Palaeogeography, Palaeoclimatology, Palaeoecology, 66 (1988): 77-100 77 Elsevier Science Publishers B.V., Amsterdam - Printed in The Netherlands

LATE PLlOCENE TO EARLY MID-PLEISTOCENE MAMMALS IN EURASIA: FAUNAL SUCCESSION AND DISPERSAL EVENTS

A. AZZAROLP, C. DE mULl!, G. FICCARELLI2 and D. TORRE 1

'Earth Sciences Department, University of Florence, Florence (Italy) 2 Earth Sciences Department, University of Camerino, Camerino (Italy)

(Received June 11, 1987; revised and accepted December 1, 1987)

Abstract

Azzaroli, A., De Giuli, C., Ficcarelli, G. and Torre, D., 1988. Late Pliocene to early mid-Pleistocene mammals in Eurasia: faunal succession and dispersal events. Palaeogeogr., Palaeoclimatol., Palaeoecol., 66: 77-100.

Four major dispersal events mark European Villafranchian and Galerian faunas (about 3.2-0.4 Ma). The beginning of the Villafranchian is evidenced by the arrival of Leptobos and of large cervids and felids; the fauna still retains typical forest elements: Mammut, Tapirus, etc. The Elephant-Equus event (about 2.5-2.6 Ma) brought in grassland elements (elephant, horse) while several forest dwellers disappeared. The massive arrival of a primitive wolf, a large hyaena and Leptobos etruscus approximately marks the Plio-Pleistocene boundary (Wolf event, about 1.7 Ma). This was followed by a new wave of prairie fauna: Praeovibos, "Leptobos" vallisarni, Allophaiomys, Canis arnensis (a coyote), Canis falconeri (possibly a lycaonid); Cervalces and Hippopotamus also arrived at this time. The Villafranchian-Galerian transition saw a total faunal turnover, with massive extinctions and new, previously unknown adaptations (end-Villafranchian event, 1.0-0.9 Ma). The late Pleistocene and living fauna of Eurasia took its origin at this time. Mammalian stratigraphy of Asia is more poorly known but faunas are easily correlated with European ones and the end-Villafranchian event is clearly recognised. Faunal events are correlated with climatic and physiographic changes (late Himalayan orogeny).

Introduction provided evidence that major faunal events tend to coincide with good approximation with In the palaeontological record the succes­ changes in vegetation, in climate, sea level sion of mammalian faunas is not characterised fluctuations, temperature changes in sea and throughout their history by steady, gradually ocean waters. progressing evolution but is interspersed with The time interval examined here spans the sudden, abrupt changes: extinctions, rapid late Pliocene, the early Pleistocene and part of evolutionary progress, appearance of new the middle Pleistocene, i.e. the Villafranchian adaptations, migrations. Such discontinuities and Galerian stages of European vertebrate were once interpreted as artifacts due to stratigraphy: in terms of absolute chronology, incompleteness ofthe record; but as knowledge from little before 3 Ma to approximately 0.4 progressed and gaps were being filled in it Ma. became apparent that they represent real revolutions of the faunal assemblages, "faunal Villafranchian events" of continental scope (Repenning, 1980). Calibration of faunal succession with pollen The term Villafranchian was introduced by sequences, isotopic and palaeomagnetic scales Pareto (1865) for a fauna collected from lacus-

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trine deposits near Villafranca d' Asti, Pied­ mation; its end has not been established with other hand do not show significant changes sites in Italy, in the French Central Massif and mont, northwestern Italy; the term was ex­ the same accuracy, but will not be dealt with in from the Ruscinian: Anancus arvernensis, in Spain (Fig. 1). tended by its author to include mammalian this paper. Unlike the Villafranchian, the Mammut borsoni. In Spain the first signs of The accompanying flora is rich and is faunas of the upper and lower valley of the Galerian fauna includes several taxa still the transition can be detected in the Layna indicative of a warm climate (Lona and Ber­ Arno in Tuscany, central Italy. Since then the living today, or taxa differing from present local fauna where Chasmaporthetes, Lepto­ toldi, 1973; Suc and Zagwijn, 1983). term Villafranchian has been used rather ones only at specific or subspecific level. In fact bos and "Lepus" are recorded for the first The lacustrine beds of the .type localities loosely, mainly by Italian, French and Swiss the present faunal assemblage of Eurasia took time. around Villafranca d' Asti represent the de­ authors to designate faunal assemblages, its origin in this time interval. The Galerian is The immigration of so many taxa points to posit of a shallow coastal lake. To the east they mostly from Italy and from southern France, also characterised by the appearance of new, some environmental change, possibly the are replaced by littoral facies with terrestrial which share some features: occurrence of previously unknown adaptations. establishment of a less dense forest cover vertebrates and marine fossils, in which the proboscideans associated with large bovids, (appearance of Acinonyx, Pliotragus etc.). Fau­ Triversa unit may be correlated, somewhat deer and equids, and above all the fact that The beginning of the Villafranchian - nal assemblages of this type occur in various indirectly, with the Globoratalia puncticulata- they are entirely composed of taxa now extinct. The "Leptobos" event The Villafranchian was initially believed to correspond to the late Pliocene: Gignoux (1916) The transition from Ruscinian (early Plio­ AUVEcRGNE correlated it with his Calabrian stage. At the cene) to early Villafranchian faunas (Triversa ET VE LAY 18th International Geological Congress faunal unit: Azzaroli, 1977) is marked by (London, 1948) it was agreed to place the several changes. The Ruscinian antelopes Par­ Calabrian at the base of the marine Pleisto­ abos and Alephis are replaced by the more cene. Several years passed before it was advanced Leptobos. Ruscinian deer are of realised that the so-called Villafranchian fau­ small size and only Croizetoceros ramosus has nas are far from uniform and actually span a long, branched antlers. The early Villafran­ large part of the Pliocene and the early chian deer fauna is richer and is represented Pleistocene (Azzaroli, 1963, 1970, 1977; Heintz, by several advanced taxa of large size: 1967). As a matter of fact the term Villafran­ Cervus pardinensis and a similar deer, more chian survives now for traditional reasons derived in the development of its antlers, in more than for its intrinsic value, and is western Tuscany; Cervus perrieri, Arvernoceros practically meaningless if not used with some ardei. Croizetoceros ramosus survived from the qualification (early, middle, later Villafran­ Ruscinian, with more progressive subspecies chian). Azzaroli (1977) divided the Villa­ (Heintz, 1970, 1974). Another immigrant among franchian into six more or less well defined ruminants is the caprine Pliotragus; among faunal units. The beginning of the Villafran­ perissodactyls, the slender Dicerorhinus miguel­ chian itself, of some of its units and its end are crusafonti, and then jeanvireti, replaced the characterised by pronounced dispersal events; more massive D. megarhinus and among canids other units were distinguished on the basis of Nyctereutes megamastoides replaced N. donne­ minor events or more gradual changes. zani. Tapirus, a survivor from the Ruscinian, is a common element of the fauna, whereas Galerian Hipparion was now rare in France and possibly extinct in Italy; it survived in larger numbers The name Galerian was introduced by Am­ in Spain, at Villaroya. The large carnivores brosetti et al. (1972) to designate a distinct made their first appearance: Chasmaporthetes, faunal assemblage in Europe; the same assem­ Acinonyx, Megantereon, Homotherium. Ursus FAUNAS blage, with minor changes, is found to charac­ minimus may be a progressive descendant of terise the same time interval in Asia. U. ruscinensis: it has no relationships with • Montopoli -St. Vallier units The Galerian spans a much shorter time later Villafranchian bears and its skeleton than the Villafranchian, of the order of 0.5 Ma, suggests a partly arboreal adaptation (Berzi, Fig.I. Pliocene local faunas, discussed in text. Triversa unit (+): 1 = Layna; 2= Villaroya; 3= Les Etouaires; 4 = Vialette; 5=Villafranca d'Asti. Montopoli-St. Vallier units (e): 6= Rinc6n; 7=La Puebla de Valverde; 8= Rocaneyra; 9=Chilhac; and is faunistically much more uniform. Its 1966). "Lepus" seems to have made now its first 10=Seneze; l1=Le Coupet; 12=La Roche Lambert; 13= St. Vidal; 14= St. Vallier; 15= Montopoli; 16=Red Crag; beginning has been dated with good approxi- appearance in Europe. Proboscideans on the 17= Tegelen; 18=Khapry; 19=Livenzovka; 20= Kuruksay.

