AMQ 712 Iannucci et al 36,1 75-90.pub


   

 Available online http://amq.aiqua.it 
ISSN (print): 2279-7327, ISSN (online): 2279-7335 
 
 

Alpine and Mediterranean Quaternary, 36 (1), 2023, 75-90 

1. INTRODUCTION 
 
During the Quaternary, marked climatic and envi-

ronmental changes took place at a global scale, most 
notably affecting meteorological and oceanographic 
circulations, the intensity of the glacial activity in the 
Northern Hemisphere, and the frequency and amplitude 
of oscillations between cool-arid and warm-humid condi-
tions (Shackleton, 1995; Flesche Kleiven et al., 2002; 
Gibbard et al., 2005; Ehlers & Gibbard, 2007; Etourneau 
et al., 2010; Gibbard & Head, 2020). In turn, these 
changes promoted substantial reorganizations of the 
large mammal fauna. Forty years ago, Azzaroli (1983) 
recognized and named three major “dispersal 
events” (Repenning, 1967, 1980), referring to them as 
“short periods of rapid intercontinental migrations and 
faunal replacements” (Azzaroli, 1983, p. 117), namely 
the “Elephant-Equus event” (between 3.0 and 2.5 Ma), 
the “Wolf event” (~1.7 Ma), and the “end-Villafranchian 
dispersal event” (~1.0-0.9 Ma). In the following years, 
the proposed timing of these events was further dis-

cussed and refined (Azzaroli et al., 1988; Azzaroli, 1995; 
Gliozzi et al., 1997), eventually correlating the Elephant-
Equus event with what is currently recognized as the 
Pliocene-Pleistocene transition (~2.6 Ma), the Wolf 
event with the Gelasian-Calabrian transition (~1.8 Ma; 
the Pliocene-Pleistocene boundary prior to 2009), and 
the end-Villafranchian event with the Early-Middle Pleis-
tocene transition (~0.8 Ma).  

Whilst Azzaroli (1983) regarded the aforemen-
tioned dispersal events as moments of marked faunal 
renewals and recognized in some cases the gradual 
character of the associated turnover, the approximate 
synchronicity between dispersal events and relevant 
geological transitions exacerbated their geochronologi-
cal importance. This resulted in a widespread use of the 
appearance of the “representative” species after which 
the events were named (e.g., the genus Equus for the 
Elephant-Equus event) as a stratigraphic datum (see 
Iannucci & Sardella, 2023, for discussion). Essentially, 
there are at least two aspects of Azzaroli’s dispersal 
events that resulted in ambiguous applications. First, 

https://doi.org/10.26382/AMQ.2023.03 

BEWARE OF THE “WOLF EVENT” - REMARKS ON LARGE MAMMAL DISPERSALS IN EUROPE 
AND THE LATE VILLAFRANCHIAN FAUNAL TURNOVER 

 
Alessio Iannucci, Beniamino Mecozzi, Raffaele Sardella 

 
Dipartimento di Scienze della Terra (PaleoFactory lab.), Università Sapienza, Roma, Italy. 

 
Corresponding author: A. Iannucci <alessio.iannucci@uniroma1.it>  

 
ABSTRACT: The “Wolf event” is a prominent concept in large mammal biochronology of western Europe. It was defined in the 
1980s as an intercontinental “dispersal event”, best represented by the arrival of a “primitive wolf”, Canis etruscus, but also invol-
ving other species. The Wolf event denoted the late Villafranchian faunal turnover, first expressed in Italy in the Olivola Faunal 
Unit. This event was also considered approximately coincident with the Pliocene-Pleistocene boundary adopted prior to 2009 (~1.8 
Ma, Gelasian-Calabrian transition), hence indicating important environmental changes and representing a relevant tool for correla-
tion. Whilst it became soon clear that sporadic finds of modern canids (and, to some extent, other species) pre-dated the age as-
sumed for the Wolf event, several authors continued to use the term and to associate it to the late Villafranchian, referring to the 
“massive expansion” of the species involved, rather than their first appearance in the European fossil record. Several bioevents 
traditionally included in the Wolf event and others that have been considered to occur later are today already documented in midd-
le Villafranchian faunas. The “Pachycrocuta brevirostris event”, proposed as a replacement term for the Wolf event, based on 
current evidence would be characterized by the arrival in Europe of the giant hyena P. brevirostris and Panthera gombaszoegen-
sis, and the increase in the documentation of other species traditionally included in the Wolf event. However, this does not corre-
spond to a sharp faunal turnover as traditionally envisioned for the Wolf event and it is possibly heralded in faunas slightly older 
than Olivola at ~2.0 Ma. In other terms, available evidence highlights the rather diachronic nature of large mammal dispersal oc-
curred in the late middle and early late Villafranchian (late Gelasian, ~2.2–1.8 Ma), pushing to critically evaluate the biochronologi-
cal, paleoecological, and paleobiogeographical significance of each bioevent. For instance, the arrival of Hippopotamus in Europe 
is now attested since ~2.2 Ma, documenting an African dispersal of a species linked to humid conditions in a context that is gene-
rally deemed to denote the spread of open-adapted faunal elements of mainly Asian affinities. 
 
Keywords: Biochronology, bioevent, faunal renewal, Pleistocene, Villafranchian.  
 



   

 

 
 

76 

 
 

Iannucci A. et al. 

although named after one or few taxa, these periods of 
faunal turnover involved, by definition, many species. 
Second, they were envisioned as “short”, but not proper-
ly geologically instantaneous intervals. To this list can 
be added that Azzaroli (1983) emphasized the link be-
tween faunal and environmental changes, or, in other 
terms, he inherently included a paleoecological compo-
nent in the biochronological event.  

Of course, this is not to say that implementations 
of the dispersal event concept with a meaning that devi-
ate from that used by Azzaroli (1983) are necessarily 
wrong. Indeed, as in general for other biochronological 
terms, concepts, principles, and practices, they are not 
formally defined within a stratigraphic code, which 
means that there exist a (legit) variety of approaches 
among different researchers (e.g., Gradstein et al., 
1985; Lindsay, 2003; Palombo & Sardella, 2007; Pal-
ombo, 2009). However, this unavoidably leaves room 
for ambiguity, most notably when divergent adoptions of 
a dispersal event are discussed as if referring to the 
same thing, while different authors might have invested 
the same term with different meanings. Conceptual di-
chotomies aside, over the years several discoveries 
have pushed to reconsider the timing of specific bio-
events (i.e., those linked to a single taxon), and thus, 
partly depending on the different acceptances with 
which Azzaroli’s dispersal events have been used, au-
thors opinions diverge as to whether reinterpret, reimag-
ine, or abandon the use of specific dispersal events. 

The Wolf event played a special role in Quaternary 

large mammal biochronology, especially in western Eu-
rope (Fig. 1), given the approximate coincidence be-
tween this event (in its original formulation) and the for-
merly adopted Pliocene-Pleistocene boundary at ~1.8 
Ma (i.e., prior to the placement of the Pleistocene 
Epoch/Series at ~2.6 Ma, in 2009; Aguirre & Pasini, 
1985; Pillans, 2004; Clague, 2006; Gibbard & Head, 
2010; Gibbard et al., 2010; Head & Gibbard, 2015; 
Capraro & Maiorano, 2023). 

Here, using the Wolf event as a case study, we 
offer a discussion, partly conceptual, partly factual, on 
the dispersal of wolf-like canids into Europe and on 
some other specific bioevents currently recognized to 
have occurred in the late middle and late Villafranchian 
(late Gelasian, ~2.2–1.8 Ma) that have been the subject 
of debate and in some instances considered in the 
framework of the Wolf event.  

 
2. THE “WOLF EVENT” 

 
Azzaroli (1983) remarked that turnover associated 

with the Wolf event was overemphasized in the Italian 
fossil record due to the lack of middle Villafranchian 
faunas, but he included in the Wolf event the arrivals of 
Leptobos etruscus (replacing L. stenometopon), Sus 
strozzii, Pachycrocuta brevirostris, and Canis etruscus. 
The latter is the “wolf” after which the event was named. 
This turnover would correspond to the beginning of the 
late Villafranchian, and to the Olivola Faunal Unit (FU). 
Other groups of Canis were considered to appear only in 

Fig. 1 - Main fossiliferous localities mentioned in this work: 1 - Fonelas P-1; 2 - La Puebla de Valverde; 3 - Perrier-Les Etouaires, Roca-
Neyra; 4 - Senèze; 5 - Vialette; 6 - Saint Vallier; 7 - Oosterschelde; 8 - Olivola, Quercia; 9 - Valdarno Superiore (several localities); 10 - 
Montagnola Senese; 11 - Vigna Nuova, Chiusi Basin, Torre Picchio; 12 - Monte Riccio; 13 - Bocchignano, Castel San Pietro; 14 - Coste 
San Giacomo, Fontana Acetosa; 15 - Elis, Aetorráchi, Hághios Demétrios, 16 - Gerakarou 1. 



   

 

 
 

77 

 
 
“Wolf event” and large mammal dispersal 

and the proper spread of the group since the beginning 
of the late Villafranchian. 

Further evidence of middle Villafranchian Canis 
has piled up in the following decades, some of which 
perhaps related to wolf-like canids, either deriving from 

the subsequent Tasso FU (Azzaroli, 1983). 
Azzaroli et al. (1988) remarked the coinci-
dent disappearance of Nyctereutes and 
Gazella, the appearance, possibly related 
to evolutionary changes, of Eucladoceros 
dicranios and Dama nestii (currently often 
referred to ‘Pseudodama’, see Cherin et 
al., 2022, and references therein), and the 
arrival of another species, Panthera tosca-
na (today referred to P. gombaszoegen-
sis), alongside the bioevents previously 
recognized. Concerning the latter, howev-
er, Azzaroli et al. (1988, p. 84) referred to 
the “massive expansion in Europe” of the 
considered species, rather than their earli-
est appearance in the fossil record, again 
leaving room for divergent interpretations. 
Therefore, there are different aspects of 
Azzaroli’s Wolf event that need to be criti-
cally evaluated based on the available 
evidence. It is worth considering: first, 
whether the dispersal of wolf-like canids in 
Europe is coincident with the beginning of 
the late Villafranchian in correspondence 
of the Olivola FU or not; second, how 
many of the aforementioned bioevents are 
indeed part of the same “dispersal event”. 
In the following, these two points are briefly 
discussed. 

When Azzaroli (1983) schematized 
the Wolf event and placed it in correspond-
ence of the Olivola FU, remains of possibly 
older Canis were already known at least 
from the French site of Senèze (Martin, 
1973), but the chronological consistency of 
the faunal assemblage recovered from this 
locality was often doubted (Masini & Torre, 
1990; Mazza & Rustioni, 1994; Rook & 
Torre, 1996). Further canid remains were 
recovered from Coste San Giacomo 
(predating Olivola and now dated at ~2.2 
Ma, Florindo et al., 2021; Coste San Gia-
como FU) and attributed to Canis etruscus 
by Rook & Torre (1996), in line with ongo-
ing investigation. Rook & Torre (1996) also 
reported the presence of a partial hemi-
mandible of a canid recovered from Quer-
cia (near Olivola and stratigraphically lower 
than it; Iannucci, 2023), accepted a middle 
Villafranchian age for the canids of Se-
nèze, and concluded that the earliest dis-
persal of modern dogs occurred during the 
middle Villafranchian. Nonetheless, Rook 
& Torre (1996, p. 499) argued that “the 
meaning of the “wolf-event” for the begin-
ning of the Late Villafranchian does not 
loose its value of “faunal event” in the 
sense of marked change in faunal assem-
blages”. In the seminal synthesis on Italian biochronolo-
gy by Gliozzi et al. (1997), the status of the Wolf event 
was not directly discussed, but the authors reiterated the 
first occurrence of wolf-like canids (C. etruscus) in Italy 
from the middle Villafranchian of Coste San Giacomo 

 

Fig. 2 - Distribution of Canis (a) in the middle Villafranchian of western Europe: 1 - 
Coste San Giacomo, Fontana Acetosa; 2 - Oosterschelde; 3 - Roca-Neyra; 4 - Se-
nèze; 5 - Chilhac 2; 6 - Saint Vallier; 7 - Quercia; 8 - Montagnola Senese; 9 - Vigna 
Nuova, Torre Picchio. Silhouettes of canids modified from: Iurino et al. (2022; Canis 
etruscus, Canis sp.); Bartolini Lucenti & Spassov (2022; Canis (X.) lycanoides); Wiki-
media Commons (https://commons.m.wikimedia.org/wiki/File:Canis_arnensis_restora-
tion.jpg; Canis arnensis). Distribution of Hippopotamus (b) in the middle Villafranchian 
of western Europe: 1 - Senèze; 2 - Chiusi Basin; 3 - Castel San Pietro; 4 - Coste San 
Giacomo, Fontana Acetosa; 5 - Hághios Demétrios, Aetorráchi. Silhouette of hippopot-
amus modified from PhyloPic, by Zimices (phylopic.org/image/c2d68ebb-50ec-45f4-
8cd1-6cf52ad02286). 



