.R., Petronio C., Rook L., Sala B., Sardella R., Zanalda E., Torre D. (1997) - Biochronology of selected Mammals, Molluscs, Ostracods from the Middle Pliocene to the Late Pleistocene in Italy. The state of the art. Rivista Italiana di Paleontologia e Strati- grafia, 103 (3), 369-388. Grossouvre A.D., Stehlin H.G. (1912) - Les sables de Rosières, près Saint-Florent (Cher) - Bulletin de la Société géologique de France, 4, 194-212. Guérin C. (1990) - Biozones or mammal units? Methods and limits in biochronology. In: Lindsay, E.H., Fahl- bush, V., Mein, P. (Eds.), European Neogene Mammalian Chronology. NATO ASI Series a Life Sciences, 180. Plenum Press, New York, 119-130. Guérin C., Dewolf Y., Lautridou J. P. (2003) - Révision d'un site paléontologique célèbre: Saint-Prest (Chartres, France). Geobios, 36(1), 55-82. Guérin C., Faure M. (1997) - The wild boar (Sus scrofa priscus) from the post-Villafranchian Lower Pleis- tocene of Untermassfeld. In Kahlke, R.D. et al. (eds), Das Pleistozän von Untermassfeld bei Mein- ingen (Thüringen), Teil 1. Monogr. Röm.-Germ. Zentralmus. Mainz, 40(1), 375-384. Hassanin A., Ropiquet A., Couloux A., Cruaud C. (2009) - Evolution of the mitochondrial genome in mam- mals living at high altitude: new insights from a study of the tribe Caprini (Bovidae, Antilopinae). Journal of molecular evolution, 68(4), 293-310. Head M.H., Gibbard, P.O. (Eds) (2005) - The Land- Ocean Early-Middle Pleistocene Transitions: Evi- dence. Geological Society of London Special Pub- lication 251. The Geological Society Publishing House, Bath UK, pp. 336. Hedberg H.D. (Ed) (1976) - International Stratigraphic Guide. John Wiley & Sons, New York, pp. 200. Heintz E. (1970) - Les Cervidés villafranchiens de France et d'Espagne. Mémoires du Muséum na- tional d’Histoire Naturelle de Paris C 22, pp. 206. Hemmer H. (2001) - Die Felidae aus dem Epivillafran- chium von Untermassfeld. Monographien des Römisch-Germanischen Zentralmuseum, 40, 699- 782. Hemmer H., Kahlke R.D., Vekua A.K. (2011) -The chee- tah Acinonyx pardinensis (Croizet et Jobert, 1828) s.l. at the hominin site of Dmanisi (Georgia) - a potential prime meat supplier in Early Pleistocene ecosystems. Quat. Sci. Rev., 30, 2703-2714. 162 M.R. Palombo ont. Italiana, 40(2), 217-223. Koufos G.D. (2014) - The Villafranchian carnivoran guild of Greece: implications for the fauna, biochronol- ogy and paleoecology. Integrative zoology, 9(4), 444-460. Koufos G.D., Kostopoulos D.S., Sylvestrou I.A. (1997) - Equus apolloniensis n.sp. (Mammalia, Equidae) from the latest Villafranchian locality of Apollonia, Macedonia, Greece. Paleontologia i Evoluciò, 30- 31, 49-76. Kurtén B. (1968) - Pleistocene Mammals of Europa. Weidenfeld and Nicolson, London, pp. 317. Kurtén B., Poulianos A. (1977) - New stratigraphic and faunal material from Petralona cave with special reference to the carnivora. Anthropos Athens, 4, 47-130. Langlois A. (2002) - Présence de Panthera gom- baszoegensis Kretzoi, 1938 à la Grotte XIV (Cénac-et-Saint-Julien, Dordogne). Paléo. Revue d'archéologie préhistorique, 14, 213-220. Lawler J.J., Shafer S.L., White D., Kareiva P., Maurer E.P., Blaustein A.R., Bartlein P.J. (2009) - Pro- jected climate-induced faunal change in the West- ern Hemisphere. Ecology, 90(3), 588-597. Leroy S.A.G., Arpe K., Mikolajewicz U. (2011) - Vegeta- tion context and climatic limits of the Early Pleisto- cene hominin dispersal in Europe. Quaternary Science Reviews, 30(11), 1448-1463. Lindsay E.H. (1990) - The setting. In: Lindsay E.H., Fahlbush V., Mein P. (Eds), European Neogene Mammalian Chronology. NATO ASI Series a Life Sciences, 180. Plenum Press, New York, 1-14. Lindsay E.H. (2003) - Chapter 10: Chronostratigraphy, Biochronology, Datum Events, Land Mammal Ages, Stage of Evolution, and Appearance Event Ordination. Bulletin of the American Museum of Natural History, 212-230. Lindsay E.H., Tedford, R.H. (1990) -. Development and application of land mammal ages in North America and Europa, and comparison. In: Lindsay E.H., Fahlbusch, W., Mein, P. (Ed), European Neogene Mammalian Chronology. NATO ASI Series a Life Sciences, 180. Plenum Press, New York, 601-624. Lisiecki