t 80 81

G. crassaformis transition of marine Pliocene selected as the type of the Montopoli faunal Savage and Curtis, 1970). The fauna, which equated with the Triversa faunal unit, the stratigraphy (beginning of the late Pliocene) unit (Azzaroli, 1977). unfortunately is rather poor, provides the Khaprovian with the Montopoli and St. Vallier (De Giuli et aI., 1984). Radiometric ages are The fauna was collected from a single oldest record of Equus and the last record of units. available for two French localities: Vialette pocket. It is rich and is characterised by Hipparion in France (Eisenmann and Brunet, In China a rather sharp break in mammalian (between 3.3 and 2.6 Ma: Bandet et aI., 1978) the arrival of highly significant elements: a 1973). faunas and in vegetation occurred approxi­ and les Etouaires. For this site Couthures and primitive elephant, Archidiskodon gromovi, a The Dutch Praetiglian has yielded rather mately at the Gauss-Matuyama transition (Liu Pastre (1983) gave an age between 3.3 and 2.6 mono dactyl equid of large size, Equus cf. poor vertebrate remains, which however seem Tung-sheng and Ding Meng-Lin, 1982; Song Ma; on the other hand Ly et al. (1983) proposed livenzovensis, closely related to Equus stenonis to fit in the Montopoli unit; Anancus arvernen­ Zhi-Chen et aI., 1982; Wang Fu-bao and Li an age younger than 2.6-2.5 Ma. The latter and to the North American E. simplicidens. sis, Archidiskodon cf. gromovi (referred to as Bing-yuan, 1982). date was not obtained directly from the fossil The cervid Eucladoceros appeared at this time, Archidiskodon planifrons by Van der Vlerk In the Indian subcontinent the arrival of bearing beds but by indirect correlation. The but is poorly represented at Montopoli. At the and Florschutz, 1950); Eucladoceros falconeri, Equus, Rhinoceros, Elephas hysudricus and error limits are rather broad, especially for the same time the most typical forest elements the most primitive species of this genus; Equus Stegodon insignis at the Gauss-Matuyama les Etouaires date (for this site, Savage and disappeared from the fauna: Mammut, Tapirus, sp. ind. is also present, alongside Odobaenus transition marks the boundary between the Curtis, 1970, reported a date of 3.4-3.5 Ma in an Sus minor, Ursus minimus. and Choneziphius. The pollen flora of the Tatrot and Pinjor faunal complexes (Lindsay et ash underlying the fossil bed). In spite of the The faunal change from the Triversa to the Praetiglian characterizes a cold interval aI., 1980; Azzaroli, 1985b). great discrepancy between datings, ages pro­ Montopoli assemblage is sharp and from the spanning approximately from 2.5 to 2.2 Ma posed are not incompatible with the attribu­ ecological viewpoint much more significant (Zagwijn, 1974; Zagwijn and Suc, 1984). The middle Villafranchian tion of the Etouaires fauna to an assemblage than the change from the late Ruscinian The English Red Crag has a similar fauna: older than the elephant-Equus event. Lindsay (Perpignan) to the Triversa assemblages. The Odobaenus, Anancus, Eucladoceros falconeri, a The Montopoli faunal assemblage is fol­ et al. (1980) calibrated with the palaeomag­ fauna is indicative of a more open, parkland large Equus referred to as E. robustus by lowed without sharp break by faunas contain­ netic scale a section in the type area of and savannah landscape. The Montopoli fauna Hopwood (1937) but possibly belonging to ing similar taxa, but generally showing more Villafranca d' Asti and proposed an age of comes from littoral deposits capping a marine E. livenzovensis (several teeth, partly unde­ derived features. Archidiskodon gromovi is 3.01-3.05 Ma (a short episode of normal polar­ sequence of the Globorotalia crassaformis zone scribed, in the British Museum (Nat. Hist.); succeeded by an elephant resembling A. meri­ ity between the Mammoth and Kaena reversed (De Giuli et aI., 1984). Lindsay et aI., (1980) an elephant, referred to as Elephas meri­ dionalis in dental features (Boeuf, 1983) but episodes). tentatively calibrated it with the transition dionalis by Leith Adams (1881, pl.26, fig.2), more primitive in its skull. Cervus pardinensis The faunas of the Triversa unit are enriched from the Gauss (normal) to the Matuyama seems to fit better in Archidiskodon gromovi was replaced by C. rhenanus Dubois, 1905 by several taxa as compared with Ruscinian (reversed) palaeomagnetic epochs, about 2.46 (a fragmental molar, with a very thick [ = C. philisi Schaub, 1941, according to one of faunas and appear more advanced in their Ma. An elephant skeleton was however col­ enamel). The molluscan fauna of the late Red us (A.A.)], Croizetoceros ramosus ramosus by composition (occurrence of large carnivores lected in marine and brackish beds at Laiatico, Crag is rich in Arctica islandica, which C.r. medius and possibly C.r. minor in France and antlered deer). From the point of view of to the south west and downsection of Monto­ clearly indicates a cold climate. It should and C. ramosus pueblensis in Spain (Heintz, environment they do not show any great poli, so that the faunal event may be somewhat be remembered however that the Red Crag 1974). Dicerorhinus jeanvireti was superseded change from the previous situation and still older. mammalian fauna is mixed. Besides Pliocene by the smaller D. etruscus. Cervus perrieri and keep the character of warm climate, predomi­ An assemblage referable to the Montopoli elements it includes post-Villafranchian ele­ Arvernoceros ardei are no longer recorded nantly forest assemblages. Isotopic analyses in unit is represented by the Rincon 1 local fauna ments (Azzaroli, 1970; Megaceros verticornis, while the more derived Eucladoceros tegulen­ marine and lacustrine series provide evidence in the Jucar valley, eastern Spain (Alberdi et Equus cf. caballus), which obviously come sis Dubois, 1905 [= E. senezensis Deperet and of a moderate climatic deterioration between aI., 1983; Leone, 1985). The fauna is character­ from a younger deposit, possibly some channel Mayet, 1910, according to one of us (A.A.)] may 3.2 and 3.1 Ma, followed by a return to higher ised by abundant Gazella borbonica and an filling. be a descendant of the more primitive E. fal­ temperatures, with mean values only slightly Equus of large size, attributed to E. stenonis In southern Russia the elephant-Equus coneri. Immigrants are few, among these the below those of the Ruscinian (Shackleton et but possibly identifiable with E. livenzovensis. event i5 also recognized and corresponds to the uncommon, medium sized felid Viretailurus aI., 1984; Leone, 1985). This fauna occurs in the late Gauss epoch and transition from the Moldavian to the Khapro­ schaubi from St. Vallier (Viret, 1954), and the may be dated to approximately 2.6 Ma. The vian "complex". The latter is typified by the bovids Gallogoral meneghinii and Gazellospira The elephant-Equus event appearance of this fauna corresponds to a faunas of Khapry near the Azov Sea and of torticornis, which both survive into the late climatic deterioration evidenced by isotopic Livenzovka near Rostov on Don, with Archi­ Villafranchian. A sharp break marks the end of the Triversa oxygen and carbon analysis in the lacustrine diskodon gromovi, Equus livenzovensis etc. The limits and duration of the middle fauna, and the onset of a new assemblage, beds (Leone, 1985). (Bajgusheva, 1971, 1978). The boundary be­ Villafranchian have not been defined in detail typified by the later early Villafranchian Another fauna which may be referred to the tween Moldavian and Khaprovian is placed at and will be discussed below. faunas. The Montopoli local fauna in the Montopoli unit is Roca Neyra in the French approximately 2.5 Ma by Russian authors The richest site of the middle Villafranchian Lower Valdarno basin, Tuscany, has been Central Massif, dated 2.5 Ma (K-Ar date, (Schanzer, 1982); the Moldavian may be is St. Vallier in the Rhone valley, which is also

