Quaternaire
Revue de l'Association française pour l'étude du
Quaternaire
vol. 18/1 | 2007
Q5 Le Quaternaire, Limites et spécificités - Première
partie
The Pliocene-Pleistocene boundary: which
significance for the so called “Wolf Event”?
Evidences from Western Europe
Quelle importance a le « Wolf event » pour la définition de la limite entre
Pliocène et Pléistocène ? Les évidences données par les faunes de l’Europe sud
occidentale
Raffaele Sardella and Maria Rita Palombo
Publisher
Association française pour l’étude du
quaternaire
Electronic version
URL: http://quaternaire.revues.org/969
DOI: 10.4000/quaternaire.969
ISSN: 1965-0795
Printed version
Date of publication: 1 mars 2007
Number of pages: 65-71
ISSN: 1142-2904
Electronic reference
Raffaele Sardella and Maria Rita Palombo, « The Pliocene-Pleistocene boundary: which significance
for the so called “Wolf Event”? Evidences from Western Europe », Quaternaire [Online], vol. 18/1 | 2007,
Online since 01 March 2010, connection on 02 October 2016. URL : http://quaternaire.revues.org/969
; DOI : 10.4000/quaternaire.969
The text is a facsimile of the print edition.
© Tous droits réservés
Quaternaire, 18, (1), 2007, p. 65-71
THE PLIOCENE-PLEISTOCENE BOUNDARY:
WHICH SIGNIFICANCE FOR THE SO CALLED “WOLF EVENT”?
EVIDENCES FROM WESTERN EUROPE
䡲
1
Raffaele SARDELLA & Maria Rita PALOMBO
1
ABSTRACT
The radiation of canids referable to Canis ex gr. etruscus in Western Europe, the so-called “Wolf Event” by Azzaroli (1983), is an important
part of a faunal renewal that occurred approximately at Pliocene-Pleistocene transition, strongly related to climatic and environmental changes.
Actually, taking into account the recently available data, the meaning of the so called “Wolf-Event” must be reconsidered in a wider sense,
and it has to be considered as different dispersal events occurred in a quite short time span to be placed at the end of Pliocene. The reconsideration of
the available fossil record suggests that:
1) The “Wolf Event”, in its original acception, is just one of the different bioevents (dispersal of Canis, Lycaon, but also Pachycrocuta and
Panthera) part of the important faunal renewal that affected the carnivore guilds in Western Europe;
2) New hunting strategies (patch hunters) took place in Western Europe at this moment;
3) The dispersal of different lineages of canids has to be related to the diffusion of open habitats;
4) The “Wolf Event”, in a new and wider acception, occurred during latest Pliocene, thus it cannot be considered as a “marker” of the beginning of Pleistocene.
Key words: Canids, Carnivora, Richness, Turnovers, Pliocene, Pleistocene, North Western Mediterranean.
RÉSUMÉ
QUELLE IMPORTANCE A LE « WOLF EVENT » POUR LA DÉFINITION DE LA LIMITE ENTRE PLIOCÈNE ET PLÉISTOCÈNE ? LES
ÉVIDENCES DONNÉES PAR LES FAUNES DE L’EUROPE SUD OCCIDENTALE
La diffusion des espèces du genre Canis est un « bioevent » dont l’importance a été reconnue depuis longtemps. L’apparition de Canis etruscus (« Wolf Event ») avait été vue comme le signal du début du Pléistocène. Canis etruscus a été reconnu pour la première fois dans la faune locale
de Olivola (Valdarno supérieur, Italie) dont l’âge était considéré comme Pléistocène inférieur.
Des découvertes plus récentes nous obligent à placer le « Wolf Event » dans le Pliocène: Canis cf. C. etruscus est présent en Italie à Costa S.
Giacomo (MN17) et les trois canidés Villafranchiens (C. etruscus, Canis arnensis et Lycaon falconeri) sont déjà présents dans le Pliocène final à
Fonelas. Par ailleurs, dans le Bassin du Valdarno, C. etruscus et C. arnensis ont été trouvés en association avec Chasmaporthethes lunensis et
Pachycrocuta brevirostris dans des dépôts d’âge Pliocène final, corrélés avec l’événement paléomagnétique d’Olduvai.
