A Case Study of Large Mammals from the Italian Peninsula

Il Quaternario Italian Journal of Quaternary Sciences 22(2), 2009 - 291-306

BIOCHRONOLOGY OF TERRESTRIAL MAMMALS AND QUATERNARY SUBDIVISIONS: A CASE STUDY OF LARGE MAMMALS FROM THE ITALIAN PENINSULA

Maria Rita Palombo Dipartimento Scienze della Terra Università La Sapienza, CNR, Istituto di Geologia Ambientale e Geoingegneria, P.le Aldo Moro, 5 00185 Roma, Italy. E-mail: [email protected]

ABSTRACT: Palombo M.R., Biochronology of terrestrial mammals and Quaternary subdivisions: a case study of large mammals from the Italian peninsula (IT ISSN 0394-3356, 2009). Defining and subdividing the Quaternary on the basis of the mammalian fossil record from the continental realm is not a simple task due to the low degree of succession continuity and the scattered palaeontological evidence. Moreover, even if the approaches to the Quaternary are basically interdisciplinary and may combine many different chronological scales, establishing correlations between biochronology, biostratigraphy, chronostratigraphy, climatostratigraphy, and composite regional stratigraphy can often be very problematical. As far as biochronology is concerned, the marked geological, environmental and climatic diversity affecting different continental regions makes a correlation based on biological events difficult. Indeed, “biochronological units” represent a time span during which faunas have a degree of taxonomic homogeneity and the corresponding “faunal complexes” have to be regarded as non- over- lapping and "ecologically adjusted groups of animals with specific geographic limits and chronological range" (TE D F O R D , 1970). Nevertheless, the stratigraphic lowest and highest occurrences of fossil remains (stratigraphic datum) within a given geographical area do not necessarily correspond to their actual first/last appearances (palaeobiological events) in time. This is due to diachroneity in palaeobiological events (i.e. local first and last appearances are strictly linked to dispersal, and the physical and biotic factors causing local evolution and extinctions) coupled with discontinuity in the continental sedimentary record, the rarity of deposits formed in a regime of virtually continuous sedimentation, the presence of important ecological barriers (that prevent some taxa from dispersing), environmental conditions (that affect the structure of palaeocommunities), and taphonomic and sampling biases. As a result, ongoing research, continuously yielding new data, make chronological frameworks thus far outlined, even if recent, open to significant improvements, and causing biochrons to be updated. This fact prevents any detailed biochronological framework from having widespread geographical significance. Thus, only higher ranking biochronological units (Land Mammal Ages, LMAs) - whose separation is based on palaeobiological events which have a wide territorial significance - could be useful for chronological correlations. Nonetheless, the transition between successive LMAs does not always correspond to the boundaries separating marine Series or Stages. For instance, the Villafranchian LMA approximately began with the Late Pliocene (Piacentian) and the transition from the early to the middle Villafranchian LMA happened around the Pliocene/Pleistocene (Piacentian/Gelasian) boundary, whereas the transition to the late Villafranchian took place during the latest Gelasian, and those from the Villafranchian to Galerian LMA and from the Galerian to Aurelian LMA respectively predated the beginning of the Middle and Late Pleistocene.

RIASSUNTO: Palombo M.R., Biocronologia dei mammiferi terrestri e suddivisioni del Quaternario: l’esempio della faune a grandi mammiferi della penisola italiana. (IT ISSN 0394-3356, 2009). Definire e suddividere il Quaternario sulla base del record fossile dei mammiferi continentali non è un compito facile ed anche se gli approcci al Quaternario sono fondamentalmente interdisciplinari e tendono a combinare diverse scale cronologiche, stabilire correla- zioni tra biocronologia, biostratigrafia, cronostratigrafia, climatostratigrafia, e stratigrafia regionale è spesso problematico. Sebbene le unità basate su eventi biologici e climatici siano di largo uso nel Quaternario continentale, il loro status non è stato formalmente definito e questo alimenta dubbi nell’approccio metodologico e confusione tra teoria (dato Paleobiologico) ed operatività (dato stratigrafico). Dal punto di vista teorico, le " unità biochronologiche" rappresentano un lasso di tempo durante il quale le faune locali hanno un certo grado di omogeneità tassonomica ed i "complessi faunistici” che caratterizzano le singole unità devono essere considerati “non-overlapping and ecologically adjusted groups of animals with specific geographic limits and chronological range" (TEDFORD, 1970). Tuttavia, l'’evidenza stratigrafica della prima e ultima presenza di un fossile nelle successioni affioranti in una determinata area geografica (dato stratigrafico) non corrisponde necessariamente all’effettiva prima e ultima comparsa di quel taxon nel tempo (evento paleobiologico). Ciò si deve alla diacronicità degli eventi paleobiologici (ad esempio gli eventi locali di comparsa e scomparsa sono strettamente legati rispettivamente alla dispersione ed ai fattori fisici e biotici che regolano diffusione ed estinzione), alla discontinuità del record sedimen- tario continentale, alla rarità di depositi formati in regime di sedimentazione continua, alla presenza di barriere ecologiche (che limita o impedisce la dispersione di alcuni taxa in certuni territori), alle condizioni ambientali locali (che influenzano la struttura delle Paleocomunita), a problemi tafonomici e di campionamento. Questi limiti rendono instabili schemi biocronologici dettagliati basati su un record fossile regionale, in quanto essi sono fortemente dipendenti dal progredire delle conoscenze sul campo. Fin dalla prima introduzione delle unità faunistiche (FUs) (Azzaroli 1977), sulle quale si fonda la biocronologia dei grandi mammiferi della penisola italiana, il progredire delle ricerche e la scoperta di nuove faune ha portato sia alla ridifinizione e/o eliminazionedi alcune FUs sia alla crea- zione di nuove. Ne consegue che, proprio in funzione della possibilità di discriminare singoli bioeventi, ogni schema biocronologico delle faune italiane risulta quanto più dettagliato tanto meno stabile nel tempo, nonché difficilmente confrontabile con gli schemi pro- posti per l’Europa continentale. In realtà, l’utilizzo di unità biochronologiche di più alto rango (Land Mammal Age, LMA) - la cui separa- zione è basata su eventi paleobiologici validi a grande scala e cambi di struttura dei complessi faunistici – meglio si presta a correlazio- ni su vasta scala. La transizione tra successive LMA, tuttavia, non sempre corrisponde ai limiti che separano Serie e Stadi. Per esempio, il Villafranchiano inizia con il Pliocene superiore (Piacenziano), la transizione al Villafranchiano medio potrebbe coincidere grosso modo con il limite Pliocene/Pleistocene (Piacenziano / Gelasiano), ma il passaggio al Villafranchiano superiore avviene nel tardo Gelasiano, e le transizioni Villafranchiano/Galeriano e Galeriano/Aureliano precedono, rispettivamente, l'inizio del Pleistocene medio e superiore.