z 82 83 the type local fauna of the St. Vallier unit. The lack of Pachycrocuta brevirostris, Panthera the one observed in Seneze is remarkably good. of their rather broad stratigraphic distribu­ Other local faunas referable to this unit are toscana and Canis etruscus, all common ele­ The mammalian fauna seems to come from the tion, or of doubtful identification. Only the Chilhac, St. Vidal, La Roche Lambert, Le ments in late Villafranchian faunas, is sur­ TC1-TC6 interval (Zagwijn, 1974); however, panther calls for attention. It was described as Coupet (younger fauna), in the French Central prising. Sus strozzii is alien to the St. Vallier most fossils were collected by quarry workers Felis (Panthera) schreuderi by Von Koenigs­ Massif; La Puebla de Valverde in Spain. and other mid-Vallafranchian assemblages but in old times and their stratigraphic location wald (1960); Ficcarelli and Torre (1968) con­ Boeuf (1983) reported several, partly discor­ is recorded at Tegelen: in Seneze it is repre­ was not exactly recorded. Freudenthal et al. sider it a synonym of Panthera toscana Schaub, dant radiometric ages for a basalt flow over­ sented by a single specimen, with somewhat (1976) retrieved a rich assemblage of molluscs a species typical of the late Villafranchian (the lying the Chilhac fauna, which is reversely primitive marks (retention of the third upper and small mammals from level TC5 (temperate) name gombaszogensis Kretzoi has time priority magnetised, and concluded for a probable age incisors: Azzaroli, 1954). The fauna fits well in in the Egypte quarry. This is clearly a homo­ over toscana but was based on mixed material of 1.9 Ma for the basalt. a mid-Villafranchian (late Pliocene) assem­ geneous assemblage; on the other hand the list and is represented by a rather poor type: the The fauna of Seneze, in the French Central blage. of large vertebrates raises some problem. The meaning of the term gombaszogensis is am­ Massif, poses a definite problem. The site is a On the other hand, faunallists include some occurrence of Tapirus is surprising and, to us, biguous: Ambrosetti et al., 1979). Von Koenigs­ maar surrounded by a partly eroded volcanic elements that do not fit in such an old context suspect; this genus was not recorded in former wald's sample consists of dentitions and frag­ apparatus. Bout (1960, 1972) made reference to and clearly point to a late Villafranchian age: papers (Schreuder, 1945; Van der Vlerk and ments of skulls, representing at least three lacustrine beds of the maar and to superposed Cervalces gallicus, Canis arnensis, Equus bres­ Florschiitz, 1950). individuals. The specimens were collected slope deposits as sources of the fossils and san us and a small equid which may possibly be The elephant was reported as "Archidisko­ between 1943 and 1948; unfortunately the exact Elhay (1969) described the pollen association Equus stehlini. Megalovis may also be a don meridionalis, archaic form" by Van der stratigraphic position was not recorded. It may from the cores of a 175 m deep well drilled in member of this younger fauna. Vlerk and Florschiitz; it is based on fragmental be presumed that they were derived from the the maar. The floral association is character­ Associating these remarks with Bout's de­ molars to which the thick enamel gives a uppermost part of the section, where the pollen ised by a long interval with temperate flora at scription of two fossil bearing horizons the rather archaic appearance; it may possibly diagram marks a trend to a climatic deteriora­ the base, followed by a shorter cool interval idea comes quite naturally that Seneze con­ represent A. gromovi or some transitional tion forestalling the Eburonian cold phase. and then by a succession of oscillations from tains in fact two faunas of very different age: form. The remains of the large deer, called The time span of the Tiglian is roughly temperate to cool. It recalls the floral spectrum one, which is also the richest in species and Eucladoceros ctenoides by Freudenthal et al. between 2.2 and 1.7 Ma (Zagwijn, 1974), its of St. Vidal, a typical middle Villafranchian individuals, representing the middle Villafran­ (1976), were referred to Cervus dicranios and to mammalian fauna belongs to the middle Villa­ site, and of the Dutch Tiglian. The beds of the chian and is about 2 Ma old, we assume the C. teguliensis by Bernsen (1930), and to Eucla­ franchian and may possibly reach the begin­ maar are reversely magnetized, with a short palaeomagnetic interpretation to be correct (it doceros teguliensis by Kunst (1937). Compari­ ning of the late Villafranchian in the highest normal episode 20 m below the surface and is supported by the pollen record, see below); son with deer from other sites is not easy. Only part of the section. Correlations of the Tiglian about 10 m below the main fossil bearing the second representing the latest Villafran­ two nearly complete antlers are known, and with marine sequences have not been estab­ horizon. Bout interpreted this as the Gilsa chian, about 1 Ma younger. antlers of Eucladoceros are highly variable. lished but Nilsson (1983) proposes a correla­ event and inferred an age of 1.6 Ma for the The fauna of Tegelen in Limburg, The The specimen figured by Bernsen, pl.2, fig.1, fits tion with the Icenian. Seneze fauna. According to Ehrlich (quoted by Netherlands, is more difficult to evaluate well with E. senezenis; the other specimen is A mid-Villafranchian fauna was also re­ Bout, 1972) the filling of the lake may have because of its fragmentary state. Freudenthal juvenile and less characteristic. Freudenthal corded from San Giacomo near Anagni, Cen­ taken 0.2-0.3 Ma. et al. (1976) gave the following list: Mammu­ et al. omitted to mention Cervus rhenanus. This tral Italy (Biddittu et al., 1979). The fauna has Thouveny and Bonifay (1984) interpret in a thus meridionalis, Anancus arvernensis, Tapi­ is a species of small size, with two bifurcations not been fully studied but shares several different way the palaeomagnetic record and rus arvernensis, Dicerorhinus etruscus, Equus in the adult antlers, and according to Kunst elements with the faunas of St. Vallier, Seneze correlate the short normal interval with the bressanus, Leptobos sp., Eucladoceros ctenoides, conforms very well with the small deer from (older fauna) and Tegelen. Among the most older Reunion event (2.12-2.14 Ma). Sus strozzii, Ursus etruscus, Enhydrictis ardea, Seneze, Cervus philisi, which in 1937 had not diagnostic species for its age are Cervus The mammalian association of Seneze shows Pannonictis pliocaenica, Pachycrocuta perrieri, yet received a scientific name. rhenanus, Croizetoceros ramosus, Gazella bor­ some anomalies which do not allow a satisfac­ Panthera gombaszogensis, Castor fiber, Tro­ We consider the two species synonyms, and bonica, Equus stenonis, Archidiskodon meri­ tory correlation with other faunas and clearly gontherium cuvieri, Mimomys pliocaenicus, M. Cervus rhenanus Dubois 1905 has priority. dionalis. point to a mixed assemblage. The deer: Croize­ newtoni, Macaca sylvana florentina. Some com­ More important, we have here a species typical toceros ramosus, Cervus rhenanus (= philisi), ments may be made on the stratigraphy and of the middle Villafranchian. Eucladoceros The "Wolf" event and the beginning of Eucladoceros tegulensis, point to a correlation fauna of the site. The pollen diagram of the tegulensis (Dubois) 1905 also has priority over the late Villafranchian with middle Villafranchian faunas; Nyctereutes Tiglian shows in its lower part a pronounced E. senezensis (Deperet) 1910. and Dolichopithecus also point to a Pliocene temperate episode (TA) followed by a shorter The record of Trogontherium cuvieri is pro­ A pronounced faunal break marks the transi­ age. Leptobos etruscus was reported with a cool period (TB) and then a series of more rapid bably incorrect: Van der Vlerk and Florschiitz tion to the following unit; this was called the question mark by Schaub (1943) and has not oscillations from temperate to cool (TC1 to record T. boisvilletti. The rest of the fauna is "wolf" event (Azzaroli, 1983). The most charac­ been identified satisfactorily at species level. TC6). The agreement of this succession with composed of species less characteristic because teristic faunal changes are: the disappearance