   

 

new discoveries or reconsiderations of old collections 
(Fig. 2a). Two lower incisors assigned to Canis sp. were 
reported by Fondi (1972) from Montagnola Senese, 
whose fauna has been referred to the Coste San Giaco-
mo FU (Bona et al., 2015). A fragmentary premolar and 
a portion of a distal metapodial of Canis sp. are known 
from Torre Picchio (Girotti et al., 2003), also referred to 
the Coste San Giacomo FU (Bona et al., 2015). During 
the revision of the carnivoran record of Saint Vallier, 
Argant (2004) identified the presence of a partial neu-
rocranium referable to the genus Canis. The locality is 
customarily biochronologically placed between Mon-
topoli and Coste San Giacomo FUs, with estimated 
ages ranging from 2.5 to 2.0 Ma (Viret, 1954; Azzaroli, 
1970, 1977; Torre et al., 1992; Gliozzi et al., 1997; 
Guérin, 2004; Nomade et al., 2014; Brugal et al., 2020). 
Reumer & Piskoulis (2017) assigned to Canis cf. 
etruscus a mandibular fragmented recovered from the 
Oosterschelde estuary, whose Early Pleistocene fossil 
fauna, although resedimented in Late Pleistocene de-
posits and collected during dredging activity, is referred 
to the middle Villafranchian, or to ~2.4–2.1 Ma (Scager 
et al., 2017). Azzarà et al. (2022) described a portion of 
the axial skeleton of a single individual of Canis sp. re-
covered from the middle Villafranchian (Coste San Gia-
como FU) site of Vigna Nuova, considering it very simi-
lar to comparable material of C. etruscus known from 
Olivola (Torre, 1967). Finally, concerning the long de-
bate on the possibility of age heterogeneity in the faunal 
assemblage from Senèze (see Delson et al., 2006), 
Pastre et al. (2015) performed 40Ar/39Ar datings that 
constrain the sequence between ~2.2 and 2.09 Ma. 
Recently, U-Pb dating on volcanic zircons was also 
applied at Senéze, with results providing a mean age of 
of 2.100 ± 0.029 Ma (Paquette et al., 2021). 

An even older record of Canis sp. would be that of 
Vialette (Heintz et al., 1974), which Lacombat et al. 
(2008) referred to 3.14 Ma. However, the attribution of 
the material and the chronology of the site have been 
contested. The problems concerning the chronology of 
the site or, more precisely, the mixing of different faunal 
elements among those labelled as from Vialette in the 
collection of the Musée Crozatier (Le Puy-en-Velay), 
has been commented by several authors (Guérin, 2005; 
Van der Made et al., 2014; Palombo & Alberdi, 2017). 
Perhaps the clearest example of chronological hetero-
geneity is represented by the identification of some re-
mains belonging to Late Miocene suids, namely as-
signed to Listriodon and cf. “Microstonyx” major (Van 
der Made 2005; Van der Made & Moullé, 2005), the 
latter species being more commonly placed in Hippopot-
amodon in recent literature (Iannucci et al., 2021a). As 
for the taxonomic attribution of the canid material to 
Canis, Lacombat et al. (2008, p. 67) argued that: “The 
morphology, the size and the proportions of these re-
mains allow us to exclude genera occurring in the Early 
Pliocene”, while Böhme et al. (2021) recently suggested 
it could rather belong to Eucyon. On the other hand, 
Böhme et al. (2021) reported material of wolf-like canids 
from Perrier-Les Etouaires, referring it to 2.78 Ma follow-
ing Nomade et al. (2014). This chronology calls for fur-
ther discussion. Nomade et al. (2014) provided this age 
for pumices of la Côte d’Ardé, which they considered 

stratigraphically close to Perrier-Les Etouaires “classical 
site”. However, as pointed out by Iannucci & Sardella 
(2023), this age cannot be accepted uncritically for all 
the mammal remains recovered from Perrier-Les 
Etouaires. Indeed, it is known that the fauna of Perrier-
Les Etouaires contains early Villafranchian and middle 
Villafranchian species, which originated from layers of 
different ages (Poidevin et al., 1984; Palombo & Valli, 
2004). In any case, following the dating and correlations 
proposed by Nomade et al. (2014) for the entire area of 
Perrier, the fossil findings should be constrained be-
tween ~3.1 and 2.6 Ma, an age that would be thus 
enough to make the record of wolf-like canids from Per-
rier-Les Etouaires reported by Böhme et al. (2021) the 
earliest (excluding Vialette) in Europe (Iannucci & Sar-
della, 2023), although the material has not been de-
scribed. Indeed, the presence of Canis aff. etruscus 
from Etouaires was reported by Heintz et al. (1974) 
based on material part of the Bravard Collection housed 
in the British Museum listed by Lydekker (1885, p. 126). 
However, the material mentioned by Lydekker (1885) is 
possibly from younger deposits at Tour-de-Boulade, and 
indeed the author considered it undistinguishable from 
C. lupus. This view was reaffirmed by Torre (1979), who 
stated that these fossils were of large size, similar to that 
observed in wolves of the last Glaciations (which would 
correspond to the second part of Late Pleistocene). 

Marciszak et al. (2023) also reported unpublished 
material of Canis cf. etruscus from Węże 2, referring it to 
2.9–2.6 Ma, also listing further material from Poland 
potentially of middle Villafranchian age. 

In sum, while the evidence for some of the afore-
mentioned findings might not be conclusive, it has long 
been quite clear that the arrival of wolf-like canids in 
Europe is at least as early as the Coste San Giacomo 
FU (middle Villafranchian), and there is further putative 
evidence that would suggest an even older age. It is 
worth reiterating that the occurrence of wolf-like canids 
already in the middle Villafranchian did not discourage 
several authors to continue using the Wolf event to refer 
to the late Villafranchian faunal turnover as a whole, 
emphasizing the increase in the abundance of wolf-like 
canids, rather than their earliest appearance in the fossil 
record (e.g., Rook & Torre, 1996). This late Villafran-
chian “massive expansion”, as it has been referred to by 
Azzaroli et al. (1988) and generally agreed in subse-
quent research (e.g., Gliozzi et al., 1997), is well docu-
mented in Italy, for instance by historical samples from 
Olivola and the Upper Valdarno, and by the abundant 
record of Pantalla (Cherin et al., 2013, 2014). 

 
3. THE “HIPPO EVENT” 

 
The earliest dispersal of Hippopotamus in Europe 

was not among the bioevents listed by Azzaroli (1983) 
as part of the Wolf event, but we consider particularly 
appropriate to discuss it here as there is now evidence 
of late middle Villafranchian (late Gelasian) hippopota-
muses from several European localities, some of which 
also yielded early Canis (Fig. 2b). In particular, a frag-
mentary incisor of Hippopotamus sp. is known from 
Coste San Giacomo (Bellucci et al., 2012, 2014; Sar-
della, 2012). The age of the fossil horizon was consid-

 
 

78 

 
 

Iannucci A. et al. 



   

 

chian hippopotamus in Europe (dated at ~2.2 Ma), but it 
is not the only. Cassoli & Segre Naldini (1984) listed 
Hippopotamus sp. among the faunal remains recovered 
from Fontana Acetosa, like Coste San Giacomo, another 
locality of the Anagni Basin in central Italy. The authors 
remarked the similar nature of the fossiliferous levels of 
Coste San Giacomo and Fontana Acetosa, yellow 
sands, although somewhat more clayey at Fontana Ace-
tosa, but did not exclude the possibility of a slightly 
younger age for the latter site. The presence of Hippo-
potamus seemed indeed at odds with the then accepted 
first appearance of the species in the Tasso FU 
(Azzaroli, 1983). When Bellucci et al. (2012) recognized 
the presence of Hippopotamus at Coste San Giacomo, 
they also accepted the correlation between the two 
sites, although the fauna of Fontana Acetosa would be 
in need of a systematic study to clarify the biochronolog-
ical value of the species reported. 

During the revision of the fossil collection stored at 
Faculté des Sciences de l'Université Claude Bernard, 
Lyon I, a first phalange (FSL 211082), previously as-
cribed to Equus sp., was attributed to Hippopotamus cf. 
antiquus by Mazza & Rustioni (1994). The fossil was 
recovered from Domeyrat, one of the toponyms general-
ly considered as part of Senèze locality. The presence 
of hippopotamuses at Senèze was previously listed by 
Jung (1946) and Bout (1960), but then excluded from 
subsequent faunal lists (e.g., Heintz et al., 1974), possi-
bly due the supposedly later arrival of Hippopotamus in 
Europe. It seems conceivable that reports older than the 
work by Mazza & Rustioni (1994) were based on further 
undescribed material and, in any case, even after that 
Mazza & Rustioni (1994) reaffirmed the occurrence of 
hippopotamuses at Senèze, these were seldom consid-
ered in subsequent biochronologic schemes (but see 
Arribas et al., 2009), likely due to the uncertainty on the 
chronology of the locality (see previous section above). 
Nonetheless, as discussed in the previous section, sev-
eral radiometric estimates now constrain the succession 
of Senèze at ~2.2–2.1 Ma (Nomade et al., 2014; Pastre 
et al., 2015; Paquette et al., 2021), approximately coeval 
with Coste San Giacomo. 

 Several findings of middle Villafranchian hippopot-
amuses are known from the area of Elis, in Greece, as 
recently reviewed by Athanassiou (2022). These re-
mains include: dentognathic material described by The-
nius (1955), which according to Athanassiou (2022) can 
be considered of earliest Pleistocene age owing to inver-
tebrate biochronology; further osteological and dental 
finds mentioned by Symeonidis & Theodorou (1986) 
from Hághios Demétrios; a juvenile fragmented cranium 
from Aetorráchi (Reimann & Strauch, 2008).  

Other hippopotamus remains attributed to H. an-
tiquus were recovered from the Chiusi Basin (Cuscani 
Politi, 1966, 1971; Mazza, 1995; Pandolfi & Petronio, 
2015). An accompanying faunal assemblage was also 
reported by Pandolfi & Petronio (2015), including Gazel-
la sp., Axis (=‘Pseudodama’) cf. nestii and Eucladoceros 
sp., although not described and made up of surface 
finds, the authors referred it to the Coste San Giacomo 
or Olivola FUs. The occurrence of Gazella would point 
towards the former, considering that the taxon is not 
recorded in Italian localities younger than the Coste San 

ered around 2.1 Ma by Bellucci et al. (2014), based on 
magnetostratigraphy, pollen, and small mammals, and 
further refined at ~2.2 Ma by Florindo et al. (2021). Bel-
lucci et al. (2012) firstly identified the specimen as be-
longing to a hippopotamus, although a more precise 
taxonomic attribution is clearly precluded by the scanty 
nature of the sample. Sardella et al. (2018) clarified that 
the hippopotamus incisor from Coste San Giacomo was 
not collected during systematic excavations but is part of 
the 1980s field collection, causing some authors to 
doubt the age of the finding (Marra et al., 2018; Martino 
& Pandolfi, 2022). However, field activities and excava-
tions at Coste San Giacomo co-directed by one of us 
(RS) have pointed out that the vertebrate assemblage 
comes from a single fossiliferous level, from which the 
old collections can also be related (Bellucci et al., 2012, 
2014; Bona et al., 2015; Strani et al., 2015; Palombo et 
al., 2017). It is worth noting that the fauna of Coste San 
Giacomo has long been considered a homogeneous 
assemblage, besides representative of the homonymous 
FU, the latest of the middle Villafranchian (Gliozzi et al., 
1997). 