Key words: Biochronology, Biostratigraphy, Plio-Pleistocene, Large Mammals, Itay. Parole chiave: Biocronologia, Biostratigrafia, Plio-Pleistocene, grandi Mammiferi, Italia. 292 M.R. Palombo

1. INTRODUCTION 2. ABOUT CHRONOSTRATIGRAPHY, BIOCHRO- NOLOGY, BIOSTRATIGRAPHY AND RELATED Defining and subdividing the Quaternary on the ITEMS: A SHORT SKETCH basis of the mammalian fossil record from the continental realm is not a simple task due to the low degree of As stated by Lindsay (2003) chronostratigraphy - succession continuity and the scattered palaeontologi- the dominant method applied in the oceanographic- cal evidence. Moreover, even if the approaches to the marine realm - and biochronology - the dominant Quaternary are basically interdisciplinary and can com- method for the terrestrial realm - are the prime concep- bine many different chronological scales, establishing tual methods for relating biological events to the geo- correlations between biochronological, chronostrati- logical time scale. The chronostratigraphical concept graphical, climatostratigraphical frames and composite was introduced by SCHENCK & MULLER (1941), who pro- regional stratigraphy can be problematical (concerning posed a new ‘‘chronostratigraphical’’ hierarchy Italy, see e.g. DE GI U L I et al., 1983; RA V A Z Z I , 2003; (System, Series, and Stage) for the stratigraphical rep- PA L O M B O , 2004a; PA L O M B O & SA R D E L L A , 2007 and refe- resentation of equivalent chronological intervals rences therein). (Period, Epoch, and Age). Recently, LINDSAY (2003, pg. With regard to terrestrial fauna, especially mam- 227) defined chronostratigraphy as: “the sequential mals, diachroneity in palaeobiological events coupled ordering of geologic events, using biostratigraphic, iso- with discontinuity in the continental sedimentary record, topic-radiometric, and paleomagnetic data”. The con- the rarity of deposits formed in a regime of virtually cept of biochronology dates back the beginning of the continuous sedimentation, the presence of important 20th century, when WI L L I A M S (1901) defined a ecological barriers (that prevent some taxa from dis- “biochron” as the absolute time extent of a peculiar persing), environmental conditions (that affect the faunal or vegetal association. Later, TE I C H E R T ( 1 9 5 8 ) structure of palaeocommunities), and taphonomical and sampling biases make the definition of isochronous described biochronology as ‘‘the dating of geological boundaries, maybe of boundaries themselves, prob- events by biostratigraphical methods’’, but did not lematical. Therefore, any biochronological framework is characterise biochronological terms, such as predisposed to change depending on new discoveries. ‘‘biochronological unit’’, and concepts relative to the During the latter half of the last century, study on biostratigraphical ones. Due to that loose definition and the terrestrial mammals of the Italian peninsula was to its ambiguous application, biochronology has never developed mainly on the basis of large-sized mam- been discussed in any stratigraphical code (LI N D S A Y , malian taxa. The Italian mammal biochronological 2003). Up to the present day, the operation of organis- framework builds on the pioneering work of Azzaroli i n g geological time on the basis of evidence provided (AZZAROLI, 1977, 1982), who first introduced the “Faunal by continental mammal faunas has continued to be a units”(FU’s) as “biochrons”, defined on the basis of all field plagued by interpretative and semantic confusion. the species from selected local faunal assemblages BE R G G R E N & VA N CO U V E R I N G (1978, pag. 39) tried to (LFA’s). Like Land Mammal Ages (LMAs) which are the avoid ambiguity defining biochronology as ‘‘the organi- primary unit of North American vertebrate chronology zation of geological time according to the irreversible (see TEAFORD,1970; LINDSAY & TEDFORD, 1990), FU’s rep- process of evolution in the organic continuum’’. resent and are defined by typical associations of mam- Subsequently, LI N D S A Y (2003, pag. 227) defined malian taxa (biological entities), each living in non-over- biochronology as the “sequential ordering of biologic lapping slices of time. As suggested by Lindsay (2003, (and geologic) events, using morphologic differences pg. 222) for LMA’s, FU’s could be defined as “a rela- that result from organic evolution (when applied to tively short interval of geological time that can be rec- Earth history)”. ognized and distinguished from earlier and later such On the other hand, it is widely known that the dis- units (in a given region or province) by a characteristic cipline of biostratigraphy establishes the stratigraphi- assemblage of mammals”. Although sometimes criti- cal ranges of the taxa remains within superimposed cised (e.g. GUERIN, 1982, 1990), FU’s are general con- sections, and verifies the relative age of rocks using cepts, especially used by the Italian scientific communi- the fossil data. Biostratigraphy is the element of ty. Since the time of the FU’s introduction, Italian stratigraphy which focuses on “the distribution of fos- palaeontologists have proposed new or revised some sils in the stratigraphical record and the organization FU’s/LMA’s (see e.g. DE GIULI, et al. 1983, TORRE, 1987; of strata into units on the basis of their contained fos- AZ Z A R O L I et al., 1988; SA L A , et al., 1992; TO R R E , et al. sils” (SA L V A D O R , 1994, pag.55). But as claimed by 1992; CALOI & PALOMBO, 1990, 1996; GLIOZZI et al., 1997, SA L V A D O R (1994, pag.55) biostratigraphical units, PE T R O N I O & SA R D E L L A , 1999; PA L O M B O et al., 2004; including biozones, “vary greatly in thickness and PALOMBO, 2004a, 2007a; MASINI & SALA, 2007; PETRONIO geographic extent...... The time they represent may et al.,.2007) (Table 1). The introduction of new likewise vary widely”. To avoid this issue, WA L S H biochrons mainly depends on acquiring new data and (1998) defined the biochronostratigraphical units as discovering new fossiliferous assemblages, which makes possible, by means of ever more detailed bios- “the sets of rock formed during biochrons, without ref- tratigraphical settings, increasingly accurate discrimina- erence to any particular stratigraphic section”. As a tion between bioevents. result, a biochronological unit should be characterised This paper’s aim is to outline the pricipal prob- and defined as a span of time defined by biological lems related to biostratigraphy/biochronology of large evidences (palaeobiological datum), while a biostrati- Italian mammals, and to update the previous graphical unit as a body of rock strata defined by its biochronological schemes according to the most recent fossil content (stratigraphical datum) (SC H O C H , 1989; discoveries (Table 2.3). SA L V A D O R , 1994). Biochronology of terrestrial mammals and quaternary subdivisions: ... 293 294 M.R. Palombo

2.1. In search of a stable biochronological frame for recent, open to significant improvements. Some the terrestrial large mammals: utopia or feasible biochrons, consequently, will require revisions and undertaking? updated definitions.