Le « Wolf Event » va donc se produire en Méditerranée Nord occidentale au cours du Pliocène supérieur, bien que de façon différente dans
chaque région, comme le montre l’observation des richesses spécifiques et des taux d’apparition et d’extinction. De plus, ce bio-évènement n’inclut
pas seulement la dispersion de Canis et Lycaon, mais également de Pachycrocuta et, au moins pour l’Italie, de Panthera, produisant un renouvellement important de la guilde des carnivores en Europe occidentale.
En conclusion, puisque ce renouvellement va se développer en grande partie durant le Pliocène final, le « Wolf Event », dans sa nouvelle
acception plus étendue, ne peut pas être considérer comme un “marqueur” du début du Pléistocène.
Mots clés : Canidés, Carnivores, Richesse spécifique, renouvellement faunique, Pliocène, Pléistocène, Méditerranée Nord occidentale.
1 - INTRODUCTION
Since the Late Pliocene, a serie of climatic cycles
have caused latitudinal displacements of vegetational
cover and biomes in Europe. Terrestrial mammals reacted to such climate changes by expanding their diffusion area or migrating. The radiation of canids
referable to Canis ex gr. C. etruscus in Western Europe,
the so-called “Wolf Event” (WE) by Azzaroli (1983), is
an important part of a faunal renewal that occurred approximately at Pliocene-Pleistocene transition. The
term “WE” initially defined the arrival in Western Europe of the first representative of the genus Canis
“sensu stricto” and the beginning of the late
Villafranchian (Olivola Faunal Unit), at the time regarded as roughly coincident with the beginning of the
Early Pleistocene (as formally defined by Aguirre &
Pasini, 1985; see Pillans 2004 and Clague 2006 for a
1
Dipartimento di Scienze della Terra, Università “La Sapienza, CNR - Istituto di Geologia Ambientale e Geoingegneria, Piazzale A. Moro,
5 - 00185 ROMA, Italy. E-mail: [email protected]
Manuscrit reçu le 08/07/2006, accepté le 09/10/2006
66
discussion). In this paper a concise overview of the
most recent discoveries is presented together with a reconsideration of the biochronological value of important bioevents as suggested by the analyses of the
Pliocene and Pleistocene carnivore guilds in the Mediterranean region (Palombo, in press; Palombo et al., in
press).
2 - CANIS DISPERSAL IN THE WESTERN
EUROPE AND RELATED ISSUES
The origin of the genus Canis is matter of debate.
Many authors agree in suggesting an Asian origin for
the genus. An Early Pliocene occurrence of Canis in
Europe has to be considered with caution, being present in coeval deposits taxa very similar in morphology
and size to true dogs, but that can be referred to the genus Eucyon (see Spassov & Rook, 2006 for a discussion).
Actually, wolf-like canids dispersal bioevent occurred during the Late Pliocene when such animals, already reported in North China around 3.0 Ma (Mazegu
Formation), dispersed from Asia into Europe (Flynn et
al., 1991). Turner (1995) pointed out that the genus
Canis, represented by several species of jackals, first
appeared in Africa approximately at 2.5 Ma, after
dispersion from Eurasia.
Recent discoveries in central Italy (Anagni and
Valdarno Basins) provide data for updating the
biochronological framework of such bio-event.
At Costa S. Giacomo, Canis cf. C. etruscus has been
retrieved from a faunal assemblage including several
middle Villafranchian taxa (Cassoli & Segre Naldini,
1984, 1993). Size and morphology of those specimens
(a fragmentary right lower carnassial and two premolars) suggest close relationships to the Villafranchian
wolf-like canids Canis ex gr. C. etruscus (Rook &
Torre, 1996). On one hand, new magnetostratigraphic
calibrations have demonstrated a Late Pliocene age for
faunas belonging to Olivola FU such as the “late
Villafranchian” Poggio Rosso local fauna (LFA), in
which Canis arnensis marks its lower occurrence together with Pachycrocuta brevirostris, but where
Chasmaporthetes lunensis is still present (Mazza et al.,
2004). Poggio Rosso is one of the richest and most interesting sites in the Upper Valdarno basin, whose
latest Pliocene age has been proved by
magnetostratigraphical evidences (upper part of the
Olduvai chron; Napoleone et al., 2001, 2003). Taking
into account the recently available data the so called
“WE” has to be considered as different dispersal events
occurred in a quite short time span to be placed at the
end of Pliocene.