The reason why the hippopotamus record of Coste 
San Giacomo has often been considered controversial 
(e.g., Martínez-Navarro et al., 2015; Pandolfi & Petronio, 
2015) seems rather contingent. Traditionally, the earliest 
occurrence of Hippopotamus in Italy was placed in the 
Tasso FU, based on several historical findings from the 
Upper Valdarno (Nesti, 1820; Leonardi, 1947; Azzaroli, 
1977; Gliozzi et al., 1997). Napoleone et al. (2003) sug-
gested that the hippopotamuses might be of a younger 
age than that of the rest of the main Upper Valdarno 
fauna, considering the lack of their remains among 
those collected during modern excavations in the area. 
This possibility was also previously evoked by Faure 
(1985) and Mazza (1991). It is worth noting that most of 
the fossils from the Upper Valdarno is part of historical 
collections gathered at least since the nineteenth centu-
ry (Rook et al., 2013). Moreover, the absence of hippo-
potamus remains from certain deposits might also be 
related to their ecological requirements, namely their 
sensitivity to the presence and amount of water (Mazza 
& Bertini, 2013). Martínez-Navarro (2004, 2010) and 
Rook & Martínez-Navarro (2010), following the sugges-
tion of Napoleone et al. (2003) on the younger age of 
the Upper Valdarno hippopotamuses, remarked that 
consequently Venta Micena yielded the oldest European 
record of Hippopotamus antiquus (although other older 
records were known, e.g., that from Monte Riccio, Maz-
zini et al., 2000). Mazza & Bertini (2013, p. 195) also 
argued that “The first certified occurrence of Hippopota-
mus antiquus is from the Early Pleistocene locality of 
Venta Micena”, but not excluding the occurrence of earli-
er findings of uncertain taxonomic status (e.g., the au-
thors quoted Fontana Acetosa). In brief, the finding of 
the hippopotamus of Coste San Giacomo was published 
at a time when some researchers were hypothesizing a 
later chronology for the arrival of Hippopotamus into 
Europe (Bellucci et al., 2012; Sardella, 2012). This was, 
however, not universally accepted (e.g., Arribas et al., 
2009). 

In general, the finding of Coste San Giacomo rep-
resents the earliest occurrence of a middle Villafran-

 
 

79 

 
 
“Wolf event” and large mammal dispersal 



   

 

 
 

80 

Giacomo FU (Masini et al., 2013; Bellucci & Sardella, 
2015). 

The scheme proposed by Azzaroli (1983) featured 
the first appearance of Hippopotamus in the Tasso FU, 
which has been most influential in the biochronological 
correlation of several faunas. The implications for the 
inferred age of three localities are especially worth men-
tioning here, recognizing the presence of hippopotamus-
es in Europe already in the middle Villafranchian, name-
ly Castel San Pietro, Bocchignano, and Monte Riccio 
(all in Italy). At Castel San Pietro, several fossil mammal 
remains were collected and partly described during the 
nineteenth century, as a by-product of the exploitation of 
a lignite mine (Tuccimei, 1889a, 1889b, 1891, 1898; 
Maxia 1949). The presence of hippopotamuses from the 
area of Castel San Pietro was reported by Meli (1882) 
and Tuccimei (1891). Pandolfi et al. (2017) carried out a 
revision of the scanty mammal remains, and further 
analysis based on ostracods and paleofloristic remains. 
Their results converged to suggest a Gelasian deposi-
tion of the lignite beds of Castel San Pietro (~2.5–1.8 
Ma) in a warm-humid and forested area. Concerning the 
hippopotamuses, the authors casted doubt on their as-
sociation with the rest of the fauna and commented that 
the whereabouts of the material are unknown. Marra et 
al. (2018) argued for the presence of two different as-
semblages at Castel San Pietro, the older with Anancus 
arvernensis, the younger with H. antiquus. Nonetheless, 
the cooccurrence of A. arvernensis and Hippopotamus 
cannot be ruled out and, if confirmed, would point to a 
reference to the Coste San Giacomo FU, as exemplified 
by the record from the reference fauna of this FU 
(Bellucci et al., 2014). 

Bocchignano is another site whose hippopotamus-
es remains have been considered indicative of faunal 
mixing (Marra et al., 2018). The fossil remains recov-
ered from the site were reported and described by Tuc-
cimei (1889b, 1891, 1893). Kotsakis (1988) revised the 
taxonomic attribution of the arvicoline rodents, referring 
them to Mimomys polonicus or Mimomys pliocaenicus. 
Recently, Marra et al. (2018) suggested the presence of 
two different assemblages, the oldest with Mimomys 
(Saint Vallier to Olivola FUs), the youngest with Hippo-
potamus (Tasso FU). As in the case of Castel San 
Pietro, the cooccurrence of the two species is arguably 
conceivable (if the arvicoline belongs to M. pliocaenicus) 
and in agreement with a correlation of the fauna with the 
Coste San Giacomo or Olivola FUs (Bellucci et al., 
2014; Bona et al., 2015). 

Unlikely of middle Villafranchian age but worth of 
consideration here is the diverse fauna of Monte Riccio, 
including Prolagus sp., cf. Mammuthus meridionalis, 
Sus strozzii, H. antiquus, Leptobos cf. etruscus, Procap-
reolus sp., Eucladoceros ctenoides, Axis (= 
‘Pseudodama’) nestii, Stephanorhinus cf. etruscus, 
Equus stenonis, Vulpes cf. alopecoides, Canis etruscus, 
and Megantereon cultridens (Mazzini et al., 2000). In-
deed, this assemblage has been referred to the Tasso 
FU due to the presence of H. antiquus (Mazzini et al., 
2000; Marra et al., 2018), while Croitor (2012) suggest-
ed a correlation with the Olivola FU or even a slightly 
older age, based on the ruminants. 

In brief, there is evidence for a middle Villafran-

chian dispersal of hippopotamus into Europe ~2.2–2.1 
Ma, partly documented by remains collected from some 
of the localities that have been also important to recog-
nize an earlier (than what proposed by Azzaroli, 1983) 
arrival of modern canids, namely Coste San Giacomo 
and Senèze. The “Hippo event” is of paleobiogeographic 
and paleoecological relevance, in that it testifies to the 
dispersal into Europe of a taxon of clear African origin 
and of distinct ecology. Indeed, though not necessarily 
limited to warm environments, remains of Hippopotamus 
are indicative of the presence of permanent water bod-
ies and humid climatic conditions (Candy et al., 2006, 
2010; Bellucci et al., 2012; Russo Ermolli et al., 2010; 
Mazza & Bertini, 2013; Adams et al., 2022). Moreover, 
morphological considerations and dietary proxies sug-
gest that H. antiquus was even more adapted to an 
aquatic lifestyle than H. amphibius (Palmqvist et al., 
2003, 2008; Adams et al., 2022). As mentioned previ-
ously, perhaps these ecological requirements explain 
the patchy distribution of middle and late Villafranchian 
occurrences of hippopotamuses in Europe, whereas 
since ~1.5 Ma Hippopotamus become a common ele-
ment of the European terrestrial ecosystems (Caloi et 
al., 1980; Faure, 1985; Kahlke, 1989, 2001; Mazza, 
1991, 1995; Kahlke et al., 2011; Van der Made et al., 
2017; Fidalgo et al., 2021; Mecozzi et al., 2021; Adams 
et al., 2022; Strani et al., 2022). In this regard, it is worth 
mentioning that paleoenvironmental reconstruction 
based on ungulate dietary adaptations attest the pres-
ence of humid subtropical-like environments at Coste 
San Giacomo (Strani et al., 2015), whereas comparable 
analyses point to the occurrence of more open condi-
tions at Olivola, where no hippopotamuses are known 
(Strani et al., 2018).  

 
4. OTHER BIOEVENTS 

 
Apart from the arrival of C. etruscus, several other 

bioevents were listed by Azzaroli (1983) and Azzaroli et 
al. (1988) as part of the Wolf event, and others have 
been proposed and discussed over the years. In particu-
lar, Azzaroli (1983) listed L. etruscus, S. strozzii, and P. 
brevirostris. The giant hyena is the only species of this 
contingent whose first appearance in Europe is still relat-
ed to the late Villafranchian faunal turnover (see section 
5). Indeed, while the “massive expansion” (Azzaroli et 
al., 1988) of L. etruscus and S. strozzii can still be 
placed in correspondence of Olivola and Tasso FUs, 
remains of both species are already documented in mid-
dle Villafranchian faunas (Masini & Sala, 2007; Cherin et 
al., 2018, 2019; Iannucci et al. 2020; Sorbelli et al., 
2023).  

Isolated dental remains of large-sized suids, which 
are generally referred to S. strozzii, are known from 
several middle Villafranchian sites, including  Saint Val-
lier, Valdeganga II, Coste San Giacomo, Quercia, and 
Vigna Nuova, among others, and the locality of Senèze 
yielded an almost complete skeleton (Schaub, 1943; 
Mein et al., 1978; Azzaroli et al., 1988; Faure, 2004; 
Cherin et al., 2018; Iannucci et al., 2020; Azzarà et al., 
2022; Iannucci, 2023).  

Remains of L. etruscus are also known from Se-
nèze (Masini, 1989; Cherin et al., 2019). Moreover, the 

 
 

Iannucci A. et al. 



   

 

debate surrounding the taxonomy and evolutionary rela-
tionships of middle and late Villafranchian Leptobos 
spp., the presence of chronologically overlapping differ-
ent lineages, and the possible appearance per-evolution 
(rather than per-dispersal) of the species in Europe, 
clearly push to exercise caution in biochronological cor-
relations (Cherin et al., 2019; Sorbelli et al., 2023).  

Objective difficulties in the attribution of isolated 
remains and disagreement on the taxonomy also com-
plicate identifying and resolving the timing of species-
level event of Eucladoceros spp. and ‘Pseudodama’ 
spp. between the middle and late Villafranchian, as re-
flected by the many samples left in open taxonomy 
(e.g., Bellucci et al., 2014; Pandolfi & Petronio, 2015; 
Azzarà et al., 2022).  

Apart from evolutionary changes in the aforemen-
tioned cervid lineages, Azzaroli et al. (1988) also men-
tioned the extinctions of Nyctereutes and Gazella. Both 
are still considered typical middle (or even early) middle 
Villafranchian taxa, but at least Gazella is present in 
some faunas that could be regarded as late Villafran-
chian or as transitional between the middle and the late 
Villafranchian (e.g., Fonelas P-1 and Gerakarou 1, see 
section 5).  

The arrival of the Caprinae Procamptoceras bri-
vatense was listed among the bioevents characterizing 
the late Villafranchian turnover by Gliozzi et al. (1997), 
based on its occurrence at Olivola (Azzaroli, 1950), but 
the species is already present in Senèze and tentatively 
listed in older faunas (Heintz et al., 1974; Palombo & 
Valli, 2004). In general, the record of P. brivatense is 
arguably too patchy to be stressed in a biochronological 
context, although this might change with further discov-
eries.  