As defined above, “biochronological units” represent a time span during which faunas are assumed to 3. FAUNAL TURNOVERS, BIOCHRONS AND QUA- be characterised by taxonomical homogeneity and TERNARY SUBDIVISIONS should be regarded as non- overlapping and an “ecologically adjusted group of animals within specific geo- During the Pliocene and Pleistocene a series of cli- graphic limits and chronologic range” (TEDFORD, 1970). matic cycles caused latitudinal displacements of vege- First/last appearance bioevents (palaeobiological tation and biomes in Europe and exerted great influ- events on which biochronology is based) have been the ence on dispersal and dispersion of mammalian principal bases for establishing the chronological set- species. As a result, the composition of regional faunas ting of continental mammalian faunas. However, the changed because of origination/ extinction and disper- marked geological, environmental and climatic diversity sal bioevents. On the Italian peninsula, climatic affecting different continental regions, as well as the changes were a driving factor, at least as far as the two discontinuity in the continental sedimentary record, the most important detectable faunal renewals are con- rarity of deposits formed in a regime of virtually contin- cerned: the Early to Middle Villafranchian LMA uous sedimentation, the diachroneity in palaeobiologi- (Piacentian to Gelasian, ~2.7–2.5 Ma) and Early to cal events (i.e. local first and last appearances are Middle Galerian LMA, transition (latest Early strictly linked respectively to dispersal, and the physical Pleistocene, ~1.1–0.8 Ma). Faunas changed also at the and biotical factors causing extinctions), taphonomical end of the Gelasian (Middle to Late Villafranchian tran- and sampling biases are responsible for the fact that sition) and at the end of the Middle Pleistocene the stratigraphical order of the highest and lowest (Galerian to Aurelian transition), but to a lesser degree occurrences (stratigraphical datum) of fossil remains of (PALOMBO, 2007b and references therein). some taxa within a given geographical area, does not necessarily reflect the temporal order of the actual 3.1 The beginning of the Villafranchian and the warn- first/last appearances (palaeobiological events) of each ing signs of the already called “Glacial Pliocene” taxon in time. Indeed, a stratigraphical discontinuity The list of early Villafranchian taxa have been within a sedimentary succession (whether due to ero- established on a number of LFA’s from Piedmont (sev- sional phases or sedimentation lacking) corresponds to eral localities in the Villafranca d’Asti area) and central a time span during which palaeobiological events do Italy (e.g. Lower and Upper Valdarno, Arcille, Spoleto). not yield a stratigraphical record. Moreover, fossil Palaeomagnetic data at Fornace R.D.B. (Villafranca remains of taxa characterising a given biochronological d’Asti) (LINDSAY et al., 1980, 1995) as well paleomagnet- unit do not necessarily appear in each section because ic calibration of the long sedimentary sequence out- they were absent from the depositional environment for cropping at Santa Barbara quarry (lignitic lacustrine ecological or taphonomical factors. Therefore, the time silty clays and sands, Meleto clays of Castelnuovo dei range of a paleobiological entity (either in general or in a Sabbioni Lower Synthem of the Upper Valdarno basin) given geographical area) cannot be unquestionably (TORRE et al., 1996; BERTINI & ROION, 1997; NAPOLEONE et established by its fossil evidence. The lowest and high- al., 2003; GHINASSI et al., 2004) illustrate that, at about est stratigraphical records of a taxon, respectively cor- 3.2 Ma, the faunal association regarded as typical of respond to the time “ante” or “post quem” when it the early Villafranchian was already present on the actually originated and became extinct (PA L O M B O Italian peninsula. 2004a). Accordingly, it should be appropriate to dis- The early Villafranchian LMA (Triversa FU) retains criminate taxon longevity from time slice of its fossil subtropical affinities typical of Ruscinian mammals, record, e.g. first/last historical appearance bioevents which lived during the Early Pliocene, the most recent (FHA/LHA = First/Last Historical Appearance) from interval with a climate definitively warmer than today. “known data” of lowest and highest occurrences in a Pachyderms, middle sized Artiodactyla, and local stratigraphical section (LlSDk/HlSDk = local Perrissodactyla, mainly browsers or mixed-feeders Lowest/Highest known Stratigraphical Datum) (WA L S H , inhabiting dense or clear forest, were still present along 1998; PA L O M B O & SA R D E L L A , 2007). On the other hand, with arboreal/scansorial viverrids and arboreal omni- mammalian biochronology is actually founded on fossil vores, such as Mesopithecus monspessulanus, records available in the continental rocks, FHA/LHA Parailurus hungaricus and Ursus minimus. On the other bioevents can only be inferred from the lowest/highest hand, the appearance of some taxa, such as Leptobos occurrences of fossil remains in fossiliferous levels, s t e n o m e t o p o n and Stephanorhinus jeanvireti (= maybe based on more than one continental stratigraph- Stephanorhinus elatus) (Table 3), perhaps more linked ical section. Hence, new discoveries may substantially to wooded parkland environment, testified to a change change the chronological extent of already defined in faunal structure which can be correlated with the cli- biochrons. Indeed, the finding of a/some taxon/a matic cooling, which occurred at about 3.2/3.1 (KM2 (whose LHA/LHA were already regarded as markers of isotopic stage) (Table 2). a biochrons) in significantly older/younger extends the theoretical time range of such a biochron, without 3.2 Early to Middle Villafranchian and the Piacentian changing its theoretical definition. As a result, ongoing to Gelasian transition research, continuously providing new data, makes The following Montopoli FU is based on rich fauna chronological frameworks thus far outlined, even if from a site near Montopoli (Lower Valdarno, Tuscany), Biochronology of terrestrial mammals and quaternary subdivisions: ... 295

Table 2 - Integrated chronological scheme for the Middle Pliocene to Late Pleistocene large mammalian record of the Italian peninsula. 296 M.R. Palombo