In the early late Villafranchian faunal assemblages
(Olivola + Tasso FUs, sensu Palombo, 2005) related to
the end of the Olduvai paleomagnetic event, three
canids have been recorded: the wolf-like Canis
etruscus, the smaller coyote-like dog Canis arnensis,
and the large sized hypercarnivorous wild dog Lycaon
falconeri. The large sized Lycaon falconeri characterised the late Villafranchian assemblages referred to
Olivola and Tasso FUs, being replaced by the more
derived Lycaon lycaonoides, recorded in Italy at
Villafranchian-Galerian transition (Pirro & Colle Curti
FUs). As stated by Werdelin & Lewis (2005) such a
phyletic trend seems plausible, although it must be
noted that “leaving Lycaon with full generic status renders Canis paraphyletic” and for this reason they prefer
to refer these species, as well as the ancestral species of
the lineage “Lycaon” falconeri, to Canis. Only the revision of the genus Canis will enable the researchers to
clarify such a topic.
In France, some dental remains belonging to a medium-sized dog from Senèze had been already referred
to Canis cf. C. arnensis by Schaub (1943). 30 years
later, Martin (1973) referred these fossils to the new
species Canis senesensis because of its primitive tooth
features. Fossil remains showing great affinities with
this species have been recorded even in the late
Pliocene mammal faunas from the early middle
Villafranchian Liventsovka and Khapry LFAs (Southern Russia) (Sotnikova et al., 2002). At Liventsovka
this canid has been recorded together with the two
hyaenas Pachycrocuta brevirostris and Pliocrocuta
perrieri, while at Khapry Canis cf. C. senezensis coexists with P. perrieri. Thus the dog from Senèze could
be considered as a primitive step of the Canis arnensis
lineage (Palombo, in press).
Also recent discoveries in Spain puts in evidence the
early appearance of the three Villafranchian canids in
Europe. A rich vertebrate fauna with “middle
Villafranchian taxa”, such as Gazella borbonica, comes from Fonelas P1 (Southern Spain) (Arribas et al.,
2004). In this site Canis aff. C. etruscus, Canis aff.
C. arnensis and Lycaon falconeri have been recorded
together with three different hyaenids such as
Pachycrocuta brevirostris, Chasmaporthetes lunensis
and Parahyaena brunnea. The occurrence of a primitive form of the living brown hyaena, perhaps a synonym of Pliocrocuta perrieri, is currently an open issue.
The appearance of different canids in Europe during
the late Pliocene is testified by an increasing number of
fossil evidences and it is worth nothing that not only
Canis etruscus but also the primitive wild dog could be
appeared in correspondence of the middle
Villafranchian faunas (MN17). Indeed at Puebla de
Valverde, some dental remains belonging to a large
canid have been referred to Canis cf. C. falconeri
(Kurtèn & Crusafont Pairò, 1977). Thus, evidences for
an early occurrence in the Late Pliocene of Europe of a
primitive wild dog are becoming more and more significant and numerous even if problematic as well as their
phylogeography.
It is worth noting that in Fonelas P1 LFA Panthera
gombazsoegensis has not been recorded yet. Actually,
the origin and dispersal of the Eurasian jaguar
Panthera gombaszoegensis is still unclear. A very early
occurrence of a closed related felid is recorded in
Africa at Laetoli (Upper Unit Laetoli I beds) (Tanzania)
67
(Leakey & Harris, 1987) even if an Asian origin cannot
be ruled out, taking into account the quite large spread
of P. gombaszoegensis in Near East Asia during the
Pleistocene (Hemmer et al., 2001). Moreover, in Eastern Europe this felid has been recorded at in the Late
Pliocene site of Slivnitza (Bulgaria) together with
Canis etruscus.
Consequently, uncertainty about the migration
routes followed by canids, hyaenas and felids still persist. The wolf like and coyote like dogs seems to have
an Asian origin as witnessed by the occurrence of
closely related taxa in the Pliocene locality of Kuruksai
(Tajikistan) (Sotnikova, 1989), even if some uncertainty in the taxonomic position is pointed out by
Spassov & Rook (2006), who suggest a coyote-like adaptation for late forms of Eucyon.
An African origin is possible for the giant hyaena
Pachycrocuta brevirostris and for Lycaon, even if wild
dogs remains have been recorded in Asian localities too
(Martinez Navarro & Rook, 2004). Martinez Navarro
& Palmqvist (1995, 1996) suggested that in the Early
Pleistocene the African dirk-toothed cat Megantereon
whitei dispersed into Europe, replacing the Pliocene
Megantereon cultridens. The taxonomic position of the
Early Pleistocene Megantereon is matter of debate
(Pons Moya, 1987; Turner, 1987; Sardella, 1998;
Hemmer, 2001). A possible African origin for the Early
Pleistocene Megantereon is a crucial point in valuing
the turnover dynamic but not as regard the “turnover”
per se.