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of Nyctereutes, Gazella, Leptobos stenometopon, represented in the Upper Valdarno basin, and the massive expansion in Europe of Canis where it overlies faunas of the Olivola unit etruscus (the "wolf"), Pachycrocuta brevi­ (Azzaroli and Lazzeri, 1977; Borselli et aI., 1980; UJ UJ /}.

meridionalis (Azzaroli, 1977a). relative of the East African H. gorgops (Elan­ Ul ~ Q) +' the late Villafranchian. It survived during the Giuli and Masini (1987) point out the occur­

7 f" ------

86 87

Giuli et al. (1987) is closer to Eobison than to Berzi, 1970); at Pirro Nord (Apricena) in the Dicerorhinus aff. hemitoechus D. etruseus and may well be its direct de­ Leptobos. Gargano peninsula (De Giuli and Torre, 1984; Dicerorhinus kirchbergensis scendant. Panthera leo, P. pardus, Aeynonyx Eucladoeeros dieranios also shows a ten­ De Giuli et al., 1987); at Scoppito (Aquila) in Crocuta crocuta jubatus and Croeuta eroeuta are most likely Panthera lea dency to increase in size and antler complexity, Abruzzo; at the Creux de Peyrolles in the Panthera pardus immigrants from Africa. The origin of other but is poorly known because remains are French Central Massif (Bout and Azzaroli, Acinonyx jubatus (= Acinonyx intermedius) taxa is not known. scanty and fragmentary. In the Pirro Nord and 1953). The late Villafranchian taxa which did Selvella faunas a large Eucladoeeros occurs, Cuon prise us was recorded by Thenius from not develop new adaptations: Eucladoeeros, considered to be a species different from The Galerian fauna and the "end­ localities of possibly Galerian age (fide Kurten, Dama nestii, Leptobos etruscus and other bo­ E. dieranios (De Giuli, 1987; De Giuli et al., Villafranchian" dispersal event 1968). Hippopotamus antiquus is recorded from vids, Sus strozzii, Arehidiskodon meridionalis 1987). Outside Italy some faunas of very late the late Villafranchian and from the Galerian. and several carnivores became extinct at the Villafranchian age (Creux de Peyrolles, cen­ The Galerian is characterized by a highly Dieerorhinus etruscus was reported by several end of the Villafranchian or in the early tral France; East Runton, England: Bout and distinctive faunal assemblage which ranges, authors from Galerian deposits: Mosbach, Galerian. Azzaroli, 1953; Azzaroli, 1953) are character­ with some local variation, throughout Eurasia, Voigtstedt and Siissenborn (Schroeder, 1903; Obviously the transition from late Villa­ ised by Eucladoeeros tetraeeros; this is recorded from the Iberian peninsula and Great Britain Kahlke, 1961), Tiraspol (Nikiforova, 1971). franchian to Galerian did not take place in Italy from beds transitional from Villafran­ to eastern Siberia. It was not recorded however One of us (A.A.) considers all records of this at once. Localities are known with mixed chi an to Galerian (Selva Vecchia). from the Indian subcontinent, where faunas species to be based on erroneous identifi­ and somewhat transitional faunas: Villafran­ Another typical element of the latest Villa­ are highly endemic. cations: according to him the specimens do not chian assemblages with few Galerian ele­ franchian is Cervalees gallieus (Azzaroli, 1983), The most common species, restricted to the correspond to D. etruseus, which is restricted ments or vice versa have been recorded from which is common in the late Villafranchian of Galerian, are: to the middle and late Villafranchian. Ovibos Italy, France, Germany and Spain (Bonifay, England, has been recorded from Seneze (Azza­ was recorded by Kahlke (1969) on fragmentary 1978; Briining, 1978; Azzaroli et al., 1982; roli, 1952, 1953) and also from Italy, in the Cervalces latifrons remains from Siissenborn. Azzaroli, 1983a; De Giuli and Torre, 1984; Crostolo river near Parma (Ambrosetti and Megaceros (Megaceros) savini Most Galerian taxa have their forerunners, De Giuli et al., 1983). Sites with naturally Megaceros (Megaceroides) verticornis Cremaschi, 1976). The latter diagnosis however and immediate ancestors in late Villafranchian mixed, i.e. not reworked assemblages are in Megaceros (Megaceroides) solilhacus rests on a rather poor antler fragment. Cervus elaphus acoronatus faunas of Eurasia. Only a part of these is any case few and the transitional phase Equus bressanus is a species of large size and Soergelia sp. however known and the roots of a large part of seems to have been of geologically short distinct from the much older E. livenzovensis, Praeovibos priscus the middle and late Pleistocene fauna should duration. with which it was sometimes confused; it is Bison schoetensacki be looked for in the largely unexplored melting The result of the dispersal event was a total Equus altidens somewhat transitional between E. stenonis and Equus siissenbornensis pot of inner Asia. Cervalees latifrons is a rejuvenation of the fauna: not only new E. siissenbornensis and seems to characterize Equus verae gigantic descendant of C. gallieus (Azzaroli, adaptations were developed, but the faunal some late Villafranchian assemblages (Azza­ Ursus deningeri 1983a); Galerian giant deer (Megaeeros) seem assemblage took a modern appearance: the late Canis lupus mosbachensis roli, 1984). to have their ancestors among medium sized Pleistocene and present day faunas are com­ Xenocyon lycaonoides Latest Villafranchian fossil assemblages Oulo schlosseri Villafranchian megacerids of eastern Europe. posed of species that appeared in the Galerian, may be referred to the marine Emilian and at Homotherium latidens Bison may have evolved from a "Leptobos­ or by their direct, slightly modified descen­ least part of the Sicilian stages and in the Arvicola cantiana Bison" group of the late Villafranchian of dants. Trogontherium cuvieri Rome area underlie the Cassian erosional Eurasia; it is interesting to remark that giant In the literature faunas of early Mid-Pleisto­ phase, which was dated approximately 1 Ma cene age and similar in composition to the These species, or subspecies, are confined to deer, bison and musk ox brought about for the (Ambrosetti and Bonadonna, 1967; Ambrosetti Galerian fauna of western Europe are often the Galerian or to part of it. first time the characteristic adaptation, respec­ et al., 1972) but seems now to have had its peak referred to as Cromerian, but this term was Other common species that made their first tively, of gigantic antlers and of heavy body around 0.8 Ma (Shackleton and Opdyke, 1976; < introduced to designate a stage based on pollen appearance in the Galerian and survived into forms two kinds of structures that were Thunell and Williams, 1983). and of much shorter duration than the Gale­ later stages are: unknown before. Latest Villafranchian faunas are known in Equus siissenbornensis seem to have been rian. Russian authors speak of a Tiraspolian Tuscany; in the Chi ana valley around Farneta, complex (Gromov, 1948; Schanzer, 1982), typi­ Elephasantiquus derived from E. stenonis, while Equus eaballus the type locality of the Farneta faunal unit, Mammuthus armeniacus and probably Equus altidens are immigrants fied by a fauna in Moldavia, and this term was and in the Mugello lacustrine basin (Azzaroli, Saiga tatarica from North America. Ursus deningeri may also applied to Siberian faunas (Vangengeim 1977a; Azzaroli et al., 1982; De Giuli et al., Bos primigenius safely be assumed to be a descendant of and Sher, 1970). According to recent papers 1984); at Imola in the Po valley, the site where Rangifer tarandus (Nikiforova, 1971; Schanzer, 1982) this term Capreolus capreolus U. etruseus, Canis lupus mosbachensis of Canis Gignoux established his correlation between Sus scrofa etruscus and Xenoeyon of C. faleoneri. Dieero­ seems to refer only to the younger part of the Calabrian and Villafranchian (Azzaroli and Equus caballus rhinus aff. hemitoechus is closely related to Galerian, comprised in the Brunhes palaeo-