The arrival of Panthera gombaszoegensis was 
added by Azzaroli et al. (1988) among the bioevents 
characterizing the Wolf event. The first appearance of 
the species in the Italian fossil record is still in the Olivo-
la FU (Ficcarelli & Torre, 1968; Gliozzi et al., 1997; Pal-
ombo, 2009). The locality of Gerakarou 1 also yielded 
remains of the species (Koufos, 1992, 2014). The occur-
rence of the species at La Puebla de Valverde, calibrat-
ed between Feni and Olduvai subchrons at ~2.12–1.92 
Ma and generally considered close in age to but more 
progressive than Saint Vallier (Sinusía et al., 2004; 
Cuccu et al., 2023), was listed by Madurell-Malapeira et 
al. (2014), but not reported by Kurtén & Crusafont-Pairó 
(1977) and Cuccu et al. (2023). 

When Azzaroli (1983) discussed the Wolf event, C. 
etruscus was the only modern canid whose arrival was 
correlated with the Olivola FU, while the first appear-
ance of Canis arnensis and Canis falconeri was placed 
in the following Tasso FU. Both species are now already 
documented in middle Villafranchian faunas of western 
Europe. Recently, Bartolini Lucenti & Spassov (2022) 
substantially extended the chronological range of the 
wild dog Canis (Xenocyon) falconeri, referring to this 
taxon a mandible from Roca-Neyra (~2.6 Ma), which 
suggests a patchy distribution of the species in Europe 
since the beginning of the middle Villafranchian. The 
earliest record of C. arnensis known to date is from Se-
nèze, based on the available dating for the site and the 
generally accepted synonymy between C. arnensis and 

C. senezensis (Brugal & Boudadi-Maligne, 2011; Bartoli-
ni Lucenti & Rook, 2016). Several early occurrences of 
Canis sp. listed previously (see section 2) might also 
belong to species other than C. etruscus (Fig. 2a). 

 
5. THE “PACHYCROCUTA BREVIROSTRIS EVENT” 
AND THE BEGINNING OF THE LATE VILLAFRAN-
CHIAN 

 
The term “Pachycrocuta brevirostris event” or simi-

lar expressions have been sporadically used since the 
1990s, e.g., by Masini & Torre (1990; “Pliohyaena brevi-
rostris Event”), Torre et al. (1992; “Pachycrocuta brevi-
rostris dispersal event”), Palombo et al. (2008; 
“Pachycrocuta event”), but it is only with the work of 
Martínez-Navarro (2010) that this concept acquired the 
meaning of a replacement term for Azzaroli’s Wolf 
event. As mentioned in section 2, the evidence of middle 
Villafranchian remains of modern canids alone did not 
push other researchers to abandon the use of the Wolf 
event earlier, mainly emphasizing the increase in abun-
dance of wolf-like canids, rather than their first appear-
ance in the European fossil record (Azzaroli et al., 1988; 
Rook & Torre, 1996). Sardella & Palombo (2007) re-
viewed the concept of the Wolf event and analyzed car-
nivoran faunas of western Europe around the old Plio-
cene-Pleistocene boundary (Aguirre & Pasini, 1985; see 
Pillans, 2004 and Clague, 2006), i.e., prior to the provi-
sions adopted in 2009, following which the base of the 
Pleistocene Epoch/Series was placed at ~2.6 Ma 
(Gibbard & Head, 2010; Gibbard et al., 2010; Head & 
Gibbard, 2015). Sardella & Palombo (2007) argued that, 
considering the carnivorans, the Wolf event includes 
several diachronic bioevents, such as the dispersals of 
different canids, Panthera, and Pachycrocuta. The latter, 
involving the giant hyena Pachycrocuta brevirostris, was 
regarded as the most representative dispersion in Eu-
rope. 

According to Iannucci et al. (2021b), the earliest 
calibrated occurrence of P. brevirostris is likely from 
Fonelas P-1, placed between Feni and Olduvai sub-
chrons, hence between ~2.12 and 1.92 Ma (Arribas et 
al., 2009). Some uncertainty persists, however, consid-
ering that the taxonomic identification is based on few 
elements of the deciduous dentition (Arribas & Garrido, 
2008), with only one tooth actually exceeding the bio-
metric range known for Pliocrocuta perrieri, a species 
also present in the site (Iannucci et al., 2021b). Like 
Fonelas P-1, the fauna of Gerakarou 1 is either referred 
to as middle or late Villafranchian (or as transitional), 
especially due to the cooccurrence of P. brevirostris and 
Gazella (Koufos, 1992; Konidaris et al., 2021). In Italy, 
Gazella is not recorded after the Coste San Giacomo FU 
(Masini et al., 2013; Bellucci & Sardella, 2015) and in-
deed its disappearance was listed among the bioevents 
characterizing the late Villafranchian faunal turnover 
(Azzaroli et al., 1988; Gliozzi et al., 1997). In France, it 
could have become locally extinct even earlier, as it is 
already absent in Senèze and other faunas usually con-
sidered of similar age (Heintz et al., 1974; Palombo & 
Valli, 2004). Assuming that this absence is not due to 
the paucity of the fossil record and/or biased by the envi-
ronmental preferences of Gazella, the association of 

 
 

81 

 
 
“Wolf event” and large mammal dispersal 



   

 

 
 

82 

Gazella and P. brevirostris suggests an age intermedi-
ate between Coste San Giacomo and Olivola FUs. This 
implies a certain degree of subjectivity in referring fau-
nas in which such association is documented, like Fone-
las P-1 and Gerakarou 1, as to middle or late Villafran-
chian. We argue that these faunas could simply be con-
sidered transitional between biochronological units tradi-
tionally recognized (Arribas et al., 2009; Konidaris et al., 
2021). This is probably the best choice to minimize in-
consistency in current research and with previous stud-
ies. 

The fauna of Olivola includes P. brevirostris 
(Ficcarelli & Torre, 1970) but it is not directly dated. 
Paleomagnetic investigations carried out at Poggio Ros-
so, allowed to correlate this latter locality within the Ol-
duvai subchron at ~1.92–1.77 Ma (Napoleone et al., 
2001, 2003; Mazza et al., 2004). The fauna of Poggio 
Rosso has been considered transitional between Olivola 
and Tasso FUs (Gliozzi et al., 1997), or in other terms it 
cannot be unequivocally ascribed to one of the two FUs, 
but in any case provides indirect constraints for both. 

The giant hyena P. brevirostris is documented by 
direct skeletal evidence in ~60 localities across Europe 

(Iannucci et al., 2021b), a number that would likely in-
crease taking into account indirect evidence (e.g., cop-
rolites, gnawing marks) and reports in open taxonomy. 
Actually, roughly coeval early occurrences are known 
from both sides of Eurasia at ~2.0 Ma (Arribas et al., 
2009; Liu et al., 2021; Iannucci et al., 2021b). The giant 
hyena is a species that played an important role as a 
taphonomic agent and has been the subject of much 
research focused on investigating its potential relation-
ships with other carnivorans and early hominins dispers-
ing out of Africa (Turner & Antón, 1996; Palmqvist et al., 
2011; Espigares et al., 2013; Madurell-Malapeira et al., 
2017; Iannucci et al., 2021b). This explains why P. 
brevirostris has been considered a representative spe-
cies of the late Villafranchian faunal turnover (Palombo 
& Sardella, 2007; Martínez-Navarro, 2010). 

The earliest material referred to Pachycrocuta sp. 
is from the Pliocene of East Africa, but African remains 
of Pleistocene age are only known from the South of the 
continent, possibly indicating that Pachycrocuta did not 
survive long in East Africa (Werdelin, 1999; Werdelin & 
Lewis, 2008). Pachycrocuta brevirostris is thought to 
derive from a large-sized population of Pliocrocuta perri-

Fig. 3 - Selected biochronological schemes and mammal ranges proposed over the years (a) and the updated scheme adopted in this 
work (b). Saint Vallier has often been placed in italics to emphasize that it is not an Italian site. Silhouettes modified from: Iurino et al. 
(2022; Canis etruscus); Iannucci et al. (2021, Pachycrocuta brevirostris); PhyloPic, by Zimices (phylopic.org/image/c2d68ebb-50ec-45f4-
8cd1-6cf52ad02286, hippopotamus).  

 
 

Iannucci A. et al. 



   

 

eri, most likely in Asia, where transitional samples are 
known (Qiu et al., 2004; Iannucci et al., 2021b). In west-
ern Eurasia, remains assigned to Pachycrocuta brevi-
rostris and Pliocrocuta perrieri are jointly known from 
Fonelas P-1, Gerakarou 1, and Dmanisi (collectively 
~2.1–1.8 Ma; Koufos, 1992, 2014; Vekua, 1995; Lord-
kipanidze et al., 2007; Arribas et al., 2009; Iannucci et 
al., 2021b). The two species share an overall similar 
morphology in many respects and, especially dealing 
with isolated remains, attributions often relied on bio-
metric comparisons. As the often-evoked “giant” size of 
P. brevirostris suggests, this approach is generally ac-
cepted, but considering the close relationship between 
the two species and the many implications associated to 
the appearance of P. brevirostris, it should be accompa-
nied by some reflection (see Iannucci et al., 2022b, for 
discussion). 

 
6. DISCUSSION AND CONCLUSIONS 

 
Azzaroli (1983) named “Wolf event” the faunal 

turnover associated to the beginning of the late Vil-
lafranchian, coinciding with the Olivola FU. This was 
also roughly coincident with the old (i.e., prior to 2009) 
Plio-Pleistocene boundary at ~1.8 Ma. Initially, this 
event was considered to document the first appearance 
in the European fossil record of Canis etruscus (the 
“wolf”, hence the name) and other large mammals 
(Leptobos etruscus, Sus strozzii, and Pachycrocuta 
brevirostris), but soon modified to include the “massive 
expansion” of the considered species and other bio-
events (Azzaroli et al., 1988, p. 84). The latter adjust-
ment was a good compromise between the need not to 
disregard occurrences of the considered species poten-
tially older than Olivola (but at the time of doubtful chro-
nology or taxonomic attribution, like those from Senèze), 
and the intention of maintaining the reference role of the 
Olivola FU. However, this definition left room for differ-
ent interpretations. It is now clear that several bioevents 
traditionally included in the Wolf event and others that 
have been considered to occur later are conversely 
already documented in middle Villafranchian faunas, 
most notably the arrival in Europe of modern canids, but 
also of S. strozzii and Hippopotamus, among others 
(Fig. 3). 

If a single species has to be taken as representa-
tive of the late Villafranchian faunal turnover, the best 
candidate based on the current evidence is the giant 
hyena Pachycrocuta brevirostris (Sardella & Palombo, 
2007; Palombo et al., 2008; Martínez-Navarro, 2010; 
Iannucci et al., 2021b). Pachycrocuta brevirostris is 
indeed the only species among those initially listed by 
Azzaroli (1983) as part of the Wolf event whose first 
appearance in the European fossil record is still coinci-
dent with the beginning of the late Villafranchian at the 
Olivola FU, and its occurrences are abundant across 
Eurasia. The first appearance in the European fossil 
record of Panthera gombaszoegensis (added in the 
Wolf event as an accompanying species by Azzaroli et 
al., 1988), is also documented at that time. However, 
the faunas Fonelas P-1 (Spain) and Gerakarou 1 
(Greece) contains Pachycrocuta brevirostris in associa-
tion with Gazella, a “typical” middle Villafranchian taxon 

(actually a holdover of even older faunas) that is not 
documented in Italy after the Coste San Giacomo FU 
(last FU of the middle Villafranchian, older than Olivola) 
(Bellucci & Sardella, 2015). Gerakarou 1 includes also 
Panthera gombaszoegensis. Therefore, even the arrival 
of Pachycrocuta brevirostris and Panthera gombaszoe-
gensis somewhat anticipates the chronology previously 
assumed. Moreover, the beginning of the late Villafran-
chian does not correspond to a sharp faunal turnover as 
traditionally envisioned for the Wolf event. Current evi-
dence highlights the rather diachronic nature of large 
mammal dispersal occurred in the late middle and early 
late Villafranchian (late Gelasian, ~2.2–1.8 Ma). In this 
regard, it would be arguably better avoiding naming this 
time-averaged phenomenon after a single species. 