As a matter of fact, during the Late Pliocene different
species of canids, hyaenids and felids dispersed into
the Mediterranean region, greatly changing the composition and the structure of the guild of the carnivores.
3 - CARNIVORA GUILD OF THE NORTH
WESTERN MEDITERRANEAN:
DIVERSITY AND TURNOVER
FROM THE MIDDLE PLIOCENE
TO THE EARLY PLEISTOCENE
3.1 - METHODS
Carnivora complexes detected according to the
biochronological scheme proposed by Palombo (in
press) (fig. 1) have been analysed in terms of richness
as well as origination and extinction rates.
Richness can be measured by the total number of
taxa actually or potentially occurring in each
biochronological interval. Nevertheless, the richness
of a single time interval may be overestimated when
first local appearances have been considered as occurring at the beginning of the interval and the taxa that
disappeared as persistent up to the end of it, whereas
they might not actually overlap in time. In order to reduce this bias, and since taxonomic diversity relates to
origination/immigration and extinction/emigration
rates, data have been analysed using the methodology
developed by Foote (2000).
Actually, species richness patterns strictly relate on
origination and extinction rates, thus richness changes
and turnover patterns are strictly connected. Increases
or decreases in richness indicate respectively dispersal
or extinction events, whereas a change in taxonomical
composition between two successive biochrons can be
regarded as a true faunal turnover when changes in species composition result from the concurrent extinction
of existing species and replacement by the immigration/origination of new species. To compute origination and extinction rates, a taxon is assumed to have
originated within the biochron where it is first observed, while a taxon is assumed to have become
extinct within its last observed biochron (Foote, 2000).
3.2 - RESULTS AND DISCUSSION
Trends in richness (fig. 2), and origination/extinction
rates (fig. 3) are quite different in the examined territories that are characterized by differences in physiography as well as in clime.
Indeed, even during the recent past, environmental
conditions affected in different ways migration and diffusion of large mammal species into the North-western
Mediterranean region.
Therefore, new occurrences in the Iberian or Italian
peninsulas (if not related to endemic speciation) resulted from a previous route through France and central
Europe. For this reason, French mammalian faunal
complexes should been more affected by immigration
waves from central Europe, as well as by taxa
latitudinal migrations.
Such a framework seems to fit quite well the main result of our analysis: both richness and origin/extinction
ratios have quite regular trends for Spain, articulated
trends for Italy, and intermediate patterns for France.
Such trend differences seem to be independent by the
quality of fossil record, even if it strongly affected any
analyses. In fact, although faunal complexes have been
taken into account in the present paper instead of local
faunal assemblages, fossil record gaps occurring in different time spans and for different countries affect the
detailed interpretation of data, but not the general
framework.
On the other hand, our results underline a moderate
richness decrease at the Middle to Late Pliocene transition (early to middle Villafranchian). A faunal renewal
has been already noted for the whole faunal data and
for herbivores. It has been related to the Pliocene climate worsening that followed the permanent Arctic
glacial cap onset (Alberdi et al., 1997; Azanza et al.,
1999, 2000; Palombo 2004, in press). As a matter of
fact, the renewal in the carnivore guild took place later
than that of the herbivore one, and just in the Late Pliocene such a guild is modified by appearances and
extinctions (Palombo, in press). It confirms the importance of the structural changes that characterized the
68
Fig. 1: Biochronological range chart of selected Carnivora from North Western Mediterranean (biochronological framework from Palombo,
2005, in press) dating from the middle Pliocene to the early Pleistocene.
Fig. 1 : Comparaison de la distribution chronologique durant le Pliocène moyen et supérieur et le Pléistocène inférieur des Carnivores d’Espagne,
France et Italie (cadre biochronologique d’après Palombo, 2005, sous presse).