t 88 89

magnetic epoch, i.e. younger than 0.7 Ma. Sher estimate, the difference in time is probably extinct before the end of the Galerian; the two should correspond to the Jaramillo or possibly (1971) introduced the term of Olyorian suite, real. immigrant species, horse (Equus caballus) and to some minor fluctuation within the late more a lithostratigraphic term, for a sequence The Galerian fauna spread over Europe and the hemione-like Equus altidens, represent Matuyama. in the Kolyma lowland of western Siberia with the middle and high latitudes of Asia and also different lineages, better adapted to highly a faunal assemblage of Galerian type, and extended to the Near East in Syria. Maps ofthe seasonal and even very cold climates. The pre-glacial Pleistocene of England straddling the late Matuyama and early distribution of some of its typical representa­ Brunhes epochs (Sher and Kaplina, 1979); on tives were published by Kahlke (1969, 1971: Age of the end-Villafranchian event Correlation of vertebrate bearing deposits of another occasion Sher writes of a "Mindel" large cervids; the Italian localities were unfor­ the English North Sea coast and of continental fauna (Sher, 1975). New details on the Olyor tunately omitted), Vangengeim and Sher (1970: Late Villafranchian mammals were recorded Europe were discussed in a previous paper faunal complex were added recently by Sher several species in Siberia), Sher (1975: elk, from the top of a Calabrian (Santernian) (Azzaroli, 1983a: p. 127). The picture in its (1984), and the fossil bearing sections of the Sorgelia, Praeovibos, Equus, in Russian Asia, sequence in a position immediately underlying broad lines is now fairly clear but some minor Kolyma lowland were calibrated with the eastern and central Europe and Great Britain), the Cassian erosional phase (Ambrosetti and problems still await for a final solution. palaeomagnetic scale by Virina et al. (1984). Vangengeim (1977: several species in Siberia). Bonadonna, 1967; Ambrosetti et aI., 1972) in The Red Crag is readily correlated with The fauna may be divided into an older, In such a vast area local variations are to be the Rome area, and in a position immediately the Praetiglian, i.e. the Montopoli faunal unit Chukochian, and a younger, Akanian unit. expected. Cervalces has not been recorded from below the Jaramillo event in the Danube valley of Villafranchian stratigraphy (not with the Characteristic of the former are Praedicro­ Spain but was recently recorded from Italy; (Kukla, 1977). The Lakhuti 2 fauna of Tadjikis­ middle Villafranchian, as in Azzaroli, 1970). stonyx capitalis, Allophaiomys pliocaenicus, Megaceros on the other hand does not seem to tan is a Galerian assemblage with few late Equivalents of the Tegelen fauna are possibly Oulo cf. schlosseri, Praeovibos beringiensis, extend east of Tadjikistan; Bos is lacking in Villafranchian holdovers (Azzaroli, 1983a) and represented by rather scanty remains from the possibly Cromeromys intermedius and an ele­ the high latitudes, Praeovibos is rare in Europe is situated at the top of the Jaramillo (Dodo­ cliffs at Easton Bavents, reported by Stuart phant, provisionally indicated as Elephantidae and common in Siberia, while Equus appears nov, 1980). (1982: pp. 104-105). West (1980) correlates the gen. novo sp. 1. The Akanian is characterised to have been extremely adaptable and is The fauna ofPonte Galeria in the Tiber delta Antian stage of Easton Bavents with the by Elephantidae gen. novo sp. 2 (more derived practically ubiquitarian. is comprised between the Cassian and the Olduvai event, in agreement with this view, than sp. 1), Dicrostonyx renidens, Microtus It was stated in a previous section that the Flaminian erosional phases (Ambrosetti et while a correlation with the Olivola faunal spp., and in a correlated section at Ulakhan­ Galerian witnessed new body adaptations, aI., 1972); the fauna of Isernia La Pineta in unit, proposed by Azzaroli (1983a: p. 127) seems Sudar on the Adych river Equus (Equus) unknown in previous times. These are repre­ central Italy, also a typically Galerian assem­ incorrect in as much as Olivola is of younger mosbachensis (= Equus caballus mosbachen­ sented by the heavy, massively built bovids blage, falls in the late Matuyama reversed age (Eburonian). Some identifications reported sis). Allophaiomys pliocaenicus is restricted to Bison, Bos, Ovibos, which developed from epoch and is dated by volcanics to 0.73 0.04 by Stuart do not fit however in this picture and the base of this unit. Other species, character­ ancestors of smaller size and more slender ± may perhaps need a revision: Eucladoceros istic of the entire Olyor complex, are: Clethrio­ build; of the giant deer of the genus Megaceros, and 0.73 ± 0.07 Ma (Coltorti et aI., 1982); the nomys ex gr. rutilus, Trogontherium cf. cuvieri, which also evolved from smaller, still poorly Galerian fauna of Solilhac in the French falconeri belongs to an older stage, Mimomys Canis lupus cf. mosbachensis, Xenocyon cf. known ancestors and equalled in the Galerian Central Massif has been calibrated with the blanci to a younger one. lycaonoides, Homotherium sp., Equus (Plesip­ the body size and antler span of the late Jaramillo episode (Thouveny and Bonifay, In Norfolk, on the coast between Happis­ pus) verae, Cervalces aff. latifrons, Soergelia sp., Pleistocene Megaceros giganteus; of the gigan­ 1984). burgh and Weybourn, the oldest vertebrate Praeovibos cf. priscus, Bison ex gr. schoeten­ tic Cervalces latifrons, which stood about 2 m The strati graphic position of the Isernia bearing sediments contain an early Pleistocene sacki and others. at the shoulder, with antlers spanning well fauna is in contrast with calibrations proposed (late Villafranchian) fauna and directly overlie The Chukochian unit begins in the Matu­ over 2.00 m, supported by 40-mm thick frontals for some classic sites, as Stranska Skala 2, the chalk. Their age ranges through Pre­ yama reversed epoch, at an estimated date of of solid bone. Rhinoceroses and bears also Voigtstedt and Sussenborn. Biostratigraphic­ Pastonian and Pastonian in the pollen scale approximately 1.2 Ma. The transition to the evolved to larger size during the Galerian and ally Isernia should be younger than these (West, 1980). These are overlain by Cromerian Akanian has been dated more accurately and continued their evolution in the later Pleisto­ because of the occurrence of Arvicola; on the deposits and by glacial till. The series is falls near the top of the late Matuyama, at a cene. Equids on the other hand reacted in a other hand the normal magnetic polarities extremely condensed, formations vary in thick­ date of approximately 0.80-0.78 Ma. The Olyo­ different way. They seem to have reached their observed at Stranska Skala 2 and Voigtstedt ness from a few meters to less than one meter rian complex is followed by a gap and its upper largest size at a somewhat earlier date, in the were referred to the Brunhes, thus indicating and are discontinuous, the sequence being cut limit cannot be exactly dated. latest Villafranchian, with Equus bressanus, an age younger than Isernia. In as much as by gaps and erosion channels. Most fossils It will be noticed here that the onset of the the stature of which may be evaluated between paleomagnetic and radiometric dates of Isernia were collected on the beach after storms had Olyorian faunal complex in eastern Siberia 170 and 180 cm at the withers. But Equus seem reliable, and assuming that Arvicola is a eroded the cliffs and their provenance may be would predate the end-Villafranchian event by bressanus is a zebra (sub genus Dolichohippus), valid marker of continental scope, the sites reconstructed on indirect evidence. It is how­ approximately 0.2 Ma; although the date of the closely related to almost equally large Equus of central Europe referred to above should ever clear that two distinct faunas are present, beginning of the Chukochian is a rather broad sussenbornensis (Azzaroli, 1984), which became be older, and their normal magnetic fields one late Villafranchian, the second Cromerian