Here, we dedicated space to discuss the “Hippo 
event” (Section 3), as we think that this case eloquently 
speaks for the need of critically evaluating the biochron-
ological, paleoecological, and paleobiogeographical 
significance of each bioevent. Indeed, the presence of 
Hippopotamus in Europe is now attested since ~2.2 Ma, 
documenting an African dispersal of a species linked to 
humid conditions in a context that is generally deemed 
to denote the spread of open-adapted faunal elements 
of mainly Asian affinities. However, this bioevent was 
once considered younger than ~1.8 Ma and placed in 
the Tasso FU (Azzaroli, 1983; Gliozzi et al., 1997). Con-
sequently, the presence of Hippopotamus at Coste San 
Giacomo and Senèze was perceived at odds with a 
middle Villafranchian age and engendered ideas of mix-
ing with substantially younger faunas, which are, howev-
er, not supported by available dating and field evidence 
(Delson et al., 2006; Bellucci et al., 2014; Palombo et 
al., 2017; Paquette et al., 2021). Several other faunas 
have been considered mixed or younger than they really 
are due to the presence of Hippopotamus, including, for 
instance, Castel San Pietro, Bocchignano, and to some 
extent Monte Riccio (e.g., Marra et al., 2018). 

It might be argued that since the arrival of many 
species once considered to indicate the beginning of the 
late Villafranchian is now recognized to be older, then 
the passage between the middle and late Villafranchian 
could be moved back as well. However, such an ap-
proach would be inconsistent with a huge body of previ-
ous research in which the beginning of the late Villafran-
chian is typified by the Olivola FU, and hence it would be 
unavoidably ambiguous. Moreover, while it is true that 
the arrivals of many species traditionally included in the 
Wolf event are now already recorded in middle Villafran-
chian faunas, their “massive expansion” (sensu Azzaroli 
et al., 1988) can still be correlated with the Olivola FU. 
This is the case of Leptobos etruscus, Sus strozzii, and 
not least Canis etruscus. We argue that the earliest oc-
currence of Pachycrocuta brevirostris and Panthera 
gombaszeogensis heralds the late Villafranchian faunal 
turnover, which is then best expressed by the increase 
in the documentation of the other species. To minimize 
inconsistent approaches, “transitional” faunas like Fone-
las P-1 and Gerakarou 1 could be simply referred to as 
such. 

In general, resolving the exact timing of large mam-
mal bioevents in the late Gelasian is complicated by the 
paucity of middle Villafranchian localities, as already 

 
 

83 

 
 
“Wolf event” and large mammal dispersal 



   

 

 
 

84 

pointed out by Azzaroli (1983). Moreover, of the two 
most important large mammal middle Villafranchian 
faunas postdating the Montopoli FU listed in the synthe-
sis of Gliozzi et al. (1997), namely those of Collepardo 
and Coste San Giacomo, only the correlation of the 
latter has been confirmed (Bellucci et al., 2012, 2014; 
Bona et al., 2015). Conversely, although the fauna of 
Collepardo was initially considered close in age to Saint 
Vallier (Gliozzi et al., 1997), it is now recognized as 
markedly older than the French site and referred to the 
Triversa FU (early Villafranchian; Bellucci et al., 2019; 
Iannucci et al., 2022a). On the other hand, a huge num-
ber of localities, more or less abundant in terms of fossil 
remains, including mainly either sporadic (i.e., not com-
ing from systematic excavations) and/or historical find-
ings, have yielded Early Pleistocene large mammal 
faunas. As shown by the recent case of Vigna Nuova 
(Azzarà et al., 2022), proper examinations of these sam-
ples might reveal further middle Villafranchian faunas 
simply not recognized as such. 

 
ACKNOWLEDGEMENTS 

We are thankful to D.A. Iurino and F. Strani for 
discussion and support. Comments and suggestions 
received during the revision of this manuscript from the 
reviewers were appreciated. This work was funded by 
Sapienza University of Rome - Grandi Scavi 2021 
(Grant no: SA12117A87BC3F0A) and Grandi Scavi 
2022 (Grant no: SA1221816893E2AB) to RS; Sapienza 
University of Rome “Progetti per Avvio alla Ricerca - 
Tipo 1 anno 2020” (Grant no: AR120172B7D44B9E) to 
AI; Sapienza University of Rome “Progetti per Avvio alla 
Ricerca - Tipo 2 anno 2022” (Grant no: 
AR222181333C1B88) and “Contributi premiali per i 
ricercatori e assegnisti di ricerca per rafforzarne la con-
dizione professionale e potenziare il sistema della ricer-
ca del Lazio” call proposal of the Lazio Region (DE 
G05411, 05/05/2022) to BM. 

 
REFERENCES  
 
Aguirre E., Pasini G. (1985) - The Pliocene-Pleistocence 

Boundary. Episodes, 8(2), 116-120. 
Adams N.F., Candy I., Schreve D.C. (2022) - An Early 

Pleistocene hippopotamus from Westbury Cave, 
Somerset, England: support for a previously un-
recognized temperate interval in the British Qua-
ternary record. Journal of Quaternary Science, 37
(1), 28-41. 

Argant A. (2004) - Les Carnivores du gisement Pliocène 
final de Saint-Vallier (Drôme, France). Geobios, 
37, S133-S182. 

Arribas A., Garrido G. (2008) - Hiénidos [Pachycrocuta 
brevirostris (Aymard, 1846) y Hyaena brunnea 
Thunberg,1820] del yacimiento de Fonelas P-1 
(cuenca de Guadix, Granada). In: Vertebrados del 
Plioceno superior terminal en el suroeste de Euro-
pa: Fonelas P-1 y el Proyecto Fonelas (Ed. by 
Arribas A.), Instituto Geológico y Minero de Espa-
ña, serie Cuadernos del Museo Geominero 10, 
Madrid, 201-230. 

Athanassiou A. (2022) - The fossil record of continental 
hippopotamids (Mammalia: Artiodactyla: Hippopot-

amidae) in Greece. In: Fossil Vertebrates of 
Greece 2 (Ed. by Vlachos E.), Springer, Cham, 
281-289. 

Azzarà B., Breda M., Cirilli O., Madurell-Malapeira J., 
Ruzza F., Sorbelli L., Tancredi D., Cherin M. 
(2022) - Vigna Nuova: the first Middle Villafran-
chian mammal assemblage from the Valdichiana 
Basin, Perugia (Italy). Bollettino della Società Pa-
leontologica Italiana, 61(2), 223-247. 

Azzaroli A. (1950) - Osservazioni sulla formazione villa-
franchiana di Olivola in Val di Magra. Atti della 
Società Toscana di Scienze Naturali, Memorie 
Serie A, 57, 104-111. 

Azzaroli A. (1970) - Villafranchian correlations based on 
large mammals. Giornale di Geologia, 35, 111-
131. 

Azzaroli A. (1977) - The Villafranchian Stage in Italy and 
the Plio-Pleistocene boundary. Giornale di Geolo-
gia, 41, 61-79. 

Azzaroli A. (1983) - Quaternary mammals and the “end-
Villafranchian” dispersal event - A turning point in 
the history of Eurasia. Palaeogeography, Palaeo-
climatology, Palaeoecology, 44, 117-139. 

Azzaroli A. (1995) - The “Elephant-Equus” and the “End-
Villafranchian” events in Eurasia. In: Palaeoclimate 
and evolution with emphasis on human origins 
(Ed. by Vrba E.S., Denton G.H., Patridge T.C., 
Burckle L.H.), Yale University Press, London, 311-
318. 

Azzaroli A., De Giuli C., Ficcarelli G., Torre D. (1988) - 
Late Pliocene to early mid-Pleistocene mammals 
in Eurasia: faunal succession and dispersal 
events. Palaeogeography, Palaeoclimatology, 
Palaeoecology, 66(1-2), 77-100. 

Bartolini Lucenti S., Rook L. (2016) - A review on the 
Late Villafranchian medium-sized canid Canis 
arnensis based on the evidence from Poggio Ros-
so (Tuscany, Italy). Quaternary Science Reviews, 
151, 58-71. 

Bartolini Lucenti S., Spassov N. (2022) - Cave canem! 
The earliest Canis (Xenocyon) (Canidae, Mamma-
lia) of Europe: Taxonomic affinities and paleoecol-
ogy of the fossil wild dogs. Quaternary Science 
Reviews, 276, 107315. 

Bellucci L., Sardella R. (2015) - The last Antilopini bo-
vids from the Early Pleistocene of Italy. Quaternary 
International, 357, 245-252. 

Bellucci L., Mazzini I., Scardia G., Bruni L., Parenti F., 
Segre A.G., Segre Naldini E., Sardella R. (2012) - 
The site of Coste San Giacomo (Early Pleistocene, 
central Italy): palaeoenvironmental analysis and 
biochronological overview. Quaternary Internation-
al, 267, 30-39. 

Bellucci L., Bona F., Corrado P., Magri D., Mazzini I., 
Parenti F., Scardia G., Sardella R. (2014) - Evi-
dence of late Gelasian dispersal of Africanfauna at 
Coste San Giacomo (Anagni Basin, central Italy): 
early Pleistocene environments and the back-
ground of early human occupationin Europe. Qua-
ternary Science Reviews, 96, 72-85. 

Bellucci L., Biddittu I., Brilli M., Conti J., Germani M., 
Giustini F., Iurino D.A., Mazzini I., Sardella R. 
(2019) - First occurrence of the short-faced bear 

 
 

Iannucci A. et al. 



   

 

Agriotherium (Ursidae, Carnivora) in Italy: bio-
chronological and palaeoenvironmental implica-
tions. Italian Journal of Geosciences, 138(1), 124-
135. 

Böhme M., Spassov N., Majidifard M.R., Gärtner A., 
Kirscher U., Marks M., Dietzel C., Uhlig G., El Atfy 
H., Begun D.R., Winklhofer M. (2021) - Neogene 
hyperaridity in Arabia drove the directions of mam-
malian dispersal between Africa and Eurasia. 
Communications Earth and Environment 2, 85. 

Bona F., Bellucci L., Sardella R. (2015) - The Gelasian 
(Late Villanyan-MN17) diversified micromammal 
assemblage with Mimomys pliocaenicus from 
Coste San Giacomo (Anagni basin, central Italy), 
taxonomy and comparison with selected European 
sites. Hystrix, 26(2), 141-151. 

Bout P. (1960) - Le Villafranchien du Velay et du bassin 
hydrographique moyen et supérior de l'Allier. Cen-
tre National de la Recherche Scientifique, impri-
merie Jeanne d'Arc, Le Puy. 

Brugal J.P., Boudadi-Maligne M. (2011) - Quaternary 
small to large canids in Europe: taxonomic status 
and biochronological contribution. Quaternary 
International, 243(1), 171-182. 

Brugal J.P., Argant A., Boudadi-Maligne M., Crégut-
Bonnoure E., Croitor, R., Fernandez P., Fourvel J.
-B., Fosse P., Guadelli J.L., Labe B., Magniez P., 
Uzunidis A. (2020) - Pleistocene herbivores and 
carnivores from France: An updated overview of 
the literature, sites and taxonomy. Annales de 
Paléontologie, 106, 102384. 

Caloi L., Palombo M.R., Petronio C. (1980) - Resti crani-
ci di Hippopotamus antiquus (= H. major) e Hippo-
potamus amphibius conservati nel Museo di Pa-
leontologia dell'Università di Roma. Geologica 
Romana, 19, 91-119. 

Candy I., Rose J., Lee J. (2006) - A seasonally ‘dry’ 
interglacial climate in eastern England during the 
early Middle Pleistocene: palaeopedological and 
stable isotopic evidence from Pakefield, UK. Bore-
as, 35, 255-265. 

Candy I., Coope G.R., Lee J.R., Parfitt S.A., Preece 
R.C., Rose J., Schreve D.C. (2010) - Pronounced 
warmth during early Middle Pleistocene interglaci-
als: Investigating the Mid-Brunhes Event in the 
British terrestrial sequence. Earth-Science Re-
views, 103(3-4), 183-196. 