found in a fossiliferous level at the top of a shallow- increase in standing richness during the subsequent water marine sequence of Late Pliocene age (BENVENUTI Pliocene and up to the beginning of the Early et al., 1995), calibrated with the Gauss/ Matuyama tran- Pleistocene. sition (LI N D S A Y et al. , 1980). The Montopoli FU should A temperate interlude occurred between 2.5 and correspond to the MN16b zone and was traditionally 1.8 Ma; during this phase the Mediterranean climate included in the Early Villafranchian LMA (Table 1), but was established permanently in southwestern Europe. the marked faunal turnover characterising the transition A progressive faunal renewal took place, and some dis- from the Triversa FU to the Montopoli FU recommends persal/origination bioevents led to a moderate faunal regarding it as the basal unit of the Middle structural reconstruction of late middle Villafranchian Villafranchian (Caloi & Palombo 1990, 1996). The faunal fauna. Two FU’s have been proposed as characterising renewal from Triversa to the Montopoli FU can be this time slice, the “Saint Vallier” and the Costa San regarded as a true turnover phase, due to the high per- Giacomo FU (Table 1), but the Italian “Saint Vallier” FU centage of last and new appearances, but also to the is loosely defined as being based on sporadic finds of important structural changes in the faunal complexes, taxa that also occur in the LFA’s assigned to the Costa in both the herbivore and carnivore guilds. This is con- San Giacomo FU. Accordingly, the author is inclined to sistent with the significant global changes characteris- consider the Italian late middle Villafranchian LFA’S as ing the Piacentian to Gelasian transition (actually the belonging to only one FU. The “Costa San Giacomo” beginning of the Quaternary as proposed by several name is here retained despite the priority of “S a i n t authors, see e.g. BOWEN & GIBBARD, 2007), with the per- Vallier”. This is because of the rich fossil record of the manent ice-sheet growth in the Northern Hemisphere, corresponding LFA (Costa San Giacomo, Anagni, the first major influx of ice-rafted debris in the middle Latium), as well to stress the significance of the appear- latitudes of the North Atlantic, the onset of extensive ance on the Italian peninsula of a wolf-like canid closely loess deposition in China around the Gauss/Matuyama related to Canis etruscus (a primitive representative of boundary (marine isotopic stage, MIS, 104 to 100) (see this phyletic lineage might be already present in the inter alios RU D D I M A N et al., 1989; DI N G et al., 1997; early Villafranchian Vialette LFA, LACOMBAT et al., 2008). SHACKLETON, 1997; PARTRIDGE, 1997a, b; RIO et al. 1998; As here redefined, the Italian Costa San Giacomo FU BO W E N & GI B B A R D , 2007 and recently ratified by the should roughly correspond to the MN 17”zone”. The International Commission on Stratigraphy, GI B B A R D & faunal composition is similar to that of the preceding HE A D , 2009), the profound changes in Eurasian flora unit as regards the presence of taxa such as assemblages (see inter alios ZA G W I J N , 1974; GR I C H U K , “Pseudodama” lyra, Eucladoceros tegulensis, Gazella 1997; SU C et al., 1997). Bioevents behind this faunal borbonica, or long-surviving taxa such as the cheetah, change (e.g. the so-called “elephant-E q u u s e v e n t ” , Acinonyx pardinensis, the hunting hyaena Lindsay et al., 1980) likely occurred close to the Chasmaporthetes lunensis and the proboscidean A . Piacentian- Gelasian boundary, driven by climatic cool- a r v e r n e n s i s , but renewed because of the first appear- ing, as well as by the effects of changes in the Earth cli- ance, among others, of the middle-sized horse E q u u s mate system: 19-23 ka cycles were superseded pro- stenonis, the goat Gallogoral meneghini, and the spiral gressively by a 41-ka rhythm (orbital obliquity periodici- horned antelope Gazellospira torticornis, all parkland ty) accompanied by moderately increased amplitude dwellers, as well the large-sized and slender legged climatic oscillations. The resulting increase in aridity boar Sus strozzii. and more intense seasonality triggered vegetation to The presence in the Costa San Giacomo of Canis reconstruct (SUC et al., 1995, and references in Bertini, aff. C etruscus bears witness that the so-called “wolf- 2003) and caused the disappearance of most forest- event” (AZ Z A R O L I 1983, 1995) was already in progress. dwelling taxa, especially small carnivores and arboreal- The carnivore guild renewal was a gradual phenomenon scansorial taxa, whereas new large grazers (the horse (SARDELLA & PALOMBO 2007, PALOMBO et al., 2008), which Equus livenzovensis), mixed feeders (a primitive developed throughout the latest Gelasian and the earli- Mammuthus meridionalis, the medium-size Gazella bor- est Calabrian and involved several large and middle- b o n i c a) or even browsers (the deer Eucladoceros fal- sized carnivores, such as the powerful scavenger, coneri) appeared (Table 3). Pachycrocuta brevirostris, the jaguar-like P a n t h e r a The taxonomical composition of the Montopoli FU gombaszoegensis (= Panthera onca toscana according resulted primarily from the dispersal, mostly from to HEMMER et all., 2003), the cooperative foraging canid, Eastern Europe, of large to medium-sized herbivores, Lycaon (Xenocyon) falconeri, as well the coyote-like while evolutionary substitutions within surviving phyletic (but see OL I V E , 2006) Canis arnensis (PA L O M B O et al. , lineages were rather negligible. The faunal change indi- 2008 and references therein). cates that forests gradually gave way to more open environments (including A r t e m i s i a steppe), alternating 3.3 The Late Villafranchian and the Gelasian- with warm temperate deciduous forests (see Bertini, Calabrian boundary 2003). Indeed, from an ecological point of view, the The latest Gelasian faunal renewal involved both Triversa faunal complex, which had a relatively high fre- carnivores and herbivores: most Pliocene and Early quency of frugivores, developed in environments rather Gelasian species disappeared, particularly among her- similar to those of modern forests, whereas the struc- bivores, and new carnivores and herbivores progres- ture of the Montopoli faunal complex shows some sively appeared (Table 3). The renewal from the middle affinities with those of modern bush-woodland to late Villafranchian LMA, should be regarded as a dis- (P A L O M B O & GI O V I N A Z Z O , 2006, P A L O M B O 2 0 0 7 b ) . persal phase (more than an actual turnover), which Moreover, this event can be considered as the begin- developed before and after the Gelasian-Calabrian ning of a dispersal phase leading to a progressive boundary, and came to an end during the earliest Early Biochronology of terrestrial mammals and quaternary subdivisions: ... 297 298 M.R. Palombo