t 90 91

(Azzaroli, 1953), or better Galerian; a younger kinsi is a typically Galerian species closely In Japan a spectacular change took place in cause of the whole sequence of climatic, fauna, recorded by Azzaroli, may be present related to Megaceros verticornis, of which it the vegetation, with the appearance of an vegetational and faunal changes. The rising but is poorly documented. represents a stunted variant. It is not primi­ alpine type flora, at the base of the Jaramillo, mountain barrier caused the onset of a mon­ A purely late Villafranchian fauna occurs at tive; the dentition is relatively hypsodont, but i.e. round 0.1 Ma earlier than in the Panno­ soon climate to the south and of increasingly East Runton: here only Pre-Pastonian and the small size, the reduction of the antlers, the nian plain (Suzuki and Manabe, 1982). Sedi­ drier and more continental conditions in cen­ Pastonian deposits were recorded (West, 1980: occasionally observed disproportion between mentation of loess, a typical product of a cold tral Asia; this was presumably the area in fig.51). At West Runton the exposed sequence the thick pedicles and burrs and the slender dry climate, generally began during the late which most of the faunal evolution took place ranges from Pre-Pastonian to Cromerian: Azza­ beams (Azzaroli, 1953; fig.33D, E, fig.34E) may Matuyama in central Asia: Uzbekistan, Tad­ and from which migratory waves spread to roli (1953) recorded only Cromerian (Galerian) indicate an incipient evolution towards the jikistan (Dodonov, 1980, 1982); in northern Europe and possibly also to the Far East. vertebrates but more recently late Villafran­ reduction in size that characterizes large China it started somewhat earlier, but still The elephant-Equus event of Europe has a chian elements were reported: Archidiskodon mammals in insular environments. after 1.2 Ma (Derbyshire, 1983). In central counterpart in the transition between the so­ meridionalis, Cervalces gallicus (Stuart, 1982: Europe the loess sequence is more complex called Tatrot and Pinjor faunal complexes of p. 106). At other sites: Bacton, Mundesley, Vegetation and climate and scattered loess horizons were recorded India and Pakistan, marked by the virtual Sidestrand, Overstrand, both late Villafran­ since the Olduvai event in the Krems section disappearance of hipparions, of Merycopota­ chian and Galerian elements were collected. The Menapian cold phase entrained changes on the Danube, but loess became more fre­ mus and of several other species and the The older fauna includes elements character­ of great moment. The pollen sequences of the quent and abundant during the Jaramillo arrival of Equus, Rhinoceros, Punjiabitherium istic of the latest Villafranchian (Farneta Pannonian plain, analysed in deep boreholes, (Kukla, 1977). (a two-horned rhinoceros), Elephas hysudricus unit): Archidiskodon meridionalis croinerensis, reveal the series of oscillations from warm to and Pachycrocuta relina. No counterpart of the Eucladoceros tetraceros, Cervalces gallicus, cool and even cold climate that charcterise the Physiographic evolution Pleistocene faunal events could be detected in Equus bressanus, Canis arnensis; other ele­ whole late Pliocene and Pleistocene, but cold the Sivalik sequence as the fossil record ments are only broadly indicative of a late dry assemblages appear for the first time at the Azzaroli and Napoleone (1982) showed that becomes scanty at the beginning of the Pleisto­ Villafranchian age: Dama nestii, Eucladoceros top of the Jaramillo (Cooke, 1981). cene and almost everywhere comes to an end dicranios, (incl. E. ctenoides), Leptobos etrus­ The pollen sequence of The Netherlands was strong tectonic movements set in in the Himalayan belt around 1 Ma ago. A mountain within the early Pleistocene. cus, Equus stenonis, Dicerorhinus etruscus. The worked out in great detail. Zagwijn and De On the other hand the rather marked faunal evidence provided by small mammals is partly belt dividing the Indian subcontinent from Jong (1984) correlated the Menapian cold migration that took place in the Tasso faunal contradictory. Stuart (1982: pp. 106-107) re­ central Asia had been in existence since the (glacial) phase with a reversed paleomagnetic unit does not correlate with any obvious ported Mimomys pliocaenicus, M. reidi and M. Miocene and had already given rise to highly interval immediately predating the Jaramillo climatic crisis. Maybe western Europe was blanci from the Pre-Pastonian and the same episode. This was followed in rapid succession endemic Indian faunas (Heintz and Brunet, only the point of arrival of wide ranging species plus M. newtoni from the Pastonian. by the Bavel interglacial, corresponding to the 1982; Brunet and Heintz, 1983; Azzaroli, dispersals prompted by increasingly continen­ This is partly in contrast with the date offered Jaramillo, and then in the late Matuyama by 1985b). The sediments derived from these tal, strongly seasonal conditions that were by large mammals, as M. pliocaenicus seems to the Linge glacial, the Leerdam interglacial, mountains, the richly fossiliferous Sivalik developing in central Asia. be restricted to the older part of the late the Dorst glacial, the interglacial 1. Glacial A series of northern India and Pakistan, were Villafranchian (Olivola unit). of Dutch geologists corresponds with the at this time generally fine grained molasse Assuming that identifications are accurate, beginning of the Brunhes Epoch and is fol­ deposits. Stronger tectonic movements are The climatic record in the marine two possibilities remain open: either the Pre­ lowed by Interglacial n. Zagwijn and De Jong revealed by the onset of coarse conglomeratic environment Pastonian and Pastonian encompass more or point out that in the Bavel and Leerdam sedimentation near the top of the series. less all the late Villafranchian, from the interglacials the return of warm floral ele­ Scattered beds of coarse conglomerates occur Palaeotemperature records in marine se­ Olivola to the Farneta units; or alternatively ments did not happen suddenly as in older interbedded in the molasse in levels calibrated quences indicate a succession of climatic some of the small mammals were reworked. interglacials, but in a definite succession of with the Olduvai event near Pinjor, northern changes, some of which can probably be The Beestonian, the cold phase dividing the migrations, and this pattern of migration was India, and during the Jaramillo event sedi­ correlated with dispersal events of terrestrial Pastonian from the Cromerian, may be corre­ repeated in younger interglacials; they con­ mentation became entirely conglomeratic (Az­ faunas. Synchroneity in some cases seems lated with the Menap or the Linge glacial of clude that this may have been due to the fact zaroli and Napoleone, 1982), until the late however to be only approximate, and time lags central Europe and with the Cassian erosional that the Menap and later glacials pushed the Sivalik sequence was closed by the overthrust­ exceeding the range of random errors may be phase of the Mediterranean. warm floral elements farther away to the south ing of older Siwalik molasse and then by a observed between the events detected in the A final remark may be made: there are than the older cold periods. The Menapian series of overthrusts of increasingly older two series of phenomena. reasons to believe that England may have glacial appears thus to have caused the great­ formations. In the north Atlantic and north Pacific, witnessed a period of insularity during the est vegetational change in the Pleistocene of The strong upheaval of the Himalayas dur­ analysis of DSDP Sites (Thunell and Williams, Pleistocene. The giant deer Megaceros daw- western Europe. ing the Pleistocene may have been the primary 1983; Shackleton et aI., 1984; Rea and Schrader,