L, Raymo M. (2005) - A Pliocene-Pleistocene stack of 57 globally distributed benthic 18O re- cords. Paleoceanography, 20, PA1003. Lister A.M., Parfitt S.A., Owen F.J., Collinge S.E., Breda M. (2010) - Metric analysis of ungulate mammals in the early Middle Pleistocene of Britain, in rela- tion to taxonomy and biostratigraphy: II: Cervidae, Equidae and Suidae. Quaternary International, 228(1), 157-179. Lister A.M., Sher A.V., van Essen H., Wei G. (2005) - The pattern and process of mammoth evolution in Eurasia. Quaternary International, 126-128, 49-64. Lister A.M. (1993) - The stratigraphical significance of deer in the Cromer Forest-bed Formation. Journal of Quaternary Science, 8 (2), 95-108. Lo Bello P. (1988) - Géochronologie par la méthode 39Ar -40Ar de ponces quaternaires contaminées: Exem- ple des ponces du Mont-Dore (Massif Central, France). Utilisation d’un laser continu pour la data- tion des minéraux individuels. Thèse de Doctorat Hemmer H., Kahlke R.D., Keller T. (2008) - Cheetahs in the Middle Pleistocene of Europe: Acinonyx pardinensis (sensu lato) intermedius (Thenius, 1954) from the Mosbach Sands (Wiesbaden, Hesse, Germany). Neues Jahrb. Geol. Paläontol. Abh., 249, 345-356 Hooijer D.A. (1959) - Fossil mammals from Jisr banat Yaqub, south of Lake Huleh, israel. Bulletin of the Research Council of Israel, Section G: Geo- sciences, 8(4), 177-199. Hughes J.K., Elton S., O'Regan H.J. (2008) - Theropith- ecus and ‘Out of Africa’dispersal in the Plio- Pleistocene. Journal of Human Evolution, 54(1), 43-77. Kahlke H.D. (1997) - Die Cerviden-Reste aus dem Unterpleistozän von Untermassfeld. In Kahlke, R.D. et al. Das Pleistozän von Untermassfeld bei Meiningen (Thüringen), Teil 1. Monogr. Röm.- Germ. Zentralmus. Mainz 40(1), 181-275 Kahlke H.D. (2001) - Neufunde von Cerviden- Resten aus dem Unterpleistozän von Untermassfeld. In Kahlke, R.D. et al. (Eds), Das Pleistozän von Untermassfeld bei Meiningen (Thüringen), Teil 2. Monogr. Röm.-Germ. Zentralmus. Mainz 40(2), 185-195 Kahlke R.D. (2000) - The Early Pleistocene (Epivillafranchian) Faunal Site of Untermassfeld (Thuringia, Central Germany) - Synthesis of New Results. Early humans at the gate of Europe. ERAUL, 92, 123 -138. Kahlke R.D. (2006) - Untermassfeld - a late Early Pleis- tocene (Epivillafranchian) fossil site near Meinin- gen (Thuringia, Germany) and its position in the development of the European mammal fauna. British Archaeological Reports, International Se- ries, 1578, 1-144. Kahlke R.D. (2007) - Late Early Pleistocene European large mammals and the concept of an Epivillafran- chian Biochron. Courier Forschungsinstitut Senck- enberg, 20(259), 265-278. Kahlke R.D. (2009) - Les communautés de grands mammifères du Pléistocène inférieur terminal et le concept d'un biochrone Épivillafranchien. Quater- naire, 20(4), 415-427. Kahlke R.D., García N., Kostopoulos D.S., Lacombat F., Lister A.M., Mazza P.P., Spassov N. Titov V.V. (2011) - Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleis- tocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews, 30 (11), 1368-1395. Konidaris G.E., Tourloukis V., Kostopoulos D.S., Thompson N., Giusti D., Michailidis D., Koufos G., Harvati K. (2015) - Two new vertebrate localities from the Early Pleistocene of Mygdonia Basin (Macedonia, Greece): Preliminary results. Comptes Rendus Palevol, 14(5), 353-362. Kostopoulos D.S. (1997) - The Pilo-Pleistocene artio- dactyls (Vertebrata, Mammalia) of Macedonia 1. The fossiliferous site Apollonia-1, Mygdonia basin of Greece. Geodiversitas, 19(4), 845-875. Koufos G. (2001) - The Villafranchian mammalian fau- nas and biochronology of Greece. Boll. Soc. Pale- 163 Late Early Pleistocene mammalian assemblages from SW Europe ............ de l’Université de Nice, pp. 122. Lo Bello P., Feraud G., Hall C.M., York D., Lavina P., Bernat M. (1987) - 40Ar39Ar step-heating and laser fusion dating of a quaternary pumice from Ne- schers, Massif Central, France: The defeat of xenocrystic contamination. Chemical Geology: Isotope Geoscience section, 66(1), 61-71. Lozano-Fernández I., Cuenca-Bescós G., Blain H.A., López-García J.M., Agustí J. (2013) - Mimomys savini size evolution in the Early Pleistocene of south-western Europe and possible biochronologi- cal implications. Quaternary Science Reviews, 76, 96-101. Lozano-Fernández I., López-García J.M., Aurell-Garrido J., Alba D.M., Madurell-Malapeira J. (2015) - Data review on the small mammals from the late Early Pleistocene of Vallparadís Estació layer EVT7 (Vallès-Penedès Basin, NE Iberian Peninsula): Biochronological and palaeoenvironmental impli- cations. Quaternary International, 389, 159-166. Madurell-Malapeira J., Minwer-Barakat R., Alba D., Gar- cés M., Gómez M., Aurell- Garrido J., Ros- Montoya S., Moyà-Solà, S., Berástegui X. (2010a) - The Vallparadís section (Terrassa, Iberian Penin- sula) and the latest Villafranchian faunas of Europe. Quaternary Science Reviews, 29, 3972- 3982. Madurell-Malapeira J., Alba D. M., Espigares M.P., Vinuesa V., Palmqvist P., Martínez-Navarro B., Moyà-Solà S. (2015) - Were large carnivorans and great climatic shifts limiting factors for hominin dispersals? Evidence of the activity of Pachycro- cuta brevirostris during the Mid Pleistocene Revo- lution in the Vallparadís Section (Vallès-Penedès Basin, Iberian Peninsula), Quaternary Interna- tional (in press), doi:10.1016/j.quaint.2015.07.040 Madurell-Malapeira J., Alba D. M., Moyà-Solà S., Aurell- Garrido J. (2010b) - The Iberian record of the puma-like cat Puma pardoides (Owen, 1846) (Carnivora, Felidae). Comptes Rendus Palevol, 9 (1), 55-62. Madurell-Malapeira J., Ros-Montoya S., Espigares M.P., Alba D.M., Aurell-Garrido J.A. (2014) - Villafran- chian large mammals from the Iberian Peninsula: paleobiogeography, paleoecology and dispersal events. Journal of Iberian Geology, 40(1), 141- 155. Magri D., Palombo M.R. (2013) - Early to middle Pleisto- cene dynamics of plant and mammal communities in South West Europe. Quaternary International, 288, 63-72. Marciszak A. (2014) - Presence of Panthera gom- baszoegensis (Kretzoi, 1938) in the late Middle Pleistocene of Biśnik Cave, Poland, with an over- view of Eurasian jaguar size variability. Quater- nary International, 326, 105-113. Martínez-Navarro B., Madurell-Malapeira J., Ros- Montoya S., Espigares, M.P., Medin T., Hortola P., Palmqvist P. (2015) - The Epivillafranchian and the arrival of pigs into Europe. Quaternary Interna- tional, 389, 131-138. Martínez-Navarro B., Rook L. (2003) - Gradual evolution in the African hunting dog lineage systematic im- plications. Comptes Rendus Palevol, 2(8), 695- 702. Martínez-Navarro B., Ros-Montoya S., Espigares M.P., Palmqvist P. (2011) - Presence of the Asian origin Bovini, Hemibos sp. aff. Hemibos gracilis and Bi- son sp., at the early Pleistocene site of Venta Mi- cena (Orce, Spain). Quaternary International, 243 (1), 54-60. Martínez-Navarro B., Sardella R., Rook L., Bellucci L., Ros-Montoya S. (2012) - First occurrence of Soer- gelia (Ovibovini, Bovidae, Mammalia) in the Early Pleistocene of Italy. Quaternary International, 267, 98-102. Masini F., Palombo M.R., Rozzi R. (2013) - A reap- praisal of the Early to Middle Pleistocene Italian bovidae. Quaternary International, 288, 45-62. Masini F., Sala B. (2007) - Large-and small-mammal distribution patterns and chronostratigraphic boundaries from the Late Pliocene to the Middle Pleistocene of the Italian peninsula. Quaternary International, 160 (1), 43-56. Maslin M.A., Ridgwell A.J. (2005) - Mid-Pleistocene revolution and the ‘eccentricity myth’. Geological Society, London, Special Publications, 247(1), 19- 34. Maslin, M.A, Ridgwell A.J. (2005) - Mid-Pleistocene revolution and the ‘eccentricity myth’. Geological Society, London, Special Publications, 247(1), 19- 34. Maul L.C., Markova A.K. (2007) - Similarity and regional differences in Quaternary arvicolid evolution in Central and Eastern Europe. Quaternary interna- tional, 160(1), 81-99. Maul L.C., Parfitt S.A. (2010) - Micromammals from the 1995 Mammouth excavation at West Runton, Nor- folk, UK: Morphometric data, biostratigraphy and taxonomic reappraisal. Quaternary International, 228 91-115. 