Capraro L., Maiorano P. (2023) - Italian GSSPs of the 
Quaternary System. Alpine and Mediterranean 
Quaternary, 36(1). 

 Doi: 10.26382/AMQ.2023.02 
Cassoli P.F., Segre Naldini E. (1984) - Nuovo contributo 

alla conoscenza delle faune villafranchiane e del 
Pleistocene medio del bacino di Anagni 
(Frosinone). In: Atti della XXIV Riunione Scientifi-
ca dell’Istituto Italiano di Preistoria e Protostoria, 
Firenze, 115-118. 

Cherin M., Bertè D.F., Sardella R., Rook L. (2013) - 
Canis etruscus (Canidae, Mammalia) and its role 
in the faunal assemblage from Pantalla (Perugia, 
central Italy): comparison with the Late Villafran-
chian large carnivore guild of Italy. Bollettino della 
Società Paleontologica Italiana, 52, 11-18. 

Cherin M., Bertè D.F., Rook L., Sardella R. (2014) - Re-
defining Canis etruscus (Canidae, Mammalia): a 
new look into the evolutionary history of Early 
Pleistocene dogs resulting from the outstanding 
fossil record from Pantalla (Italy). Journal of Mam-
malian Evolution, 21, 95-110. 

Cherin M., Sorbelli L., Crotti M., Iurino D.A., Sardella R., 
Souron A. (2018) - New material of Sus strozzii 
(Suidae, Mammalia) from the Early Pleistocene of 
Italy and a phylogenetic analysis of suines. Qua-
ternary Science Reviews, 194, 94-115. 

Cherin M., D’Allestro V., Masini F. (2019) - New bovid 
remains from the Early Pleistocene of Umbria 
(Italy) and a reappraisal of Leptobos merlai. Jour-
nal of Mammalian Evolution, 26, 201-224. 

Cherin M., Breda M., Esattore B., Hart V., Turek J., 
Porciello F., Angeli G., Holpin S., Iurino D.A. 
(2022) - A Pleistocene Fight Club revealed by the 
palaeobiological study of the Dama-like deer rec-
ord from Pantalla (Italy). Scientific Reports, 12(1), 
13898. 

Clague J. (2006) - Status of the Quaternary: Your opin-
ion sought. Quaternary International, 144, 99-100. 

Croitor R., Brugal J.P. (2010) - Ecological and evolution-
ary dynamics of the carnivore community in Eu-
rope during the last 3 million years. Quaternary 
International, 212(2), 98-108. 

Cuccu A., Valenciano A., Azanza B., DeMiguel D. 
(2023) - A new lynx mandible from the Early Pleis-
tocene of Spain (La Puebla de Valverde, Teruel) 
and a taxonomical multivariate approach of medi-
um-sized felids. Historical Biology, 35(1), 127-138. 

Cuscani Politi P. (1966) - Resti di ippopotami provenienti 
dalla zona di Chiusi. Atti dell’Accademia dei Fisio-
critici di Siena, 12, 1-44. 

Cuscani Politi P. (1971) - Natura dei sedimenti in cui 
sono stati rinvenuti i resti di ippopotamo (della 
zona di Chiusi) precedentemente studiati. Atti 
dell’Accademia dei Fisiocritici di Siena, 14, 1-13. 

Delson E., Faure M., Guérin C., Aprile L., Argant J., 
Blackwell B.A.B., Debard E., Harcourt-Smith W., 
Martin-Suarez, E., Monguillon A., Parenti F., 
Pastre J.-F., Sen S., Skinner A.R., Swisher C.C., 
Valli A.M.F. (2006) - Franco-American renewed 
research at the late Villafranchian locality of Se-
nèze (Haute-Loire, France). Courier Forschungsin-
stitut Senckenberg, 256, 275-290. 

Ehlers J., Gibbard P.L. (2007) - The extent and chronol-
ogy of Cenozoic Global Glaciation. Quaternary 
International, 164-165, 6-20. 

Espigares M.P., Martínez-Navarro B., Palmqvist P., Ros
-Montoya S., Toro I., Agustí J., Sala R. (2013) - 
Homo vs. Pachycrocuta: Earliest evidence of com-
petition for an elephant carcass between scaven-
gers at Fuente Nueva-3 (Orce, Spain). Quaternary 
International, 295, 113-125. 

Etourneau J., Schneider R., Blanz T., Martinez P. (2010) 
- Intensification of the Walker and Hadley atmos-
pheric circulations during the Pliocene-Pleistocene 
climate transition. Earth and Planetary Science 
Letters, 297(1-2), 103-110. 

Faure M. (1985) - Les hippopotames Quaternaires non-
insulaires d’Europe occidentale. Nouvelles Ar-

 
 

85 

 
 
“Wolf event” and large mammal dispersal 



   

 

chives du Muséum d’Histoire naturelle de Lyon, 
23, 13-79. 

Faure M. (2004) - Le Sus strozzii du Pliocène final de 
Saint-Vallier (Drôme). Geobios, 37, S189-S190. 

Ficcarelli G., Torre D. (1968) - Upper Villafranchian pan-
thers of Tuscany. Palaeontographia Italica, 34, 
173-184. 

Ficcarelli G., Torre D. (1970) - Remarks on the taxono-
my of hyaenids. Paleontographia Italica, 36, 13-
33. 

Fidalgo D., Galli E., Madurell-Malapeira J., Rosas A. 
(2021) - Earliest Pleistocene European hippos: a 
review. Comunicações Geológicas, 108, 69-73. 

Flesche Kleiven H., Jansen E., Fronval T., Smith T.M. 
(2002) - Intensification of Northern Hemisphere 
glaciations in the circum Atlantic region (3.5-2.4 
Ma)-ice-rafted detritus evidence. Palaeogeogra-
phy, Palaeoclimatology, Palaeoecology, 184(3-4), 
213-223. 

Florindo F., Marra F., Angelucci D.E., Biddittu I., Bruni 
L., Florindo F., Gaeta M., Guillou H., Jicha B., 
Macrì P., Morigi C., Nomade S., Parenti F., Perei-
ra A., Grimaldi S. (2021) - Environmental evolu-
tion, faunal and human occupation since 2 Ma in 
the Anagni basin, central Italy. Scientific Reports, 
11, 7056. 

Fondi R. (1972) - Fauna cromeriana della Montagnola 
senese. Palaeontographia Italica, 68, 1-27. 

Gibbard P.L., Smith A.G., Zalasiewicz J.A., Barry T.L., 
Cantrill D., Coe A.L., Cope J.C.W., Gale A.S., 
Gregory F.J., Powell J.H., Rawson P.F., Stone P., 
Waters C.N. (2005) - What status for the Quater-
nary? Boreas, 34, 1-6. 

Gibbard P.L., Head M.J. (2010) - The newly-ratified 
definition of the Quaternary System/Period and 
redefinition of the Pleistocene Series/Epoch, and 
comparison of proposals advanced prior to formal 
ratification. Episodes, 33, 152158. 

Gibbard P.L., Head M.J., Walker M. (2010) - Subcom-
mission on Quaternary Stratigraphy. Formal ratifi-
cation of the Quaternary System/Period and the 
Pleistocene Series/Epoch with a base at 2.58 Ma. 
Journal of Quaternary Science, 25, 96102. 

Gibbard P.L., Head M.J. (2020) - The Quaternary Peri-
od. In: Geologic Time Scale 2020 (Ed. by Grad-
stein F.M., Ogg J.G., Schmitz M.D., Ogg G.M.), 
Elsevier, Amsterdam,1217-1255. 

Girotti O., Capasso Barbato L., Esu D., Gliozzi E., 
Kotsakis T., Martinetto E., Petronio C., Sardella 
R., Squazzini E. (2003) - The section of Torre 
Picchio (Terni, Umbria, Central Italy): a Villafran-
chian site rich in mammals, molluscs, ostracods 
and plants. Rivista Italiana di Paleontologia e Stra-
tigrafia, 109, 77-98. 

Gliozzi E., Abbazzi L., Argenti P., Azzaroli A., Caloi L., 
Capasso Barbato L., Di Stefano G., Esu D., Ficca-
relli G., Girotti O., Kotsakis T., Masini F., Mazza 
P., Mezzabotta C., Palombo M.R., Petronio C., 
Rook L., Sala B., Sardella R., Zanalda E., Torre D. 
(1997) - Biochronology of selected mammals, 
molluscs and ostracods from the middle Pliocene 
to the late Pleistocene in Italy. The state of the art. 
Riv. Ital. Paleontol. Stratigr., 1997, 103, 369-388. 

Gradstein F.M., Agterberg F.P., Brower J.C., 
Schwarzacher W. (1985) - Quantitative Stratigra-
phy. Reidel Publication Company and Unesco, 
Dordrecht and Paris.  

Guérin C. (2005) - Préface. In: Les grands Mammifères 
fossils du Velay. Les collections paléontologiques 
du Plio-Pléistocène du mesée Crozatier, le Puy-en
-Velay (Ed. by Lacombat F.), Phil’ Print, Yssin-
geaux, 11-13. 

Guérin C., Faure M., Argant A., Argant J., Crégut-
Bonnoure É., Debard É., Delson E., Eisenmann V., 
Hugueney M., Limondin-Lozouet N., Martín-
Suárez E., Mein P., Mourer-Chauviré, Parenti F., 
Pastre J.-F., Sen S., Valli A. (2004) - Le gisement 
pliocène supérieur de Saint-Vallier (Drôme, 
France): synthèse biostratigraphique et paléoé-
cologique. Geobios, 37, S349-S360. 

Head M.J., Gibbard P.L. (2015) - Formal subdivision of 
the Quaternary System/Period: past, present, and 
future. Quaternary International, 383, 4-35. 

Heintz E., Guérin C., Martin R., Prat F. (1974) - Princi-
paux gisements villafranchiens de France: listes 
fauniques et biostratigraphie. Mémoires du Bureau 
de Recherches géologiques et minières, 78(1), 
169-182. 

Iannucci A. (2023) - Sus strozzii (Suidae, Mammalia) 
from the historical locality of Quercia (Early Pleis-
tocene, Italy). Geobios, 77. 

 Doi: 10.1016/j.geobios.2023.03.001  
Iannucci A., Sardella R. (2023) - What does the 

“Elephant-Equus” event mean today ? Reflections 
on mammal dispersal events around the Pliocene-
Pleistocene boundary and the flexible ambiguity of 
biochronology. Quaternary, 6(1), 16.  

Iannucci A., Gasparik M., Sardella R. (2020) - First re-
port of Sus strozzii (Suidae, Mammalia) from the 
Early Pleistocene of Hungary (Dunaalmás) and 
species distinction based on deciduous teeth. The 
Science of Nature, 107(1), 5. 

Iannucci A., Cherin M., Sorbelli L., Sardella R. (2021a) - 
Suidae transition at the Miocene-Pliocene bounda-
ry: a reassessment of the taxonomy and chronolo-
gy of Propotamochoerus provincialis. Journal of 
Mammalian Evolution, 28(2), 323-335. 

Iannucci A., Mecozzi B., Sardella R., Iurino D.A. (2021b) 
- The extinction of the giant hyena Pachycrocuta 
brevirostris and a reappraisal of the Epivillafran-
chian and Galerian Hyaenidae in Europe: Faunal 
turnover during the Early–Middle Pleistocene Tran-
sition. Quaternary Science Reviews, 272, 107240. 

Iannucci A., Bellucci L., Conti J., Mazzini I., Mecozzi B., 
Sardella R., Iurino D.A. (2022a) - Neurocranial 
anatomy of Sus arvernensis (Suidae, Mammalia) 
from Collepardo (Early Villafranchian; central Italy): 
taxonomic and biochronological implications. His-
torical Biology, 34(1), 108-120. 

Iannucci A., Cherin M., Sardella R. (2022b). The lost 
hyena from Paciano (Umbria, Italy) reconsidered. 
Alpine and Mediterranean Quaternary, 35(2), 105-
117. 