t ------

92 93···~

Q) CAPE BOJADOR. SITE 397 ';0 SITE 132 ~ ,M a m m a 1 s Q) ~ Poll en 6 "O~D'_ '[ 2.0 1.0 0.0 -1.0 5,0 4,0 3.0 '"E o '0 u'" Major Erosional :0 W, Europe NE Asi a '- W, [urooe E. Europe events phases MA ] J V) z i. '.4 0 ~ w U Ost:l.tln 20 " o " l­ z ll) W Nomentanan ..J 0.. CltomeJL...Ut11 Ranuccio T.ur.a.J.JpoUal1 _ - - -100 - - - - - u'" o Ill'vicola Plaminian P.te Galeria 8 ______~ 0.. i = 3.8 1 _ (J) _ A!I1~p:;a.~ Domegliara ~ C/lulwcluC1.H: -J WA . f­ Cassian co (:?,,a . a; ;.- ... E .. W Farneta ~ z i S ui w Waa.Ua.n, I&i (,) ~ iii"" 0 _ - - - - Ti!l.9S0 in 80 200 ::::; ~ ... 0.. i: / .. Olivol", :I: / ~ W :I: .- I- ~ .- \~olf Aullan "" « 0.. :t' -' ------, ,0 l!! '" , ..... Hgllin '" ~ ~ :z 80 2_ S. Vallier ..-, w '« z :z .- w )( ~ 3.0 '" .- w ~ ... z -- --- « KtLtuya{- .0 ..J '« ... .. ~ ·" w RwveJtian Triversa Moldav.cal1 f-- ~ ~ Cl '" :z • 3- z '« w ci· I-- 120 Z 400 LeptOlJOB w '« (SHACKLE TON a"d CIl A, 19;9) '" I-- '" u - ~ ~ :1 o ~

H ui

Fig.4. Major shifts in ocean water temperatures in the late Pliocene and early Pleistocene (after Vergnaud Grazzini, 1984).

ranging crisis of extinction in the marine of Blanc (Ambrosetti et aI., 1972), the Moulou­ mollusc fauna. An earlier extinction crisis yan of the Atlantic coast of Marocco (Biber­ of molluscs in the northern Atlantic was son, 1971), the marked drop in sea level variously dated between 3.5 Ma (Stanley, 1982) described by Stipp et aI. (1967) in New Zealand. and 3.2-3.1 Ma (Raffi and Marasti, 1982); the Agreement is less satisfactory with the picture latter date agrees with a temporary tempera­ of global cycles of sea level provided by Vail et Fig.3, Correlation chart of continental stages, faunal units, major faunal events and erosional phases (negative eustatic ture drop recorded in DSDP 552A (3.3-3.1 Ma) . aI. (1977: fig.3): these authors show sharp drops oscillations of sea level), and in the Jucar basin (3.2-3.1 Ma). of sea level in the Pliocene at approximately 3 The crisis dated 2.4 Ma on DSDP 552A is or 2.9 Ma and at 2.0 Ma. The discrepancies, in 1985) shows a sharp fall in deep water tem­ between the temperature falls of the north considered the greatest climatic crisis in the our opinion, are due to inaccurate datings of perature and a sudden increase of ice rafted Atlantic and ofthe lacustrine beds ofthe Jucar Plio-Pleistocene marine record (Shackleton et sea level fluctuation in the synthesis of Vail et debris at 2.4-2.5 Ma. There is a time difference basin of Spain. The lowering of temperature in aI., 1984). It coincides with a world wide drop aI. A closer agreement with the interpretation of at least 0.1 and possibly as much as 0.3 Ma Atlantic deep water corresponds to a wide of sea level: the Acquatraversa erosional phase adopted in this paper is found in an article on

t • • r.~'Cj:.··.·· ~::1 94 95 .., >r MEDITERRANEAN m POLLEN ZONES LOCAL FAUNAS STRATI­ PALEO­ NORTH SEA 0 1= FAUNAL GRAPHY MAGNETISM CLIMATE en EVENTS 0 EAST. (Ryan, 1973) Ma >r NEDERLAND EAST ANGLiA ITALY FRANCE SPAIN CENTR. ASIA m EUROPE EUROPE 1960-1974)