15 Mayhew D.F. (2013) - West European arvicolid evidence of intercontinental connections during the Early Pleistocene. Quaternary International, 284, 62-73. Mazo A.V., Sesé C., Ruiz Bustos A., Peña J.A. (1985) - Geología y Paleontología de los yacimientos plio- pleistocenos de Huéscar (Depresión de Guadix- Baza, Granada). Estudios Geológicos 41, 467- 493. McClymont E.L., Sosdian S.M., Rosell-Melé A., Rosen- thal Y. (2013) - Pleistocene sea-surface tempera- ture evolution: Early cooling, delayed glacial inten- sification, and implications for the mid-Pleistocene climate transition. Earth-Science Reviews, 123, 173-193 Menéndes E. (1987) - Cérvidos del yacimiento del Pleis- toceno inferior de Venta Micena_2, Orce (Granada, España), Paleontol. Evolucio, 1, 129- 181. Meon H., Ballesio R., Guérin C., Mein P. (1979) - Corré- lations biostratigraphiques dans le Néogène du sud-est de la France. Ann. Géol. Pays hellén., 807 -816. Minwer-Barakat R., Madurell-Malapeira J., Alba D. M., Aurell-Garrido J., de Estevan-Trivigno S., Moyà- Solà S. (2011) - Pleistocene rodents from the Tor- 164 M.R. Palombo Bailon S. (2005) - L'environnement animal des premiers habitants de l'Europe méditerranéenne: les grands mammifères contemporains de l'homme du Vallonnet, données taxonomiques et biostratigraphiques pour la deuxième moitie du Pléistocène inférieur. BAR International Series, 1364, 105-113. Musil R. (2001) - Die Equiden-reste aus dem Unterpleis- tozän von Untermassfeld. In: Kahlke, R.-D (Ed.), Das Pleistozän von Untermassfeld bei Meiningen (Thüringen), Teil 2. Monograph. Römish-German. Zentralmus., 40 (2), 633-658 Napoleone G., Albianelli A., Azzaroli A., Bertini A., Magi M., Mazzini M. (2003) - Calibration of the Upper Valdarno basin to the Plio-Pleistocene for correlat- ing the Apennine continental sequences. Il Quater- nario, Italian Journal of Quaternary Sciences 16 (1Bis), 159-195. Nocchi G., Sala B. (1997) - The fossil rabbit from Val- demino cave (Borgio Verezzi, Savona) in the con- text of western Europe Oryctolagini of Quaternary. Palaeovertebrata, 26(1-4), 167-187. Nomade S., Pastre J.F., Guillou H., Faure M., Guérin C., Delson E., Debard E., Voinchet P., Messager E. (2014) - 40Ar/39Ar constraints on some French landmark Late Pliocene to Early Pleistocene large mammalian paleofaunas: Paleoenvironmental and paleoecological implications, Quaternary Geochro- nology 21, 2-15. O'Regan H.J. (2008) - The Iberian Peninsula-corridor or cul-de-sac? Mammalian faunal change and possi- ble routes of dispersal in the last 2 million years. Quaternary Science Reviews, 27(23), 2136-2144. Olive F. (2006) - Evolution des grands Carnivores au Plio Pléistocène en Afrique et en Europe occiden- tale. L'Anthropologie, 110(5), 850-869. Osborn H.F., Matthew W.D. (1909) - Cenozoic mammal horizons of western North America. U.S. Geologi- cal Survey Bulletin, 361, 1-138. Pacheco F.G., Santiago A., Gutiérrez J.M., López- García J.M., Blain H.A., Cuenca-Bescós G., Van der Made J., Caceres I., García, N. (2011) - The Early Pleistocene paleontological site in the sierra del chaparral (Villaluenga del Rosario, cádiz, Southwestern Spain. Quaternary International, 243 (1), 92-104. Palmqvist P., Arribas A., Martínez-Navarro B. (1999) - Ecomorphological study of large canids from the lower Pleistocene of southeastern Spain. Lethaia, 32(1), 75-88. Palmqvist P., González-Donoso J.M., De Renzi M. (2014) - Rectilinear evolution in arvicoline rodents and numerical dating of Iberian Early Pleistocene sites. Quaternary Science Reviews, 98, 100-109. Palombo M.R. (1995) - Gli elefanti del Pliocene superi- ore e del Pleistocene dell’Italia centrale peninsu- lare. Studi Geol. Camerti, vol. spec. B (1994), 447- 457. Palombo M.R. (2009) - Biochronology of terrestrial mammals and Quaternary subdivisions: a case study of large mammals from the Italian peninsula. Il Quaternario, Italian Journal of Quaternary Sci- ences, 22, 291-306. rent 20 de Vallparadís section (Terrasa, northeast- ern Spain) and biochronological 21 implications. Journal of Vertebrate Paleontology, 31, 849-865. Moigne A.M., Palombo M.R., Belda V., Heriech-Briki D., Kacimi S., Lacombat F., de Lumley M.A., Moutoussamy J., Rivals F., Quilès J., Testu A. (2006) - Les faunes de grands mammifères de la Caune de l'Arago (Tautavel) dans le cadre bio- chronologique des faunes du Pléistocène moyen italien. L'anthropologie, 110(5), 788-831. Mol D., Post K., Reumer J.W.F., de Vos J., Laban C. (2003) - Het Gat: preliminary note on a Bavelian fauna from the North Sea with possibly two mam- moth species. Deinsea, 9, 253-266. Mol D., Post K., Reumer J. W., de Vos J., Laban C. (2003) - Het Gat: preliminary note on a Bavelian fauna from the North Sea with possibly two mam- moth species. Advances in Mammoth Research. Deinsea, 9, 253-266. Moncel M. H., Despriée J., Voinchet P., Tissoux H., Moreno D., Bahain J. J., Courcimault G., Fal- guères C. (2013) - Early Evidence of Acheulean Settlement in Northwestern Europe-La Noira Site, a 700 000 Year-Old Occupation in the Center of France. PloS one 8(11), e75529. Montoya P., Alberdi M.T., Barbadillo L.J., Morales J., Murelaga X., Peñalver E., Robles F., Ruiz Bustos A., Sánchez A., Sanchiz B., Soria D., Szyndlar Z. (2001) - Une faune très diversifiée du Pléistocène inférieur de la Sierra de Quibas (province de Murcia, Espagne). Comptes Rendus de l’Acadé- mie des sciences Paris, 332, 387-393. Morales J., Soria D., Soto E. (1987) - Los carnívoros del Pleistoceno medio de Atapuerca. El hombre fósil de Ibeas y el Pleistoceno de la Sierra de Atapu- erca I. Junta de Castilla y León, Consejería de Cultura y Bienestar Social, Soria, 135-152. Moreno García D. (2011) - Datation par ESR de quartz optiquement blanchis (ESR-OB) de la région de Atapuerca (Burgos, Espagne). Application au site préhistorique de Gran Dolina (contexte karstique) et aux systèmes fluviatiles quaternaires de l'Arlan- zón et l'Arlanza.". PhD dissertation. Universitat Rovira i Virgili. Departament d'Història i Història de l'Art, Tarragona, Spain, pp. 305. Mosquera M., Ollé A., Saladié P., Cáceres I., Huguet R., Rosas A., Villalaín J. J., Carrancho A., Bourlès D., Braucher R., Pineda A., Vallverdú J. (2015) - The early Acheulean technology of Barranc de la Bo- ella (Catalonia, Spain). Quaternary International, 393, 95-111. Moullé P.E., Echassoux A., Martínez-Navarro B. (2004) - Ammotragus europaeus: une nouvelle espèce de Caprini (Bovidae, Mammalia) du Pléistocène in- férieur à la grotte du Vallonnet (France). Comptes- rendus Palevol, 3, 663-673. Moullé P.E., Lacombat F., Echassoux A. (2006) - Apport des grands mammifères de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes-Maritimes, France) à la connaissance du biochronologique de la seconde moitié du Pléistocene inférieur d’Eu- rope. L’Anthropologie, 110, 837-849. Moullé P.E., Echassoux A., Lacombat F., Desclaux E., 165 Late Early Pleistocene mammalian assemblages from SW Europe ............ Palombo M.R. (2014) - Deconstructing mammal disper- sals and faunal dynamics in SW Europe. Quater- nary Science Reviews, 96, 50-71. Palombo M.R. (2015a) - Discrete dispersal bioevents of large mammals in Southern Europe in the post- Olduvai Early Pleistocene: A critical overview. Quaternary International (in press). doi:10.1016/ j.quaint.2015.08.034 Palombo M.R. (2015b) - To what extent could functional diversity be an useful tool in inferring ecosystem responses to past climate changes? Quaternary International. ( in press) doi:10.1016/ j.quaint.2015.07.069 Palombo M.R., Ferretti M.P. (2005) - Elephant fossil record from Italy: knowledge, problems, and per- spectives. Quaternary International, 126-128, 107- 136. Palombo M.R., Milli S. (2011) - Mammalian fossil recod, depositional setting, and sequence stratigraphy in the Middle - Upper Pleistocene of Roman Basin. Il Quaternario, Italian Journal of Quaternary Sci- ences, 23 (2), 243-248. Palombo M.R., Valli A.M.F. (2004) - Remarks on the biochronology of mammalian faunal complexes from the Pliocene to the Middle Pleistocene in France. Geologica Romana, 37(2003-2004), 145- 163. Palombo M.R., Azanza B., Alberdi M.T. (2003) - Italian mammal biochronology from latest Miocene to Middle Pleistocene: a multivariate approach. Ge- ologica Romana 36 (2000-2002), 335-368. Palombo M.R., Alberdi M.T., Azanza B., Giovinazzo C., Prado J.L., Sardella R. (2009) - How did environ- mental disturbances affect carnivoran diversity? A case study of the