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The rich Quaternary fossil record from the Cantabrian region (NW Iberia) is virtually restricted to the Late Pleistocene, with Middle Pleistocene findings very rare. This work presents the study of two Middle Pleistocene palaeontological sites in the Cantabrian region: Llantrales quarry and Mestas de Con mine, both discovered and first described in the 1950s but never studied in detail thereafter. The material was here re-analysed and re-evaluated. Taxonomic attributions have been changed in most cases and, consequently, chronological assignments of the fossils have also been changed. This first detailed study of the assemblage from Llantrales quarry revealed the occurrence of the large-sized deer Praemegaceros solilhacus, which has never been reported previously in North Iberia. Cervus cf. elaphus and Stephanorhinus cf. hundsheimensis were also identified. The age of these fossils was originally regarded as Pontian (late Miocene), but new palaeontological results indicate a much younger age, most likely from around 0.8 to 0.5 Ma. A taxonomic re-evaluation of the fossils from Mestas de Con provided different results with respect to the 1950s works. The faunal association is composed of a large-sized deer (probably corresponding to Praemegaceros), Cervus elaphus, Capreolus cf. capreolus, Bison sp. (small sized), Equus sp. (large sized), Stephanorhinus hemitoechus, Ursus sp. and Homotherim latidens. This faunal association was evaluated within a western European context and proves to be highly analogous to other contemporary assemblages there. The chronology was initially regarded as late Villafranchian (in a broader sense). New results, documenting the co-occurrence of Stephanorhinus hemitoechus and Homotherium latidens, indicate a more precise and younger age, most likely between 0.6 and 0.4 Ma. These two fossil assemblages provide new and relevant information on the Middle Pleistocene faunas from a geographical region where this information, compared with other western European areas, is rare and poorly known.

Villafranchian faunas were widely spread throughout the European continent during the Plio-Pleistocene. Ungulates, and particularly ruminants, are very well represented in most of the localities of this period. The fossil site of Tegelen (Province of Limburg, the Netherlands) is a classical Villafranchian locality where cervids are especially abundant. Some of the sites from this period have been thoroughly studied in various aspects, such as faunal assemblages, biochronology, magnetostratigraphy, sedimentology or palaeoenvironmental conditions, among others. Tegelen has been as well studied, but its particular exploitation has hampered its study for years. In order to reconstruct its prevailing palaeohabitat, we have applied statistical analyses based on functional morphology for the first time for this fossil site. In this work, we have analysed the ecomorphology of the postcranial skeleton of Eucladoceros ctenoides and Metacervoceros rhenanus, the two extinct species of deer present in Tegelen. Results show that the best ecological indicators are the metapods and the tibia, while the astragalus seems to be the least informative element. The analyses reveal likewise how both taxa show adaptations to both open and forest environments, but with a slight trend to the former. These results disagree with the general assumption that E. ctenoides and M. rhenanus are indicative of forested settings, as well as with the classical interpretations of the palaeoenvironment of Tegelen made to date, which are exclusively based on the fauna. However, they go along with the appearance of some taxa typical of open habitats, such as the micromammals Hystrix and Dicrostonyx, which could help to explain their enigmatic presence. Copyright © 2014 John Wiley & Sons, Ltd.

Knowledge on the Villafranchian faunas of Greece was limited for a long time, but over the past 3 decades several new fossiliferous sites have been discovered and new material has been unearthed. The carnivores constitute a remarkable part of this material and their study has provided useful information for the taxonomy, chronology and palaeoecology. The Villafranchian carnivoran guild includes numerous taxa belonging to various families. Some of them are interesting as they are differentiated from the other Eurasian families. They cover the entire Villafranchian but the mass is known from the Middle to the Epi-Villafranchian (EVC). Despite the discontinuous data, they provide useful biochronological results. The coexistence of Pliocrocuta perrieri and Pachycrocuta brevirostris in the locality of Gerakarou 1 is strong evidence for the Middle Villafranchian (MVC)/Late Villafranchian (LVC) boundary. The composition of the Villafranchian carnivores suggests that the canids predominate, while the felids, hyenids and mustelids are well represented; the ursids are represented by a single taxon, the well-known Ursus etruscus, which seems to be common during the entire Villafranchian. The carnivoran guild structure and the multivariate analysis of the Greek Villafranchian carnivoran assemblages (MVC, LVC and EVC) in comparison with modern assembalges from open and closed environments indicate a relatively open landscape, which is in agreement with previous results for the Villafranchian of Greece. During the EVC there is a faunal renewal that is related to a more open and dry environment, like savannah bushland/grassland.

Aim Knowledge of mammal community composition across large areas is critical to the reconstruction of biogeographical trends in past ecosystems where mammalian fossils serve as palaeoecological proxies. We employed ‘ecometrics’ (taxon-free trait analysis) to compare fossil communities of the Italian Plio-Pleistocene across space and time. We tested the hypotheses that ecometric traits characterizing fossil mammal communities differ between time bins, and that they exhibit geographical correlations. Location Peninsular Italy, excluding islands (Sardinia and Sicily). Methods We explored selected ecometric traits (body weight, biomass, hypsodonty and brachial index) in 69 local fossil assemblages at the scale of nine palaeocommunities (PCOMs), and at the broader scale of ‘mammal age’, or biochronological unit, to which they belong: Villafranchian, Galerian or Aurelian. Principal components analysis, MANOVA and linear discriminant analysis were used to describe differences in the PCOM and mammal age ecospaces. We also tested for the effects of latitude and longitude on the ecometric variables and mapped their distributions during individual mammal ages. Results Spatial analysis suggested that neither latitude nor longitude relate meaningfully to any of the ecological parameters under consideration. However, PCOMs in each biochronological interval were successfully discriminated using ecometric traits, with the exception of the localities from Triversa and Pirro, which bore more resemblance to the younger Aurelian localities. The mammal ages were more clearly distinct. Main conclusions Communities differ from one another particularly in relation to both the average body weight of herbivores and the entire mammalian fauna and the biomass of prey species. This is probably related to a climatic cooling trend that occurred between the Pliocene and Pleistocene and resulted in the evolution of larger forms. Hypsodonty trends are identified and we support hypsodonty as a valuable ecometric trait, but only at larger temporal scales.