69
20
2
18
2
Spain
16
1
12
1
10
1
8
1
6
1
4
0
2
0
0
0,8
0
1,1
1,4
1,7
2
Total Diversity
Estimated mean standing diversity
2,3
2,6
2,9
Ma
3,2
0,8
1,4
1,7
2
Per-taxon Origination rate
2,3
2,6
2,9
Ma
3,2
Per-taxon Extinction rate
2
18
2
France
16
France
2
14
1
12
1
10
1
8
1
6
1
4
0
2
0
0
0
1,1
1,4
1,7
2
Total Diversity
Estimated mean standing diversity
2,3
2,6
2,9
Ma 3,2
0,8
1,1
Total Diversity minus singletons
1,4
1,7
2
Per-taxon Origination rate
20
2,3
2,6
2,9
Ma 3,2
Per-taxon Extinction rate
2
Italy
18
Italy
2
16
2
14
1
12
1
10
1
8
1
6
1
4
0
2
0
0
0,8
1,1
Total Diversity minus singletons
20
0,8
Spain
2
14
0
1,1
1,4
1,7
Total Diversity
2
2,3
2,6
2,9
Ma
3,2
Total Diversity minus singletons
Estimated mean standing diversity
Fig. 2 : “Total Diversity” and “Estimated mean standing diversity”
of Spanish, French and Italian Carnivora calculated following
Foote (2000).
Total Diversity (Ntot) = NFL + NbL + NFt + Nbt; Estimated mean standing
diversity (Nesd) = Ntot – No/2 – Ne/2.
Number of originations (No) = NFL + NFt; Number of extinctions (Ne) =
NFL + NbL.
NFL = taxa that exist only in the interval; NbL = taxa that originate before
the interval but go extinct within it; NFt = taxa that originate in the interval and persisting beyond it; Nbt = taxa that originate before the interval
and persisting beyond it.
Fig. 2 : « Diversité totale » et « Diversité estimée » des Carnivores
d’Espagne, France et Italie calculées d’après Foote (2000).
Diversité totale (Ntot) = NFL + NbL + NFt + Nbt ; Diversité estimée
(Nesd) = Ntot – No/2 – Ne/2.
Nombre d’apparitions (No) = (No) = NFL + NFt ; Nombre d’extinctions
(Ne) = NFL + NbL.
NFL = taxons qui existent seulement dans l’intervalle ; NbL = taxons
qui apparaissent avant l’intervalle et disparaissent durant l’intervalle ;
NFt = taxons qui apparaissent durant l’intervalle et qui persistent durant l’intervalle suivant ; Nbt = taxons qui apparaissent avant l’intervalle et persistent durant l’intervalle suivant.
whole North-western Mediterranean region at the beginning of Early Pleistocene, with Pachycrocuta
brevirostris and Canis etruscus as common elements.
Indeed, in France, Panthera gombaszoegensis is first
recorded only in the latest Early Pleistocene LFAs (for
instance Le Vallonnet) (Moullé, 1998), and Lycaon
falconeri and C. arnensis seems to be missing, even if
the latter has a closely related form (possibly co-specific) in Canis senezensis from Seneze (see above). As
a matter of fact, a new guild structure progressively
took place, with prevailing middle-sized and large carnivores. The “WE” (Azzaroli, 1983; Azzaroli et al.,
0,8
1,1
1,4
1,7
Per-taxon Origination rate
2
2,3
2,6
2,9
Ma
3,2
Per-taxon Extinction rate
Fig. 3: Per-taxon rates of origination and extinction of Spanish,
French and Italian Carnivora calculated following Foote (2000).
Per-taxon origination rate = NFL + NFt)/Ntot/Δt
Per-taxon extinction rate = (NFL + NbL)/Ntot/Δt
NFL = taxa that exist only in the interval; NFt = taxa originate in the interval
and persisting beyond it; NbL = taxa that originate before the interval but
go extinct within it; Δt = time interval.
Fig. 3 : Taux d’apparition et d’extinction «Per-taxon» des Carnivores
d’Espagne, France et Italie calculés d’après Foote (2000).
Taux d’apparition = NFL + NFt) /Ntot/ t ; taux d’extinction = (NFL +
NbL) /Ntot/ t.
NFL = taxons qui existent seulement dans l’intervalle ; NFt = taxons qui
apparaissent durant l’intervalle et qui persistent durant l’intervalle suivant ; NbL = taxons qui apparaissent avant l’intervalle et disparaissent
durant l’intervalle, t = intervalle de temps.
1988; Palmqvist et al., 1999) seems to be more closely
related to migrations affecting the guild of the
herbivores, than to environmental changes.
The different temporal extension of decreasing in diversity that affected the considered countries during
the late Early Pleistocene relays on local disappearances of long survivor carnivores of Pliocene origin
that occurred in the studied territories different times.