Calabrian, (PA L O M B O , 2004b, 2007a, b). The dispersal 3.4 The end of the Villafranchian LMA and the dawn phase primarily involved carnivores, since among new of the Galerian LMA taxa none belong to species which evolved in loco, During the subsequent Early Pleistocene (Farneta, whereas new appearances among herbivores were Pirro and Colle Curti FU’s) (Table 2) an important principally linked to the emergence of new species renewal of faunas occurred: most of the typical within pre-existing phyletic lineages (e.g. M a m m u t h u s , Villafranchian taxa disappeared or became rare while Equus, Eucladoceros, “Pseudodama” and large bovids scattered bioevents led to the appearance of new of subgenus Leptobos), and subordinately, to immigra- species, either newcomers or ones evolved from pre- tion of large ruminants, mainly mixed feeders (e.g. existing taxa (Table 3). The previously mentioned Procamptoceras and Praevibos). These faunal changes increase in diversity throughout the early late indicate that forests or woodlands gradually gave way Villafranchian (mainly via migrations and dispersal to more open environments (including A r t e m i s i a events) had possibly altered the palaeocommunity steppe), but alternating with warm-temperate decidu- equilibria, and climate changes possibly removed key- ous forests. The gradual faunal reconstruction from the stone species causing new intra- and inter-guild latest Gelasian to Early Calabrian took form within an dynamics, leading mammalian communities to signifi- interval of climatic transition, and was possibly driven cantly reconstruct (PALOMBO, 2007b). This is the reason both by climatic worsening (moderate average cooling why, despite the progressive appearance of new MIS 70 to58), and modification of internal competitive species (most of which will survive in the early Middle dynamics, the latter depending on the disappearances Pleistocene), last appearances prevailed, leading to a of some pre-existing key taxa and ensuing availability progressive reduction in diversity from Farneta to Colle of ecological niches (PALOMBO, 2007b). Curti FU’s (Table 3). After AZ Z A R O L I (1977) the late Villafranchian, at this time considered as coincident with the early Pleistocene 3.4.1 Farneta and Pirro FU’s (s e n s u AG U I R R E & PA S I N I , 1985), would include three FU’s: Differences in taxonomical composition and struc- Olivola, T and F (Table 1). Subsequently, a more recent, ture between Tasso and the following Farneta FU are fourth FU, Pirro was added, and the end of the late mainly related to changes in the herbivore guild and Villafranchian was dropped before the Jaramillo sub- mirror the spread of open environments due to the chron (Table 1). LFA’s assigned to the Olivola and Tasso decrease in average temperature, along with the FU’s are well-known from the Upper Valdarno Basin, increase in dryness (BE R T I N I , 2003; SU C & PO P E S C U , where fossil-bearing sediments of two stratigraphically 2005). For instance, rhinoceroses similar to superimposed units are exposed (DE GI U L I & MA S I N I , Stephanorhinus hundsheimensis were the possible eco- 1 9 8 7 ; TO R R E , 1987; AZ Z A R O L I et al., 1988). Magnetostrati- graphy of these fossiliferous stratigraphical successions logical substitutes for Stephanorhinus etruscus. T h e provides chronological constraints for Olivola plus the larger, mixed-feeder Praemegaceros obscurus replaced Tasso FU: in the Upper Valdarno area, LFA’s assigned to the browser Eucladoceros, and a more specialised form the Tasso FU have been correlated to the top of the replaced the mixed-feeder “Pseudodama” nesti. It is Olduvai subchron (NA P O L E O N E et al., 2003), which is worth noting that BR E D A & MA R C H E T T I (2007) doubtfully around the GSSP of the Gelasian-Calabrian boundary reported ?Eucladoceros ex gr. ctenoides-dicranios and (La Vrica Section; AG U I R R E & PA S I N I , 1985), whereas ?Megaloceros obscurus from the main brown coal level LFA’s of the Olivola FU (e.g. Matassino and Poggio (subunit 5) of the biogenetic unit of Leffe Formation Rosso LFA’s) are “inferred as pre-Olduvaian, not (Bergamo, Northern Italy), but the actual co-occurrence younger than 2.0 Ma.” (NA P O L E O N E et al. , 2003, pag. 308). of these large deer needs to be fully substantiated. The tructure of the faunal complexes of the Among carnivores, Megantereon whitei, a saber-tooth Olivola and Tasso FU’s are not very dissimilar, the latter cat of African origin, first appeared. being different essentially for the appearance, among The faunal renewal between the Farneta and Pirro others, of Lycaon (Xenocyon) falconeri (already present FU (Table 2) is marked by the appearance of some in the latest Pliocene Spanish fauna of Fonelas P-1, arriving African taxa, such as the large H i p p o p o t a m u s GA R R I D O , 2008), Equus stehlini (= Equus senezensis antiquus (MAGRI et al., 2009), and (doubtfully) the cerco- stehlini, according to ALBERDI et al., 1998) and the stout pithecid T h e r o p i t h e c u s, a mixed-feeding genus whose bovid Leptobos vallisarni. Praeovibos is also recorded extant species inhabits the high grasslands of Ethiopia in the Casa Frata LFA, whereas Gallogoral, Gazellospira and Eritrea (RO O K et al., 2004, but see PA T E L et al., a n d Procamptoceras were not reported (Table 3). The 2007). This evidence of African immigration to south- occurrence of Hippopotamus antiquus in the Tasso FU ernmost Europe stresses the importance of “out of is doubtful, since hippopotamuses from Valdarno Africa” migratory waves, which occurred approximately (already ascribed to the Tasso FU), was probably around 1.6/1.3 Ma (MA R T I N E Z - N A V A R R O , 2004). Other retrieved from fossiliferous layers younger than those of immigrants had an Asian/European origin, as the ovi- the Montervarchi succession (NA P O L E O N E et al., 2003). bovine M e g a l o v i s and the rather stout bovine B i s o n On the other hand, the occurrence in the same strati- (Eobison) degiulii, the latter replacing the slender lepto- graphical layer of H. antiquus and Leptobos aff. furtivus bovines. Among perissodactyls, the middle-sized, slen- at Piano dei Cavalieri (Rieti basin, Latium) (MASINI 1989, der limbed Equus altidens and the larger E q u u s GE N T I L I & MA S I N I , 2005), needs to be confirmed (see s u e s s e n b o r n e n s i s were certainly present, whereas SE G R E NA L D I N I & VA L L I and references therein). Equus stehlini is no longer recorded (see AL B E R D I & Accordingly, faunas assigned to OL I V O L A and TA S S O PALOMBO, in press for a discussion). Among carnivores FU’s seem to be part of the same, progressive faunal the middle-sized bone crusher Canis mosbachensis renewal, and the hypothesis that they could represent a first appeared and Lycaon (Xenocyon) lycaonoides sub- single FU cannot be ruled out. stituted the less advanced L. falconeri. The dispersal of Biochronology of terrestrial mammals and quaternary subdivisions: ... 299