Plio-Pleistocene Carnivora of the North-Western Mediterranean. Evolutionary Ecol- ogy, 23(4), 569-589. Palombo M.R., Mussi M., Agostini A., Barbieri M., Di Canzio E., Di Rita F., Fiore I., Iacumin P., Magri D., Speranza F., Tagiacozzo A. (2010) - Human peopling of Italian intramontane basins: The early Middle Pleistocene site of Pagliare di Sassa (L’Aquila, central Italy). Quaternary International, 223-224, 170-178. Parés J. M., Duval M., Arnold L. J. (2013) - New views on an old move: Hominin migration into Eurasia. Quaternary International 295, 5-12. Parfitt S. A., Barendregt R. W., Breda M., Candy I., Collins M. J., Coope G. R., Durbidge P., Field M.H., Lee J.R., Lister A.M., Mutch R., Penkman K.E.H., Preece R.C., James R., Stringer C.B., Symmons R., Whittaker J.E., Wymer J.J., Stuart A. J. (2005) - The earliest record of human activity in northern Europe. Nature, 438(7070), 1008- 1012. Parfitt S.A., Ashton N.M., Lewis S. G., Abel R.L., Coope G. R., Field M.H., Gale R., Hoare P.G., Larkin N.R., Lewis M.D., Karloukovski V., Maher B.A., Peglar S.M., Preece, R.C., Whittaker J.E., Stringer C.B. (2010) - Early Pleistocene human occupation at the edge of the boreal zone in northwest Europe. Nature, 466(7303), 229-233. Petronio C., Marcolini F. (2013) - Mammal Biochronol- ogy at the end of Late Villafranchian (Early Pleisto- cene): Pirro Faunal Unit. Palaeontographica Ab- teilung A, 298(1-6), 183-191. Petrucci M., Cipullo A., Martínez-Navarro B., Rook L., Sardella R. (2013) - The Late Villafranchian (Early Pleistocene) carnivores (Carnivora, Mammalia) from Pirro Nord (Italy) - Palaeontographica Ab- teilung A, 298(1-6), 113-145. Pons-Moya J. (1987) -. Los carnivoros (Mammalia) de Venta Micena (Granada, Espana) - Paleontologia i evolucio Memoir special 1, 109-128. Prideaux G.J., Roberts R.G., Megirian D., Westaway K.E., Hellstrom J.C., Olley J.M. (2007) - Mammal- ian responses to Pleistocene climate change in southeastern Australia. Geology, 35(1), 33-36. Prothero D.R. (2004) - Did impacts, volcanic eruptions, or climate change affect mammalian evolution?. Palaeogeography, Palaeoclimatology, Palaeoecol- ogy, 214(3), 283-294. Prothero D.R. (2012) - Cenozoic mammals and climate change: the contrast between coarse-scale versus high-resolution studies explained by species sort- ing. Geosciences, 2(2), 25-41. Prothero D.R. (2014) - Species longevity in North Ameri- can fossil mammals. Integrative zoology, 9(4), 383 -393. Qiu Z.X. (2006) - Quaternary environmental changes and evolution of large mammals in North China. Vertebrata PalAsiatica, 44, 109-132. Raynal J.P., Magoga L., Bulle T., Guadelli J.L., Maigne S. (1996) - Quelle Préhistoire ancienne en Basse Auvergne et Velay?. In: Tuffreau, A., (Éd.), L'Acheuléen dans l'Ouest de l'Europe, Actes du Colloque de Saint Riquier, 1989, Lille, Publications du CERP 4, 115-127. Roebroeks W., van Kolfschoten T.H. (1995) - The earli- est occupation of Europe: a reappraisal of artefac- tual and chronological evidence. In Roebroeks W., van Kolfschoten T. (Eds.), The earliest occupation of Europe. Proceedings of the European Science Foundation workshop at Tautavel (France) (1993). University of Leiden Press, Leiden, 297-315. Ropiquet A., Hassanin, A. (2005) - Molecular phylogeny of caprines (Bovidae, Antilopinae): the question of their oriin and diversification during the Miocene. Journal of Zoological Systematics and Evolution- ary Research, 43(1), 49-60. Sadori L., Bertini A., Combourieu-Nebout N., Kouli K., Lippi M.M., Roberts N., Mercuri A.M. (2013) - Paly- nology and Mediterranean vegetation history. Flora Mediterranea, 23, 141-156. Sala B., Masini F. (2007) - Late Pliocene and Pleisto- cene small mammal chronology in the Italian pen- insula. Quaternary International, 160(1), 4-16. Sardella R. (1998) - The Plio-Pleistocene Old World dirk -toothed cat Megantereon ex gr. cultridens (Mammalia, Felidae, Machairodontinae), with com- ments on taxonomy, origin and evolution. Neues Jahrbuch fur Geologie und Palaontologie- Abhandlungen, 207(1), 1-36. Sardella R., Petrucci M. (2012) - The earliest Middle Pleistocene Crocuta crocuta (Erxleben, 1777) at Casal Selce (Rome, Italy). Quaternary Interna- 166 M.R. Palombo 358. Toro-Moyano I., Barsky D., Cauche D., Celiberti V., Grégoired S., Lebegued F., Moncel M.H., de Lum- ley H. (2011) - The archaic stone tool industry from Barranco León and Fuente Nueva 3, (Orce, Spain): Evidence of the earliest hominin presence in southern Europe Quaternary International, 243 (1), 80-91. Torres T., Cobo R., Garcìa Alonso P., Grun R., Hoyos M., Julia R., Llamas J., Soler V. (1995) - Evolucion del sistema fluvial Jarama-lozoya-Guadalix du- rante el Plioceno final y el Cuaternario. Geo- gaceta, 17, 46-48 Tsoukala, E. (2004) - The early pleistocene carnivores (Mammalia) from Ceyssaguet (Haute-Loire). Paléo, 16, 193-242. Turner A., Antón M. (1996) - The giant hyaena, Pachy- crocuta brevirostris (Mammalia, Carnivora, Hyae- nidae). Geobios, 29(4), 455-468. Tzedakis P. C., Hooghiemstra H., Pälike H. (2006) - The last 1.35 million years at Tenaghi Philippon: re- vised chronostratigraphy and long-term vegetation trends. Quaternary Science Reviews, 25(23), 3416 -3430. Valli A.M.F., Caron J.B., Debard E., Guérin C. (2006) - Le gisement paléontologique villafranchien termi- nal de Peyrolles (Issoire, Puy-de-Dôme, France): résultats de nouvelles prospections. Geodiver- sitas, 28, 297-317. Vallverdú J., Saladié P., Rosas A., Huguet R., Cáceres I., Mosquera M., Gracía-Tabernero A., Estalrrich A., Lozano-Fernández I., Pineda-Alcalá A., Car- rancho A., Villalaín J.J., Bourlès D., Braucher R., Lebatard A., Vilalta J., Esteban-Nadal M., Ben- nàsar M.L., Bastir M., López-Polín L., Ollé, A., Vergés, J.M., Ros-Montoya, S., Martínez-Navarro, B., García, A., Martinell, J., Expósito I., Burjachs F., Agustí J., Carbonell E. (2014) - Age and date for early arrival of the Acheulian in Europe (Barranc de la Boella, la Canonja, Spain). PloS one, 9(7), e103634. van Dam J.A. (2003) - European Neogene mammal chronology: past, present and future. Deinsea, 10, 85-95. Van der Made J. (1998) - Ungulates from Gran Dolina (Atapuerca, Burgos, Spain)[La faune des ongulés de la séquence de Gran Dolina (Atapuerca, Bur- gos, Espagne)]. Quaternaire, 9(4), 267-281. Van der Made J., Carlos Calero J.A., Mancheno, M.Á. (2008) - New material of the goat Capra? alba from the Lower Pleistocene of Quibas (Spain); notes on sexual dimorphism, stratigraphic distribu- tion and systematics. Bollettino della Società Pale- ontologica Italiana, 47, 13-23. Van der Made J. (1999) - Ungulates from Atapuerca- TD6. Journal of Human Evolution, 37(3/4), 389- 413 Van der Made J. (2001) - Les ongulés d'Atapuerca. Stratigraphie et biogéographie. L'Anthropologie, 105 (1), 95-113. Van der Made J. (2013) - First description of the large mammals from the locality of Penal, and updated faunal lists for the Atapuerca ungulates-Equus tional, 267, 103-110. Schloss C.A., Nuñez T.A., Lawler J.J. (2012) - Dispersal will limit ability of mammals to track climate change in the Western Hemisphere. Proceedings of the National Academy of Sciences, 109(22), 8606-8611. Sher A. (1997) - An Early Quaternary bison population from Untermassfeld: Bison menneri sp. nov. In Kahlke R.D. (Ed.), Das Pleistozuan von Unter- mussfeld bei Meiningen (Thüringen), Teil 1. Rudolf Habelt Verlag, Bonn, 101-180. Siori M.S., Sala B. (2007) - The mammal fauna from the late early Biharian site of Castagnone (Northern Monferrato, Piedmont, NW Italy). Geobios, 40(2), 207-217. Siori M.S., Boero A., Carnevale G., Colombero S., Delfino M., Sardella R., Pavia M. (2014) - New data on Early Pleistocene vertebrates from Monte Argentario (Central Italy). Paleoecological and biochronological implications. Geobios, 47(6), 403- 418. Sotnikova M.V. (2001) - Remains of Canidae from the lower Pleistocene site of Untermassfeld. In: Kahlke, R.D. (Ed.), Das Pleistozan von Unter- massfeld bei Meiningen (Thüringen), Teil 2. Rudolf Habelt Verlag, Bonn, 607-632. Spassov N. (2003) - The Plio-Pleistocene vertebrate fauna in South-Eastern Europe and the megafau- nal migratory waves from the east to Europe. Re- vue de Paléobiologie, 22, 197-229. Spassov N. (2011) - Acinonyx pardinensis (Croizet et Jobert) remains from the Middle Villafranchian locality of Varshets (Bulgaria) and the Plio- Pleistocene history of the cheetahs in Eurasia. Estud. Geol., 67, 245-253. Stehlin H.G. (1910) - Remarques sur les faunules de Mammifères des couches Éocènes et Oligocènes du Bassin de Paris. Bulletin de la Societe Ge- ologique de France, 9 (4), 488-520. Steninger F.F., Piller W.E. (Eds) (1999) - Empfehlungen (Richtlinien) zur Handhabung der strati- graphischen Nomenklatur. Courier Forschungsin- stitut Senckenberg, 209, 1-19. Stewart, J.R. (2009) - The evolutionary consequence of the individualistic response to climate change. Journal of evolutionary biology, 22(12), 2363- 2375. Suc J., Popescu, S. (2005) - Pollen records and climatic cycles in the North Mediterranean region since 2.7 Ma. Geological Society, London, Special Publica- tions, 247(1), 147-154. Tedford R.H. (1970) - Principles and practices of mam- malian geochronology in North America. In Pro- ceedings of the North American Paleontological Convention, 1 (Part F), 666-703. Thenius E. (1954) - Die Caniden (Mammalia) aus dem alquartär von Hundsheim (Niederösterreich) nebst bemerkungen zur stammesgeschichte des gattun Cuon. Neues Jahrbuch fur Geologie und Paläon- tologie, 99 (2), 230-286. Thouveny N., Bonifay E. (1984) - New chronological data on European Plio-Pleistocene faunas and hominid occupation sites. Nature, 308(5957), 355- 167 Late Early Pleistocene mammalian assemblages from SW Europe ............ Ms. received: February 8, 2016 Final text received: May 19, 2016 altidens, Bison and human dispersal into Western Europe. Quaternary International, 295, 36-47. Van der Made J. (2015) - The latest Early Pleistocene giant deer Megaloceros novocarthaginiensis n. sp. and the fallow deer Dama cf. vallonnetensis from Cueva Victoria (Murcia, Spain). In Gibert L., Fer- randez-Canadell C., (Eds), Geología y Paleon- tología de Cueva Victoria. Mastia, 11-13, 269-323. Van der Made J., Dimitrijević V. (2015) - Eucladoceros montenegrensis n. sp. and other Cervidae from the Lower Pleistocene of Trlica (Montenegro) - Quaternary International, 389, 90-118. Van der Made J., Tong H.W. (2008) - Phylogeny of the giant deer with palmate brow tines Megaloceros from west and Sinomegaceros from east Eurasia. Quaternary International, 179(1), 135-162. van der Meulen A.J. (1973) - Middle Pleistocene smaller mammals from the Monte 16 Peglia (Orvieto, Italy) with special reference to the phylogeny of Micro- tus 17 (Arvicolidae, Rodentia. Quaternaria, 17, 1- 144. Vislobokova I.A. (2011) - Historical development and geographical distribution of giant deer (Cervidae, Megacerini). Paleontological Journal, 45(6), 674- 688. Vislobokova I.A. (2013) - Morphology, taxonomy, and phylogeny of megacerines (Megacerini, Cervidae, Artiodactyla). Paleontological Journal, 47(8), 833- 950. Walker M.J., López-Martínez M., Carrión-García J.S., Rodríguez-Estrella T., San-Nicolás del-Toro M., Schwenninger J.L., López-Jiménezm A., Ortega- Rodrigáñezm J., Haber-Uriartem M., Polo- Camacho J.L., García-Torre J., Campillo-Boj M., Avilés-Fernández A., Zack, W. (2013) - Cueva Negra del Estrecho del Río Quípar (Murcia, Spain): A late Early Pleistocene hominin site with an “Acheulo-Levalloiso-Mousteroid” Palaeolithic assemblage. Quaternary International, 294, 135- 159. Walsh S.L. (1998) - Fossil datum and paleobiological event terms, paleontostratigraphy, chronostratigra- phy, and the definition of land mammal “age” boundaries. Journal of Vertebrate Paleontology, 18(1), 150-179. Williams H.S. (1901) - The discrimination of time values in geology. Journal of Geology 9(7), 570-585. Woodburne M.O. (1977) - Definition and characterization in mammalian chronostratigraphy. Journal Paleontology, 51, 220-234. Woodburne M.O., Goin F.J., Bond M., Carlini A.A., Gelfo J.N., López G.M., Iglesias A., Zimicz A.N. (2014) - Paleogene land mammal faunas of South Amer- ica; a response to global climatic changes and indigenous floral diversity. Journal of Mammalian Evolution, 21(1), 1-73. Zachos J., Pagani M., Sloan L., Thomas E., Billups K. (2001) - Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292(5517), 686 -693. 168 M.R. 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