For instance, the large felid Megantereon has been latest recorded at Pirro Nord in Italy and at Venta Micena
in Spain (Sardella, 1998) and disappeared in such regions before the Jaramillo event; Puma, that has been
never recorded in Italy and was present in Spain and in
France during the Late Pliocene, survived in France
(Le Vallonnet) (Hemmer et al., 2004) and central Europe (Untermassfeld) (Hemmer, 2001) in LFAs correlated to Jaramillo event. Actually, the occurrence at Le
70
Vallonnet of the puma is matter of debate, being the
fossils (2 lower carnassials) referred to Panthera
pardus by Moullé et al. (in press). In our opinion the
limited recorded sample and its morphometrical
variability leave such taxonomical issue unsolved at
the moment.
During the late early Pleistocene, the reduction in
biodiversity among canids bases more on the disappearance of Lycaon, that survived until the end of the
Early Pleistocene at Le Vallonnet and Colle Curti (Martinez-Navarro & Rook, 2003), than of Canis arnensis
(latest recorded at Upper Valdarno) (Rook, 1993) because canids of the group of Canis mosbachensis partially occupied the niche of the later species. The “end
Early Pleistocene” biodiversity decrease of Italian carnivores (where it approximates the Jaramillo event) it
possibly as consequence of the ongoing decrease of
herbivores recorded at the beginning of Early Pleistocene (Palombo, 2005, unpublished data). In France, on
the other hand, in the LFAs related to Jaramillo, together with the Villafranchian long survivor taxa (e.g.
Puma, Acinonyx and Lycaon) both taxa found in France
for the first time (e.g. P. gombazsoegensis) and newly
appeared ones (possibly U. dolinensis, Garcia &
Arsuaga, 2001) can be recorded (Moullé, 1998;
Hemmer et al., 2004; Martinez Navarro & Rook, 2003;
Palombo & Valli, 2004). In Spain too, the Eurasian
jaguar and the primitive brown bear are recorded in
slightly younger LFAs. Differences in trends can be
due to the scantiness of well documented Early Pleistocene French LFAs. In Italy, LFAs referable to the
Jaramillo event are quite rare, with an overestimation
of uncommon species disappearances.
Accordingly the major guild reorganisation occurred
during the latest Pliocene and the latest Early Pleistocene.
4 - CONCLUSION: WHICH
BIOCHRONOLOGICAL SIGNIFICANCE
FOR THE ALREADY CALLED “WOLF EVENT”?
From the above mentioned issues a question arises:
the renewal of the carnivore guild can be related to the
climate change occurred at the Plio-Pleistocene transition? Is the “WE” a bioevent marking the Plio-Pleistocene transition?
At the beginning of Pleistocene, the Carnivora’s guild
is extensively reconstructed especially as regards to cooperative foraging canids and hyaenids. Pliocrocuta
perrieri and Chamaporthetes lunensis disappeared
whereas the powerful scavenger Pachycrocuta
brevirostris became more and more frequent, the social
canids are already present and spread in the grassland/savanna environments. On the other hand, among
the large felids, Acinonyx pardinensis and possibly
Puma pardoides persisted, while the new incomer
Panthera gombaszoegensis inhabited more close environments together with Megantereon cultridens still
present at least in the late Villafranchian faunas of
Upper Valdarno (Sardella, 1998). Accordingly, the renewal guild occurred in a rather gradual way during the
latest Pliocene and the guild of large carnivores is already definitely modified and in a new equilibrium at
the beginning of the Pleistocene.
The results obtained support the argument that the
“WE”, in a new and wider acception, actually includes
different bioevents affecting the carnivore guilds in
Western Europe (dispersal of Canis, Lycaon, but also
Pachycrocuta and, at least in Italy, Panthera), thus it
cannot be considered as a “marker” of the beginning of
Pleistocene, but, on the contrary, different quite
diachronic bioevents with the dispersal of different carnivore taxa. Among them the most representative dispersion in Europe can be considered that of
Pachycrocuta brevirostris.
ACKNOWLEDGEMENTS
We wish to thank R.-D. Kahlke and P.-E. Moullé for the critical
reading of the manuscript and the useful suggestions. Work supported
by MIUR 60% grants 2005 Prof. M.R. Palombo (“Evoluzione degli
ecosistemi continentali dell’Italia peninsulare durante il Plio-Pleistocene: un approccio multidisciplinare”).
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