Homo in the Italian peninsula is testified by the finding (BO N I F A Y , 1978; AZ Z A R O L I et al., 1988) or a distinct of lithic implements in the karstic network of Pirro Nord, biochron (e.g. Protogalerian sensu Caloi & Palombo, the local eponym of the Pirro FU (AR Z A R E L L O et al. , 1995, Epivillafranchian sensu Kahlke 2007 and previous 2007). All in all, forest-dwelling taxa dwindled, whereas papers; ‘latest Villafranchian’ sensu Koufos, 2001 or species inhabiting prairies, savanna or steppes ‘Final Villafranchian’ SPASSOV, 2003). On the other hand, increased in percentage. The prevalence of dry and if Middle Pleistocene faunas are clearly to be consid- open environments at the time of the Pirro FU (and ered as ‘modern’ when compared to those from the Colle Curti FU as well) is confirmed by the great struc- Villafranchian (PA L O M B O , 2004b), which criteria should tural compatibility of these faunas with those of present be used to define the ‘theoretical’ boundary between day savanna/bushland habitats (PA L O M B O , 2007b). the Villafranchian and the following Galerian LMA? Nevertheless, forest areas were still rather abundant, According to GL I O Z Z I et al. (1997) (Table 1), the begin- and woodlands spread in the more temperate and ning of the Galerian LMA would be conventionally humid climatic phases, as evidenced, for instance, by marked by the appearance, among others, of the presence of the browser moose Cervalces carnutu- Praemegaceros verticornis, which is first recorded in r u m in the subunit 7 of Leffe Formation (Bergamo, Italy at the Colle Curti LFA (Colle Curti FU, late Early Northern Italy) (BREDA & MARCHETTI, 2005, 2007), which Pleistocene, Jaramillo Subchron, COLTORTI et al., 1998). deposited shortly before the Jaramillo Subchron This giant deer has been recently found in the Spanish (MUTTONI et al., 2007). Fuente Nueva 3 and the Barranco Leon LFA, dated at The occurrence at Pietrafitta (Umbria, central Italy) about 1.3 Ma (AG U S T Í & MA D U R E L L, 2003), which retain of a vole close to Microtus (Allophaiomys) ex gr. M. ruf- several Villafranchian taxa, as does the Colle Curti LFA. foi, and of Microtus (Allophaiomys) chalinei a n d Moreover, the Pirro Nord (latest Villafranchian) and the Mimomys pusillus in a large mammal fauna whose taxo- Colle Curti LFA’s exhibit a relatively high coefficient of nomical composition and structure is consistent with similarity (PA L O M B O & VA L L I , 2005). Consequently, the that of the the Farneta FU (GE N T I L I et al., 1996; BA R I S O N E Colle Curti LFA still looks closer to the latest et al. , 2006; PA L O M B O & GI O V I N A Z Z O , 2006), as well the Villafranchian LFA’s than to the ensuing typical Galerian presence at Pirro Nord of a more evolved M i c r o t u s ones, despite the LlSDk of Praemegaceros verticornis, ( A l l o p h a i o m y s ) ex gr. M. ruffoi, provide a correlation of Cervalces latifrons (BREDA & MARCHETTI, 2005), and pos- the Farneta and Pirro FU with the earliest Biharian (Early sibly of subgenus B i s o n , the significant reduction in Biharian I, MA S I N I & SA L A, 2007). Accordingly, and taking biodiversity shown by carnivores, as well as the innova- into account magnetostratigraphical dating of Tuscan tive characteristics of the arvicolids (SA L A & MA S I N I , LFA’s assigned to Olivola plus the Tasso FU, the Farneta 2007). and Pirro FU’s developed during the pre-Jaramillo Early All in all, the the Pirro and Colle Curti FU’s corre- Pleistocene, but the actual chronological extent of each spond to a time slice during which the Villafranchian FU is based only on biochronological estimates. were gradually but not completely displaced by some typical Galerian large mammals foreshadowing the 3.4.2 Colle Curti FU and the beginning of the Mid coming of a new age. This evidence encourages a Pleistocene revolution (MPR) redefinition of the Galerian LMA (cf. PA L O M B O , 2004a The transition from the Early to Middle Pleistocene, and references therein, KAHLKE, 2007). Mid-Pleistocene revolution (MPR) (from about 1.2 to 0.6 Ma) marks a fundamental change in the Earth’s climate 3.5. The MPR accomplishment and the onset of typi- system (MA S L I N and RI D G W E L L , 2005) and represents a cal Galerian mammalian faunas major episode in mammalian fauna reorganisation over As above mentioned, the faunal renewal leading to the course of the Cenozoic. Orbital obliquity at 41-ka the typical Galerian faunas finds its origin in two distinct cycles was superseded progressively by a 100-125-ka trends: 1) the progressive reduction in richness charac- rhythm, sustained by four-five precessional cycles, and terising late Early Pleistocene faunas (from the Farneta accompanied by increased amplitude climatic oscilla- to the Colle Curti FU’s), 2) the subsequent dispersal tions (Table 2). Glacial phenomena intensified, tempera- and the progressive spread in Italy of taxa from Eastern ture and moisture dropped notably, vegetational “exot- and Central Europe, that triggered a rapid increase in ic” essences, quite common in earlier floras, progres- diversity at the beginning of the Middle Pleistocene sively disappeared and sustained climatic changes pre- (Slivia FU) (PALOMBO 2007b and references therein). It is vented these floral elements from becoming re-estab- noteworthy that this faunal change involved both carni- lished (BERTINI, 2003). These changes spurred a shift in vore and herbivore guilds at various times. The most environments and constrained the geographical diffu- important change in the former had already taken place sion of a number of taxa, allowing dispersal waves that prior to the Jaramillo Subchron, at the Pirro–Colle Curti drove the reorganisation of large mammal communities. FU’s transition, with the disappearance of some large Community rebuilding came to an end during the taxa, either flesh-eaters or bone-breakers, such as beginning of the Middle Pleistocene with the disappear- large canids and felids (Table 3) (PALOMBO et al., 2008 ance of the last surviving Villafranchian species, such and references therein). This is the starting point for a as Pachycrocuta brevirostris and Panthera gombazsoe- reversal in the proportion of carnivores versus herbi- gensis (Table 3). Thus, at the time of MPR, fauna was vores: indeed, across the Early to Middle Pleistocene characterised by the presence of long surviving (Slivia FU), the faunal renewal was strongly biased in Villafranchian taxa and by a progressive increase in favour of large herbivores, while the carnivore guild taxa, which also persisted into the Middle Pleistocene. changed at a slower pace. Taxa dispersed from Eastern Such a framework of “mixed faunal elements” was and Central Europe reached the Italian peninsula in regarded by authors either as a “transitional fauna” succession, thus leading an important and relatively 300 M.R. Palombo