P L M E I S A WAALlAN T o C T Ranucclo E EBURONIAN N CROMERIAN 5.5. E IGL.II u Choukoutien Mosbach } MN 17 (Graues) GLAZIAL A L'Escale Mauer Tiraspol Koshkurgan P TIGLlAN Isernla p.73 IGL.I /Wica/a Stranska P.Galeria skala L DORSTGL SlIvia ~us5enbor LEERDAM IGL. Lakhuti 2 A 0.9 L1NGE GL. End Villa- M.Peglia Sclelhac Voigtstedt PR~ETIGLIAN J Nogalsk BAVEL IGL. franchian cavaSud Vallonet Lakhutl1 P I 0.97 Mosbach MENAP GL. BEESTONIAN Domegllara Venta (Grobes) S Pirro Micena ?Nihowan 2 Farnata A WAAL IGL PASTONIAN Imola Seneze 11 R - B L N C REUVERIAN I P.PRE_PASTONIAr:"noPhaiomys Tasso Salnzelles Erpfingen A R - A EBURONIAN ?BRAMERTONIAN BAVENTIAN N Woll Olivola 1.67

0 ?ANTIAN M.Coupet Tegelen 0 1.9 Br - C TIGLlAN T Chilac Kuruksay 2 THURNIAN Senel8 I 2.01 Puebla de 2.04 R A SI.Vallier Valverde ?Kuruksay 1 .12 LUDHAMIAN 2.14 C Br - B B PRE-TIGLlAN PRE-LUDHAMIAN Elephant Montopoli Rocaneyra Khapry ?Nihowan 1 E BRUNSSUMIAN .48 Equus Rlncon 1 Uvenzovka A

N Br - A N Villaroya

~.92 Etouaires K 3.01 REUVERIAN Triversa Vialette E A .05 Lop/obos Layna M .15 N SUSTERIAN Su

3.4 Fig.5. Pollen climatic curves for the Mediterranean and North Sea, Pliocene to Pleistocene, plotted against the palaeomagnetic scale. Adapted from Zagwijn and Suc (1984). Fig.6. Correlation chart of local faunas, Pliocene to early middle Pleistocene, for Europe and Eastern Asia...... ------~------

96 97

quaternary eustatic cycles, mainly based on of the Jaramillo and the beginning of the Ambrosetti, P. L., Azzaroli, A., Bonadonna, F. P. and Table on the stratigraphic distribution of terrestrial evidence from the west Atlantic, by Beard et al. Brunhes, i.e. around 0.8 Ma. Follieri, M., 1972. A scheme of Pleistocene chronology mammalian faunas in Italy from the Pliocene to the early middle Pleistocene. Geogr. Fis. Din. Quat., 5: (1982). Ruggieri et al. (1984) showed a marked for the Tyrrenian side of Central Italy. BoIl. Soc. Geol. Ital., 91: 169-184. 55-58. The carbon isotope curve of DSDP Site 552A increase in cold water pteropods in a sandy Ambrosetti, P. L., Bartolomei, G., De Giuli, C., Ficcarelli, Bajgusheva, V. S., 1971. Fossil teriofauna of the Liven­ shows an increase in water temperature be­ bed, possibly evidencing a drop in sea level, at G. and Torre, D., 1979. La breccia ossifera di Silvia zovka sand pit. Material on the Anthropogene fauna of tween 2.1 and 1.7 Ma (the Tiglian of Zagwijn) Ficarazzi near Palermo, in the type section of (Aurisina-Sistiana) nel Carso di Trieste. BoIl. Soc. USSR. Akad. Nauk U.S.S.R., Moscow, pp. 5-29. Bajgusheva, V. S., 1978. The large horse of the Khaprovian followed by a new minimum between 1. 7 and the Sicilian, and date it again to 0.8 Ma. The Palaeontol. It., 18: 207-220. Arias, C., Azzaroli, A., Bigazzi, G. and Bonadonna, F. P., complex from the alluvium on the NW of the Azov Sea. 1.6, which may mark the beginning of the date was obtained through detailed correlation 1980. Magnetostratigraphy and Pliocene-Pleistocene (Russian) Rep. N. Caucasian high-school Sci. Cent., 1: Eburon phase of the North Sea, the Aullan with nannofossil sequences (Di Stefano and Boundary in Italy. Quat. Res., 13: 65-74. 98-102 (in Russian). erosional phase of central Italy, the Baventian Rio, 1981; Ruggieri et al., 1984). Here not only Azzaroli, A., 1950. Osservazioni sulla formazione villafran­ Bandet, Y., Donville, B. and Michaux, J., 1978. Etude geologique et geochronologique du site villafranchien de cold period of East England and the beginning the temperature drop but also the eustatic chiana di Olivola in Val di Magra. Atti Soc. Tosc. Sci. Nat. Mem., 57: 104-111. Vialette (Puy de Dome). Bull. Soc. Geol. Fr., 20: 245-251. of the Pleistocene-Late Villafranchian. oscillation of sea level turns out to be 0.1 to 0.2 Azzaroli, A., 1952. L'alce di SEmeze. Palaeontol. Ital., 47: Beard, J: H., Sangree, J. B. and Smith L. A., 1982. Temperature of the north Atlantic at this Ma younger than the end-Villafranchian 133-141, Pisa. Quaternary chronology, paleoclimate, depositional se­ time was no less severe than during the event. The difference in time between major Azzaroli, A., 1953. The Deer of the Weybourn Crag and the quences, and eustatic cycles. Bull. Am. Assoc. Pet. Geol., 66: 158-169. previous cold interval: if the impact on the changes in continental faunas and floras and Forest Bed of Norfolk. Bull. Br. Mus. (Nat. Hist.), Geol., 2: 1-96. Bernsen, J. J. A., 1930. Eine Revision der fossilen marine fauna was less dramatic this may have events in the marine environment is a chal­ Azzaroli, A., 1954. Filogenesi e biologia di Sus strozzii e di Saugetierfauna aus den Tonen von Tegelen. IX. Notizen been because the fauna had already been lenging theme for future investigations. Unfor­ Sus minor. Palaeontogr. Ital., 48: 41-76. uber die Cervidae. Natuurhist. Maandbl., 23: 38-46; selected for cold waters, warm water dwellers tunately the Ficarazzi section was not cali­ Azzaroli, A., 1963. Rinoceronti Pliocenici del Valdarno 71-77; 82-86. Berzi, A., 1966. L'orso di Gaville nel Valdarno Superiore. having been eliminated earlier. brated in the palaeomagnetic scale. We hope Inferiore. Palaeontogr. Ital., 57(1962): 11-20. Azzaroli, A., 1970. 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Introduction reactivated basement constitute the internal part of this belt whose age has been much The West Congo Basin is a vast synclino­ discussed. Certain workers believe it to be a rium, referred to as the Niari or Nyanga paired orogen in which a Pan-African Belt, Synclinorium, which trends NNW-SSE and approximately 700 Ma old (Cahen et al., 1976) parallels the Atlantic coast from Gabon to or 600 Ma old (Hossie and Caby, 1979), joins an Angola. The present study is concerned with older Mayombian Belt of about 1100-1200 Ma. the Congo part of this basin. The northeastern Others think that it is a Mayombian Orogen flank of the basin is monoclinal and rests very slightly reactivated during the Pan­ unconformably on the basement of the Chaillu African Orogenesis (Vellutini et al., 1983). Massif. The steepened and folded southwestern Recent investigations by Hossie and Caby flank makes up the external structural units of (1979), Hossie (1980) and Boudzoumou (1986) the West Congo Belt (Fig. lA). The underlying suggest the entire orogenic ensemble has Mayombian succession, which probably is been subjected to only a single, polyphased Middle Proterozoic in age, together with the orogenesis of Pan-African age.

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