Iurino D.A., Mecozzi B., Iannucci A., Moscarella A., 
Strani F., Bona F., Gaeta M., Sardella R. (2022) - 
A Middle Pleistocene wolf from central Italy pro-

 
 

86 

 
 

Iannucci A. et al. 



   

 

vides insights on the first occurrence of Canis 
lupus in Europe. Scientific Reports, 12(1), 2882. 

Jung J. (1946) - Géologie de l’Auvergne et de ses con-
fins Bourbonnais et Limousins. Collection Mé-
moires pour servir à l‘explication de la carte 
géologique détaillée de la France. Imprimerie Na-
tionale, Paris. 

Kahlke R.-D. (1989) - Die unterpleistozänen Hippopota-
mus-Reste von Würzburg-Schalksberg. Quartär, 
39, 67-94. 

Kahlke R.‐D. (2001) - Schädelreste von Hippopotamus 
aus dem Unterpleistozän von Untermassfeld. In: 
Das Pleistozän von Untermaßfeld bei Meiningen 
(Thüringen), Teil 2 (Ed. by Kahlke R.‐D.), Habelt 
Verlag, Bonn, 483-500. 

Kahlke R.-D., García N., Kostopoulos D.S., Lacombat 
F., Lister A.M., Mazza P.P., Spassov N., Titov V.V. 
(2011) - Western Palaearctic palaeoenvironmental 
conditions during the Early and early Middle Pleis-
tocene inferred from large mammal communities, 
and implications for hominin dispersal in Europe. 
Quaternary Science Reviews, 30(11-12), 1368-
1395. 

Konidaris G.E., Kostopoulos D.S., Maron M., Schaller 
M., Ehlers T.A., Aidona E., Marini M., Tourloukis 
V., Muttoni G., Koufos G.D., Harvati K. (2021) - 
Dating of the lower Pleistocene vertebrate site of 
Tsiotra Vryssi (Mygdonia basin, Greece): biochro-
nology, magnetostratigraphy, and cosmogenic 
radionuclides. Quaternary, 4, 1. 

Kotsakis T. (1988) - Biostratigraphy of Plio-Pleistocene 
arvicolids (rodents) of Italy. Modern Geology, 13, 
163-175. 

Koufos G.D. (1992) - The Pleistocene carnivores of the 
Mygdonia basin (Macedonia, Greece). Annales de 
Paléontologie, 78, 205-257. 

Koufos G.D. (2014) - The Villafranchian carnivoran guild 
of Greece: implications for the fauna, biochronolo-
gy and paleoecology. Integrative zoology, 9(4), 
444-460. 

Kurtén B., Crusafont Pairó M. (1977) - Villafranchian 
carnivores (Mammalia) from La Puebla de 
Valverde (Teruel, Spain). Commentationes Biolog-
icae, 85, 1-39. 

Lacombat F., Abbazzi L., Ferretti M.P., Martínez-
Navarro B., Moullé P.-E., Palombo M.R., Rook L., 
Turner A., Valli A.-M.F. (2008) - New data on the 
Early Villafranchian fauna from Vialette (Haute-
Loire, France) based on the collection of the 
Crozatier Museum (Le Puy-en-Velay, Haute-Loire, 
France). Quaternary International, 179(1), 64-71. 

Leonardi P. (1947) - L’ippopotamo del Valdarno. Palae-
ontographia Italica, 43, 17-44. 

Lindsay E. (2003) - Chronostratigraphy, biochronology, 
datum events, Land Mammal Ages, stage of evo-
lution, and appearance event ordination. Bulletin 
of the American Museum of Natural History, 279, 
212-230. 

Liu J., Liu J., Zhang H., Wagner J., Jiangzuo Q., Song 
Y., Liu S., Wang Y., Jin C. (2021) - The giant short
-faced hyena Pachycrocuta brevirostris 
(Mammalia, Carnivora, Hyaenidae) from Northeast 
Asia: A reinterpretation of subspecies differentia-

tion and intercontinental dispersal. Quaternary 
International 577, 29-51. 

Lordkipanidze D., Jashashvili T., Vekua A., De Léon 
M.S.P., Zollikofer C.P., Rightmire G.P., Pontzer H., 
Ferring R., Oms O., Tappen M., Bukhsianidze M., 
Agustí J., Kahlke R., Kiladze G., Martínez-Navarro 
B., Mouskhelishvili A., Nioradze M., Rook L. (2007) 
- Postcranial evidence from early Homo from 
Dmanisi, Georgia. Nature, 449, 305-310. 

Lydekker R. (1885) - Catalogue of the fossil Mammalia 
in the British Museum (Natural History) Cromwell 
Road, S.W. Part I. Containing the orders Primates, 
Chiroptera, Insectivora, Carnivora, and Rodentia. 
Taylor and Francis, London. 

Madurell-Malapeira J., Ros-Montoya S., Espigares M.P., 
Alba D.M., Aurell-Garrido J. (2014) - Villafranchian 
large mammals from the Iberian Peninsula: paleo-
biogeography, paleoecology and dispersal events. 
Journal of Iberian Geology, 40(1),167-178. 

Madurell-Malapeira J., Alba D.M., Espigares M.P., Vi-
nuesa V., Palmqvist P., Martínez-Navarro B., 
Moyà-Solà S. (2017) - Were large carnivorans and 
great climatic shifts limiting factors for hominin 
dispersals? Evidence of the activity of Pachycrocu-
ta brevirostris during the Mid-Pleistocene Revolu-
tion in the Vallparadís Section (Vallès-Penedès 
Basin, Iberian Peninsula). Quaternary Internation-
al, 431, 42-52. 

Marciszak A., Kropczyk A., Gornig W., Kot M., 
Nadachowski A., Lipecki G. (2023) - History of 
Polish Canidae (Carnivora, Mammalia) and their 
biochronological implications on the Eurasian 
background. Genes, 14(3), 539. 

Marra F., Petronio C., Salari L., Florindo F., Giaccio B., 
Sottili G. (2018) - A review of the Villafranchian 
fossiliferous sites of Latium in the framework of the 
geodynamic setting and paleogeographic evolution 
of the Tyrrhenian Sea margin of central Italy. Qua-
ternary Science Reviews, 191, 299-317. 

Martin R. (1973) - Trois nouvelles espèces de Caninae 
(Canidae, Carnivora) des gisements plio-
villafranchiens d'Europe. Travaux et Documents 
des Laboratoires de Géologie de Lyon, 57(1), 87-
96. 

Martínez-Navarro B. (2004) - Hippos, pigs, bovids, saber
-toothed tigers, monkeys, and hominids, dispersals 
through the Levantine corridor during late Pliocene 
and early Pleistocene times. In: Human Paleoecol-
ogy in the Levantine Corridor (Ed. by Goren-Inbar 
N., Speth J.D.), Oxbow Books, Oxford, 37-52. 

Martínez-Navarro, B. (2010) - Early Pleistocene faunas 
and hominin dispersals. In: Out of Africa I: The 
First Colonization of Eurasia (Ed. by Fleagle J.G., 
Shea J.J., Grine F.E., Baden A.L., Leakey R.E.), 
Vertebrate Paleobiology and Paleoanthropology. 
Springer Netherlands, Dordrecht, 207-224. 

Martínez-Navarro B., Madurell-Malapeira J., Ros-
Montoya S., Espigares M.P., Medin T., Hortola P., 
Palmqvist P. (2015) - The Epivillafranchian and the 
arrival of pigs into Europe. Quaternary Internation-
al, 389, 131-138. 

Martino R., Pandolfi L. (2022) - The Quaternary Hippo-
potamus records from Italy. Historical Biology, 34

 
 

87 

 
 
“Wolf event” and large mammal dispersal 



   

 

(7), 1146-1156. 
Masini F. (1989) - I bovini villafranchiani dell’Italia. Un-

published PhD thesis, Università consorziate: Mo-
dena, Bologna, Firenze, Roma. 

Masini F., Torre D. (1990) - Large mammal dispersal 
events at the beginning of the late Villafranchian. 
In: European Neogene Mammal Chronology (Ed. 
by Lindsay E.H., Fahlbusch V., Mein P.), Plenum 
Press, New York, 131-138. 

Masini F., Palombo M.R., Rozzi R. (2013) - A reapprais-
al of the Early to Middle Pleistocene Italian Bo-
vidae. Quaternary International, 288, 45-62. 

Maxia C. (1949) - Resti di Mammiferi rinvenuti nella 
miniera di lignite di San Pietro (Sabina). La Ricer-
ca Scientifica, 19, 346-347. 

Mazza P. (1991) - Interrelations between Pleistocene 
hippopotami of Europe and Africa. Bolletino della 
Società Paleontologica Italiana, 30(2), 153-186. 

Mazza P. (1995) - New evidence on the Pleistocene 
hippopotamuses of western Europe. Geologica 
Romana, 31, 61-241. 

Mazza P., Rustioni M. (1994) - The fossil bear from 
Senèze (Southern France). Rendiconti Lincei, 5, 
17-26. 

Mazza P.P., Bertini A. (2013) - Were Pleistocene hippo-
potamuses exposed to climate‐driven body size 
changes? Boreas, 42(1), 194-209. 

Mazza P.P., Bertini A., Magi M. (2004) - The late Plio-
cene site of Poggio Rosso (central Italy): taphono-
my and paleoenvironment. Palaios, 19(3), 227-
248. 

Mazzini I., Paccara P., Petronio C., Sardella R. (2000) - 
Geological evolution and biochronological evi-
dences of the Monte Riccio section (Tarquinia, 
Central Italy). Rivista Italiana di Paleontologia e 
Stratigrafia, 106(2), 247-256. 

Mecozzi B., Iannucci A., Mancini M., Sardella R. (2021) 
- Redefining Ponte Molle (Rome, central Italy): an 
important locality for Middle Pleistocene mammal 
assemblages of Europe. Alpine and Mediterrane-
an Quaternary, 34(1), 131-154. 

Mein P., Moissenet E., Truc G. (1978) - Les formations 
continentales du Néogène supérieur des vallées 
du Jucar et du Cabriel au NE d’Albacete 
(Espagne), biostratigraphie et environment. 
Travaux et Documents des Laboratoires de Géolo-
gie de Lyon, 72, 99-147. 

Meli R. (1882) - Sulla zona dei fori lasciati dai litodomi 
pliocenici nella calcaria giurese di Fara Sabina. 
Bollettino del Regio Comitato Geologico d’Italia, 
13, 147-155. 

Napoleone G., Albanielli A., Azzaroli A., Mazzini M. 
(2001) - The Poggio Rosso locality calibrated to 
the end-Pliocene and its significance for dating the 
late Villafranchian mammal faunas of the Upper 
Valdarno, Central Italy. Rivista Italiana di Paleon-
tologia e Stratigrafia, 107, 287-296. 

Napoleone G., Albianelli A., Azzaroli A., Bertini A., Magi 
M., Mazzini M. (2003) - Calibration of the Upper 
Valdarno basin to the Plio-Pleistocene for correla-
ting the Apennine continental sequences. Il Qua-
ternario, 16(1 bis), 131-166. 

Nesti F. (1820) - Descrizione osteologica dell'ippopota-

mo maggiore fossile dei terreni mobili del Valdarno 
superiore in Toscana. Atti della Società Italiana 
delle Scienze, Modena, 28, 1-23. 

Nomade S., Pastre J.-F., Guillou H., Faure M., Guérin 
C., Delson E., Debard E., Voinchet P., Messager 
E. (2014) - 40Ar/39Ar constraints on some French 
landmark Late Pliocene to Early Pleistocene large 
mammalian paleofaunas: Paleoenvironmental and 
paleoecological implications. Quaternary Geochro-
nology, 21, 2-15. 

Palmqvist P., Gröcke D.R., Arribas A., Fariña R.A. 
(2003) - Paleoecological reconstruction of a lower 
Pleistocene large mammal community using bio-
geochemical (δ13C, δ15N, δ18O, Sr: Zn) and eco-
morphological approaches. Paleobiology, 29(2), 
205-229. 