rapid increase in diversity at the beginning of the authors regarded as intermediate in taxonomical com- Middle Pleistocene. These changing faunas represent a position, and in age as well, between the Slivia and major large mammal community reorganisation Isernia FU’s. This, because of the persistence of throughout the Quaternary. This dispersion phase was Pachycrocuta brevirostris and Panthera gombaszoe- probably triggered by notable climatic changes that g e n s i s in the Slivia FU, the exclusive occurrence of correspond to the onset of the pronounced Hemibos galerianus in the “Ponte Galeria FU” and the glacial–interglacial fluctuations as observed in the appearance of Bos primigenius in the Isernia FU. marine isotope record during the late Early Pleistocene Nevertheless, Slivia and Ponte Galeria 2 LFA’s would (MIS 25 to 22-20), while the progressive increase in be assumed to be part of the same FU due to the diversity is seen as a consequence of the ensuing onset uncertain identification of some herbivores from Slivia of wetter climatic conditions (see P A L O M B O & LFA, and the assumption that carnivores absent from GI O V I N A Z Z O , 2006 and references therein). Instead of the Ponte Galeria CU have in fact been found in the replacing any one species, these taxa enriched the her- Slivia LFA. Unfortunately, there are no stratigraphical bivore guild, as well as the large mammal community constraints for the age of the Slivia FU, even if it can be as a whole (Palombo, 2007b and references therein). correlated with the Late Biharian, which roughly Even if, on a larger temporal scale, the progressive extends from 0.95 to 0,6 Ma, due to the occurrence of modification in the structure of large mammal faunas of M. savini associated with that of Microtus (Steno- the Italian peninsula began before the Jaramillo c r a n i u s) and Microtus (Terricola). Conversely, in the Subchron, it was shortly before the Early to Middle Roman basin, the integrated mammal biostratigraphy Pleistocene boundary that the major renewal and and sequence stratigraphy approach represents a valid important ecological reorganisation of faunal complex- tool to identify and to correlate physical and biological es took place (PALOMBO, 2007b). events. The biochronological and sequence-strati- Throughout the course of time (Table 3), the local graphical scheme proposed for the Ponte Galeria area, appearances of several pachyderms (E l e p h a s highlights that the sequence stratigraphical framework (Palaeoloxodon) antiquus, Mammuthus trogontherii, can constrain the possible chronological interval in Stephanorhinus kirchbergensis, Stephanorhinus hemi- which a bioevent, even if local, occurred (see MI L L I & t o e c h u s), large (Megaloceros savini, Praemegaceros PALOMBO, 2005). solilhacus), and medium-sized deer (Capreolus capreo- The transition from the middle to late Galerian lus suessenbornensis, Cervus elaphus acoronatus a n d (Fontana Ranuccio FU) possibly occurred around possibly Dama clactoniana), as well as large (Hemibos 550–500 ka, due to the absolute age available for the galerianus, Bos primigenius) and middle-sized bovids LFA’s ascribed to Isernia FU (Notarchirico and Isernia (Ovis ammon antiqua, Hemitragus bonali), together with LFA’s, estimated age about 600 ka by LE F É V R E & caballine horses (Equus mosbachensis), all mainly RAYNAL, 1999; Lefévre et al., 1999; and COLTORTI et al., mixed feeders or grazers, inhabiting open landscapes, 2005, respectively), and the Fontana Ranuccio FU bushlands or wooded grasslands, brought about a (Fontana Ranuccio LFA, estimated age about 458 ka, major change in the herbivore guilds (PALOMBO, 2007b). BIDDITTU et al., 1979). Among carnivores, Crocuta crocuta first appeared During the late middle Galerian (Fontana (Ponte Galeria 2 LFA, PG2 sequence, about 750-650ka, Ranuccio FU), “Villafranchian” taxa were not yet record- MILLI & PALOMBO, 2005 and references there in), where- ed and the number of very large herbivores diminished as some other large carnivores, active hunters that (Table 3). The occurrence of “Galerian” equids and were both flesh-eaters and scavengers [such as Ursus megacerines is doubtful and the specialised and slight- deningeri, Panthera fossilis, Isernia LFA, about 600 ka ly larger H. antiquus was replaced by H i p p o p o t a m u s (COLTORTI et al., 2005); a hyaena, Hyaena prisca, whose a m p h i b i u s and, among carnivores, Ursus arctos taxonomical identification is still matter of debate at the appeared as well as Felis silvestris (Table 3). Ponte Galeria 3 LFA (see CA L O I & PA L O M B O , 1980; In the second half of the Middle Pleistocene, PA L O M B O et al., 2008; TU R N E R et al., 2008), PG3 about increasingly different habitats supported increasingly 650-550 ka ( M I L L I & PA L O M B O , 2005); Panthera pardus, varied faunas (cf. TR I A N T I S et al., 2003), with new Valdemino Cave LFA, Isernia FU (SA L A , 1992) and dynamic interspecific relationships. In fact, with the end Isernia LFA (SALA, 2006) ], had their lowest stratigraphi- of the Galerian (Fontana Ranuccio FU) and the transi- cal occurrence even later. These species, together with tion to the following Aurelian LMA, the most flexible S. hundsheimensis, Equus altidens, E. suessenbornen- species became the dominant group, the percentage of sis, P. verticornis and B. shoetensacki are eponymous browsers increased and fauna progressively acquired of the “typical” Galerian faunas. modern characteristics (PALOMBO 2007b and references Most authors recognise in the “middle” Galerian therein). (sensu Gliozzi et al., 1997) two FU’s, Slivia and Isernia which are clearly differentiated by the small mammals: Mymomys savini (Slivia LFA) and Arvicola mosbachen- 3.6 The Aurelian LMA and the Middle to late sis (Isernia LFA). The small mammal association found Pleistocene boundary at the archaeological site Isernia La Pineta (Molise), GLIOZZI et al. (1997) proposed the Aurelian as a new represents the oldest Toringian fauna recorded in Italy LMA, differing from the Galerian LMA because of the to date (SALA & MASINI, 2007) and represents the main appearance of taxa that constitute the core of the mod- renewal of small mammals throughout the Galerian ern mammalian fauna, whereas some “Galerian”species LMA. (such as horses, megacerine deer, m e d i u m - s i z e d , PE T R O N I O and SA R D E L L A (1999) proposed the intro- bovid Ovis ammon antiqua and Hemitragus bonali, and duction of the Ponte Galeria FU (Table 1), by these large bovide, Bison shoetensacki) were no longer pre- Biochronology of terrestrial mammals and quaternary subdivisions: ... 301