A review on the position of the Pliocene-Pleistocene boundary is presented. Uncertainties about the Calabrian stratotype and mammal faunas ill relation to absolute datings are discussed. In the North Sea basin and elsewhere a marked cold stage, the Praetiglian, began about 2.5 million years ago. It caused a partial extinction of the flora, which in Italy too is distinctly represented in fresh-water as well as marine deposits. In the latter deposits, the extinction level coincides with the base of the Calabrian. The same type of extinction occurs at the base of the Villafranchian faunal sequence dated between about 2.5 and 3.1 million year-s ago. The Calabrian base is therefore most probably at about 2.5 m.y., not at 1.8 m.y. as previously thought. The 1948 Recommendation on the position of the Plio-Pleistocene boundary is still considered valid.

We report here on early hominid facial diversity, as part of a more extensive morphometric survey of cranial variability in Pliocene and early Pleistocene Hominidae. Univariate and multivariate techniques are used to summarise variation in facial proportions in South and East African hominids, and later Quaternary groups are included as comparators in order to scale the variation displayed. The results indicate that “robust” australopithecines have longer, broader faces than the “gracile” form, but that all australopithecine species show comparable degrees of facial projection. “Robust” crania are characterised by anteriorly situated, deep malar processes that slope forwards and downwards. Smaller hominid specimens, formally or informally assigned to Homo (H. habilis, KNM-ER 1813, etc.), have individual facial dimensions that usually fall within the range of Australopithecus africanus, but which in combination reveal a significantly different morphological pattern; SK 847 shows similarly hominine facial proportions, which differ significantly from those of A. robustus specimens from Swartkrans. KNM-ER 1470 possesses a facial pattern that is basically hominine, but which in some respects mimics that of “robust” australopithecines. Early specimens referred to H. erectus possess facial proportions that contrast markedly with those of other Villafranchian hominids and which suggest differing masticatory forces, possibly reflecting a shift in dietary niche. Overall the results indicate two broad patterns of facial proportions in Hominidae: one is characteristic of Pliocene/basal Pleistocene forms with opposite polarities represented by A. boisei and H. habilis; the other pattern, which typifies hominids from the later Lower Pleistocene onwards, is first found in specimens widely regarded as early representatives of H. erectus, but which differ in which certain respects from the face of later members of that species.

Hominids, Pebble Tools and the African Villafranchian l F. CLARK HOWELL Washington University School of Medicine (St. Louis) T HE greatly accelerated pace of prehistoric research in the postwar years has made it necessary to reexamine certain problems in paleoanthropol­ ogy. This is especially true for Africa where primitive hominids and pebble tools have been discovered in deposits of recognized Villafranchian age. These discoveries raise the question of the relationship, if any, between these earliest stone implements and the Villafranchian hominids. A related problem is the place of the Pliocene-Pleistocene boundary and the age of the Villafranchian stage in Africa \\ ith reference to the Pleistocene sequence in other parts of the Old World. A review of the evidence for Pleistocene man south of the Sahara will be presented at another time, but it is worth while here to clarify certain points about the Villafranchian stage in Africa in view of some misconcep­ tions present in the literature. The first evidence of tool-making is found in deposits of the first pluvial (Kageran) stage in equatorial and southern Africa. Pebble tools of this age have been found on the eastern shores of Victoria basin at Kanam (Leakey 1936a, 1936b), and in the Kagera and Kafu river valleys west of the basin in Uganda (Wayland 1934; Lowe 1952); artifacts of comparable age have recently been found sealed in situ in ancient gravel deposits in the Vaal river basin (Lowe 1953). These implements, referred to the Kafuan stage of the Pre-Chelles­ Acheul industry, were manufactured from waterworn pebbles of quartz or quartzite with stone percussion flaking in one direction, and on only one side, to produce a single, adze-like cutting edge. Although artifacts of Kafuan type are found rolled and abraded in deposits of post-Kageran age at various locali· ties south of the Sahara, the industry may well be restricted to the first pluvial stage. The Oldowan, a derivative stage of the African Pre-Chelles-Acheul, is found in deposits of the early part of the second pluvial (Kamasian) stage. Some Oldowan implement types are found in the final phases of the Kafuan (Lowe 1952), however, so it is possible that this industry had its beginnings in the later part of the first pluvial stage. This problem may be clarified by future field work in northwestern and central Africa. Pebble tools of first end­ pluvial age (Kageran-Kamasian interpluvial) have been found recently in southern Morocco (Mortelmans, Choubert and Hollard 1952) and also in the vicinity of Casablanca (Biberson 1953). An industry termed the Kafilian, more advanced than the Kafuan, is known from the Congo and is of the same ap­ proximate age (Mortelmans 1947, 1949). These assemblages may well fill the gap between the Kafuan and the Oldowan.