sent. According to GL I O Z Z I et al. (1997), three distinct, thus the Vitinia FU cannot be considered a valid main faunal complexes could be detected: early, middle biochron. The LFA’s previously ascribed the Torre in and late Aurelian, characterised by an increase in the Pietra and the Vitinia FU’s have to be included in a sin- percentage of modern taxa. The early and middle gle FU (named Torre in Pietra FU for priority reason) Aurelian were subdivided on the basis of faunal differ- (Table 1). ences between two already created FU’s, Torre in Summarizing, several bioevents characterised the Pietra and Vitinia (CA L O I and PA L O M B O 1990, 1996, but beginning of the Torre in Pietra FU, as well as the see below), while for the late Aurelian (MIS 5-2) no char- beginning of the Aurelian LMA (subdivided into early acteristic FU’s have correctly been designated due to and late Aurelian) as here redefined: for instance Canis the monotonous taxonomical composition of the last lupus first appeared as well as Ursus spelaeus that sub- interglacial faunas, as well the only sporadic occur- s t i t u t e d U. deningeri, Megaloceros giganteus, Dama rence of the so called “cold taxa” during the last Glacial dama “tiberina, and Bubalus murrensis (PALOMBO, 2007 (but see PETRONIO et al, 2007). and unpublished data), the “steppe horse E. hydruntinus dispersed along the Italian peninsula. Furthermore, 3.6.1 The Aurelian LMA and the question of Vitinia some new medium-sized taxa appeared, when more FU severe, arid climatic oscillations favoured the spread of The Torre in Pietra and Vitinia FU’s have been “mountain” bovids (Capra ibex and Rupicapra) towards defined on the basis of palaentological and stratigraphi- the southernmost regions of the Italian peninsula, cal evidence collected in the Roman Basin (CALOI et al., whereas Galerian megacerine deer and horses, O v i s 1998; but cf. PALOMBO et al., 2004 and references there- ammon antiqua and Bison shoetensacki are no longer in). Along the Latium coast, rich LFA’s (Torre in Pietra, recorded (Table 3). La Polledrara di Cecanibbio, Collina Barbattini, Castel All in all, with the end of the Galerian (Fontana di Guido, Malagrotta) have been recovered either from Ranuccio FU) and the transition to the Aurelian (Torre in fossiliferous deposits of the Aurelia Formation (CONATO Pietra FU), the most flexible species became the domi- et al., 1980) or the PG6 sequence, both correlated with nant group, the percentage of browsers increased and MIS 9 (MILLI & PALOMBO, 2005). In these LFA’s, assigned fauna progressively acquired modern characteristics. to the Torre in Pietra FU the LlSDk of Canis lupus, This is consistent with a moderate woodland expansion Ursus spelaeus and Megaloceros giganteus is reported. under increasingly mild climatic conditions characteris- In the same area, LFA’s uncovered in fossiliferous lev- ing the interglacial phases, as well as the increase in els of the Vitinia Formation (CONATO et al., 1980) or PG7 north to south and east to west climatic gradients lead- sequence (MILLI, 1997), correlated with MIS 7 (e.g. Torre ing to the fragmentation of Italian habitats. in Pietra upper levels, Vitinia level e, Cerveteri) (PALOMBO The structure of Aurelian mammalian faunas was et al., 2003 and references therein) were previously fully attained long before the beginning of the beginning ascribed to the Vitinia FU’s (CA L O I & PA L O M B O , 1995; Late Pleistocene, thereby the transition from Galerian to CALOI et al., 1998) essentially because of the abundance Aurelian LMA predated the Middle to Late Pleistocene of a primitive fallow deer, Dama dama tiberina ( l o c a l l y boundary. associated with the pre-existing larger Dama dama clactoniana), as well the appearance, among others of 3.6.2 The Late Pleistocene Cervus elaphus with features similar to those of modern The Last Interglacial LFA’s did not substantially dif- red deer and of Equus hydruntinus. Nevertheless, the fer from those recorded during the late Middle structure and the taxonomical composition of the Torre Pleistocene. The rare species Bubalus murrensis is no in Pietra and Vitinia faunal complexes are similar longer present, while modern fallow and red deer sub- (PALOMBO & MUSSI, 2001; PALOMBO 2004), thus the divi- stitute the late Middle Pleistocene subspecies, but the sion into two FU’s seems to be based on weak palaen- core of the early Aurelian faunas keeps roughly tological data, moreover new stratigraphical evidences unchanged. Although the Last Glacial phase caused a call for a revision of these biochrons. In the 1990’s, two remarkable shift in the vegetational cover and in the LFA’s, Sedia del Diavolo and Monte delle Gioie (today, distribution of mammals in Europe, minor changes no longer exposed due to of the intense urbanization of characterised the transition from the Last Interglacial to Rome after World War II), were assigned to the Vitinia the Last Glacial faunal complexes on the Italian penin- FU due to the abundance of Dama dama tiberina sula (PALOMBO, 2007b). During the last Glacial, the fau- remains in the alluvial deposits overlaying the “Tufo nal biodiversity increased with the arrival of some lionato” (a lithified pyroclastic flow deposit from Albani “cold” species, such as the typical, Mammuthus primi- Volcanic Complex, dated 355 ka by KARNER et al., 2001) g e n i u s, the woolly rhinoceros Coelodonda antiquitatis, where also Equus hydruntinus was found (CA L O I & the elk Alces alces, inhabiting a “mammoth steppe”-like PA L O M B O , 1995). CA L O I et al., (1998) correlated these environment extending throughout the Eastern Po bone-bearing deposits with the Vitinia Formation (MIS Valley (GALLINI & SALA, 2001). Mammoth and woolly rhi- 7), but recently the overlain ash flow has been dated at noceros, as well the wolverine, Gulo gulo, reached the 285 ± 1 ka (MARRA & ROSA, 1995; KARNER et al., 2001). Tyrrhenian coast (Latium) and were widespread in the Following this new chronological data, the deposition of south-eastern part of the Italian peninsula, where the fluvio-lacustrine sediments overlaying the “Tufo Alopex lagopus is also recorded, while Rangifer taran- lionato” at Sedia del Diavolo, occurred approximately d u s only reached the westernmost geographical during the time span between 355 and 285 ka (PALOMBO boundary of the Italian coast ( C A L O I & PA L O M B O , 1995; et al., 2004). SA L A , 2005). The spread of “cold” species across the Accordingly, the LlSDk Dama dama tiberina in Italy Italian peninsula was profoundly conditioned by micro- is documented in deposits correlated with the MIS 9, climate and physiography, and the taxonomical compo- 302 M.R. Palombo

sition and structure of LFA’s could vary greatly, based on palaeobiological events which have a wide depending on the geographical position of fossiliferous territorial significance - could be useful for chronologi- sites. That, coupled with the high percentage of extant cal correlations. Nonetheless, the transition between species, prevent any FU to be fairly defined. Moreover, successive LMAs does not always correspond to the the faunal assemblages were progressively depleted of boundaries separating marine Series or Stages. For some long-surviving Galerian species. Pachyderms instance, the Villafranchian LMA approximately began were most affected by climatic deterioration. with the Late Pliocene (Piacentian) and the transition H i p p o p o t a m u s survived only until the end of the MIS from the early to the middle Villafranchian LMA hap- 5a, whereas Elephas antiquus occurred also during MIS pened around the Pliocene/Pleistocene (Piacentian/ 4 and S t e p h a n o r h i n u s was still present during MIS 3, Gelasian) boundary, whereas the transition to the late while Panthera spelaea survived till the beginning of the Villafranchian took place during the latest Gelasian, and Holocene (BOSCHIAN et al., 1995). those from the Villafranchian to Galerian LMA and from All in all, the mammal faunas on the Italian main- the Galerian to Aurelian LMA respectively predated the land maintained “temperate-cool” characteristics even beginning of the Middle and Late Pleistocene. during the cold-arid peaks of the Last Glacial. Broad- Concerning the Villafranchian to Galerian LMA transi- leaved wooded release “refugial” areas always existed tion, from what has been previously reported on the on the Tyrrhenian side of the peninsula, while the cli- biochronological patterns of large mammals, it possibly mate on the Adriatic side was probably drier. Therefore, occurred close to MIS 25, where CI T A & CA S T R A D O R I some “warm” species remained on the Italian peninsula (1995, but see CI T A et al. , 2006) previously proposed longer than elsewhere, and turnovers in successive lowering the Early to Middle Pleistocene Boundary. communities were less dramatic than in northern Europe. With the deglaciation phase at the beginning of the Holocene, dramatic moisture and vegetation ACKNOWLEGEMENT change led to marked changes in mammalian faunas, particularly in their abundance, leading to the megafau- Thanks are due to the Organising Committee of na extinction. the AIQUA Congress “Il Quaternario nella cartografia CARG2 for the opportunity to present this paper and to the anonymous reviewers for their helpful comments on 4. CONCLUSION the manuscript. The English version was revised by Steven T. Haire Several evidences show that, at least as far as the Work supported by the project MIUR Università Italian peninsula is concerned, global climatic changes 2007 C26A07LPWJ. are a contributing factor, especially in the first appearances of large mammals. This permitted the dispersal of new taxa, because mammals frequently reacted to the Plio-Pleistocene climate changes, expanding their REFERENCES distribution area or migrating rather than by evolving and producing new species. 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