Palmqvist P., Pérez-Claros J.A., Janis C.M., Figueirido 
B., Torregrosa V., Grocke D.R. (2008) - Biogeo-
chemical and ecomorphological inferences on prey 
selection and resource partitioning among mam-
malian carnivores in an early Pleistocene commu-
nity. Palaios, 23(11), 724-737. 

Palmqvist P., Martínez-Navarro B., Pérez-Claros J.A., 
Torregrosa V., Figueirido B., Jiménez-Arenas J.M., 
Patrocinio Espigares M., Ros-Montoya S., De 
Renzi M. (2011) - The giant hyena Pachycrocuta 
brevirostris: Modelling the bone-cracking behavior 
of an extinct carnivore. Quaternary International 
243, 61-79. 

Palombo M.R. (2009) - Biochronology of terrestrial 
mammals and Quaternary subdivisions: a case 
study of large mammals from the Italian peninsula. 
Il Quaternario, 22, 291-306. 

Palombo M.R., Valli A.M.F. (2004) - Remarks on the 
biochronology of mammalian faunal complexes 
from the Pliocene to the Middle Pleistocene in 
France. Geologica Romana, 37(2003-2004), 145-
163. 

Palombo M.R., Sardella R. (2007) - Biochronology ver-
sus biostratigraphy: a true dilemma or a false trou-
ble? The example of the Plio-Pleistocene large 
mammalian faunas from the Italian peninsula. 
Quaternary International, 160, 30-42. 

Palombo M.R., Alberdi M.T. (2017) - Light and shadows 
in the evolution of South European stenonoid hors-
es. Fossil Imprint, 73, 115-140. 

Palombo M.R., Sardella R., Novelli M. (2008) - Car-
nivora dispersal in Western Mediterranean during 
the last 2.6 Ma. Quaternary International, 179, 176
-189. 

Palombo M.R., Alberdi M.T., Bellucci L., Sardella R. 
(2017) - An intriguing middle-sized horse from 
Coste San Giacomo (Anagni Basin, central Italy). 
Quaternary Research, 87(2), 347-362. 

Pandolfi L., Petronio C. (2015) - A brief review of the 
occurrences of Pleistocene Hippopotamus 
(Mammalia, Hippopotamidae) in Italy. Geologia 
Croatica, 68(3), 313-319. 

Pandolfi L., Spadi M., Martinetto E., Kotsakis T., Esu D. 
(2017) - New data on the lower Pleistocene 
(Gelasian) lignite beds of Castel San Pietro (Rieti, 
Central Italy). Rivista Italiana di Paleontologia e 
Stratigrafia, 123(2), 335-346. 

 
 

88 

 
 

Iannucci A. et al. 



   

 

Paquette J.L., Médard E., Poidevin J.L., Barbet P. 
(2021) - Precise dating of middle to late Villafran-
chian mammalian paleofaunae from the Upper 
Allier River valley (French Massif Central) using U-
Pb geochronology on volcanic zircons. Quaternary 
Geochronology, 65, 101198. 

Pastre J.F., Debard E., Nomade S., Guillou H., Faure 
M., Guérin C., Delson E. (2015) - Nouvelles don-
nées géologiques et téphrochronologiques sur le 
gisement paléontologique du maar de Senèze 
(Pléistocène inférieur, Massif Central, France). 
Quaternaire. Revue de l'Association française 
pour l'étude du Quaternaire, 26(3), 225-244. 

Pillans B. (2004) - Proposal to redefine the Quaternary. 
Episodes, 27(2), 127. 

Poidevin J.L., Cantagrel J.M., G.U.E.R.P.A. (1984) - Un 
site unique du Plio-Pleistocene en Europe: le plu-
teau de Perrier (Puy-de-Dôme) - Confrontation 
des donneés volcanologiques, stratigraphiques et 
paléontologiques. Revue des Sciences naturelles 
d'Auvergne, 50, 87-95. 

Psarianos P. (1954) - Über das Vorkommen von Hippo-
potamus auf Kephallinia (Griechenland). Praktiká 
Akademías Athinón, 28, 408-412. 

Qiu Z., Deng T., Wang B. (2004) - Early Pleistocene 
mammalian fauna from Longdan, Dongxiang, 
Gansu, Chian. Palaeontologica Sinica N. Ser. C, 
27, 1-198. 

Reimann K.C., Strauch F. (2008) - Ein Hippopotamus-
Schädel aus dem Pliozän von Elis (Peloponnes, 
Griechenland). Neues Jahrbuch für Geologie und 
Paläontologie-Abhandlungen, 249(2), 203-222. 

Repenning C.A. (1967) - Nearctic mammalian dispersal 
in late Cenozoic. In: The Bering Land Bridge (Ed. 
by Hopkins D.M.), Stanford University Press, Palo 
Alto, 208-311. 

Repenning C.A. (1980) - Faunal exchanges between 
Siberia and North America. Canadian Journal of 
Anthropology, 1, 37-44. 

Reumer, J.W.F., Piskoulis P. (2017) - A specimen of 
Canis cf. C. etruscus (Mammalia, Carnivora) from 
the Middle Villafranchian of the Oosterschelde. 
Netherlands Journal of Geosciences, 96(1), 3-7. 

Rook L., Torre D. (1996) - The wolf-event in western 
Europe and the beginning of the Late Villafran-
chian. Neues Jahrbuch für Geologie und Paläon-
tologie-Monatshefte, 1996(8), 495-501. 

Rook L., Martínez-Navarro B. (2010) - Villafranchian: 
the long story of a Plio-Pleistocene European 
large mammal biochronologic unit. Quaternary 
International, 219, 134-144. 

Rook L., Croitor R., Delfino M., Ferretti M.P., Gallai G., 
Pavia M. (2013) - The Upper Valdarno Plio-
Pleistocene vertebrate record: an historical over-
view, with notes on palaeobiology and stratigraph-
ic significance of some important taxa. Italian Jour-
nal of Geosciences, 132(1), 104-125. 

Russo Ermolli E., Sardella R., Di Maio G., Petronio C., 
Santangelo N. (2010) - Pollen and mammals from 
the late Early Pleistocene site of Saticula 
(Sant'Agata de'Goti, Benevento, Italy). Quaternary 
International, 225(1), 128-137. 

Sardella R. (2012) - Evidence of Hippopotamus remains 

in the middle Villafranchian faunal assemblages of 
Anagni basin (Central Italy): evidence for an early 
dispersal of the genus in Europe. Quaternary Inter-
national, 279-280, 427. 

Sardella R., Palombo M.R. (2007) - The Pliocene-
Pleistocene boundary: which significance for the 
so called “Wolf Event”? Evidences from Western 
Europe. Quaternaire, 18, 65-71. 

Sardella R., Bellucci L., Bona F., Cherin M., Iurino D.A., 
Rook L. (2018) - Before and after the earliest Ho-
mo dispersal in Europe: Evidence from the early 
Pleistocene sites of the Italian Peninsula. Comptes 
Rendus Palevol, 17(4-5), 287-295. 

Scager D.J., Ahrens H.J., Dieleman F.E., Van den Hoek 
Ostende L.W., De Vos J., Reumer J.W.F. (2017) - 
The Kor & Bot collection revisited, with a biostrati-
graphic interpretation of the early Pleistocene 
Oosterschelde Fauna (Oosterschelde estuary, the 
Netherlands). Deinsea, 17, 16-31. 

Schaub S. (1943) - Die oberpliocaene Säugetierfauna 
von Senèze (Haute-Loire) und ihre verbreitung-
sgeschichtliche Stellung. Eclogae Geologicae Hel-
vetiae, 36, 270-289. 

Shackleton N.J. (1995) - New data on the evolution of 
Pliocene climatic variability. In: Palaeoclimate and 
evolution with emphasis on human origins (Ed. by 
Vrba E.S., Denton G.H., Patridge T.C., Burckle 
L.H.), Yale University Press, London, 242-248. 

Sinusía C., Pueyo E.L., Azanza B., Pocoví A. (2004) - 
Datación magnetoestratigráfica del yacimiento 
paleontológico de la Puebla de Valverde (Teruel). 
Geotemas, 6(4), 339-342. 

Sorbelli L., Cherin M., Kostopoulos D.S., Sardella R., 
Mecozzi B., Plotnikov V., Prat-Vericat M., Azzarà 
B., Bartolini-Lucenti S., Madurell-Malapeira, J. 
(2023) - Earliest bison dispersal in Western Pa-
learctic: Insights from the Eobison record from 
Pietrafitta (Early Pleistocene, central Italy). Quater-
nary Science Reviews, 301, 107923. 

Strani F., DeMiguel D., Sardella R., Bellucci L. (2015) - 
Paleoenvironments and climatic changes in the 
Italian Peninsula during the Early Pleistocene: 
evidence from dental wear patterns of the ungulate 
community of Coste San Giacomo. Quaternary 
Science Reviews, 121, 28-35. 

Strani F., DeMiguel D., Sardella R., Bellucci L. (2018) - 
Resource and niche differentiation mechanisms by 
sympatric Early Pleistocene ungulates: the case 
study of Coste San Giacomo. Quaternary Interna-
tional, 481, 157-163. 

Strani F., Bellucci L., Iannucci A., Iurino D.A., Mecozzi 
B., Sardella R. (2022) - Palaeoenvironments of the 
MIS 15 site of Cava di Breccia-Casal Selce 2 
(central Italian Peninsula) and niche occupation of 
fossil ungulates during Middle Pleistocene intergla-
cials. Historical Biology, 34(3), 555-565. 

Symeonidis N.K., Theodorou G.E. (1986) - New locali-
ties with fossil Hippopotamus in Northwestern Pel-
oponnese. Annales Géologiques des Pays Hel-
léniques, 33, 51-67. 

Thenius E. (1955) - Hippopotamus aus dem Astien von 
Elis (Peloponnes). Annales Géologiques. des Pays 
Helléniques, 6, 206-212. 

 
 

89 

 
 
“Wolf event” and large mammal dispersal 



   

 

Torre D. (1967) - I cani villafranchiani della Toscana. 
Palaeontographia Italica, 63, 113-138. 

Torre D. (1979) - The Ruscinian and the Villafranchian 
dogs of Europe. Bollettino della Società Paleonto-
logica Italiana, 18(2), 162-165. 

Torre D., Ficcarelli G., Masini F., Rook L., Sala B. 
(1992) - Mammal dispersal events in the early 
Pleistocene of Western Europe. Courier For-
schungsinstitut Senckenberg, 153, 51-58. 

Tuccimei G. (1889a) - Alcune recenti osservazioni sul 
Villafranchiano della Sabina Bollettino della Socie-
tà Geologica Italiana, 8, 566-568. 

Tuccimei G. (1889b) - Il Villafranchiano nelle valli sabine 
e I suoi fossili caratteristici. Bollettino della Società 
Geologica Italiana, 8, 95-131. 

Tuccimei G. (1891) - Alcuni mammiferi fossili delle pro-
vicie Umbra e Romana. Memorie della Pontificia 
Accademia dei Nuovi Lincei, 7, 89-152. 

Tuccimei G. (1893) - Resti di Arvicola nel Pliocene lacu-
stre della Sabina. Memorie della Pontificia Acca-
demia dei Nuovi Lincei, 9, 35-45. 

Tuccimei G. (1898) - Sopra alcuni cervi pliocenici della 
Sabina e della Provincia di Roma. Memorie della 
Pontificia Accademia dei Nuovi Lincei, 14, 33-55 

Turner A., Antón M. (1996) - The giant hyaena Pa-
chycrocuta brevirostris (Mammalia, Carnivora, 
Hyaenidae). Geobios, 29, 455-468. 

Van der Made J. (2005) - Le Pliocène Moyen, le Villa-
franchien inférieur. La faune du Velay vers 3 mil-
lions d’années. - cf. “Microstonyx” major. In: Les 
grands Mammifères fossils du Velay. Les collec-
tions paléontologiques du Plio-Pléistocène du 
mesée Crozatier, le Puy-en-Velay (Ed. by 
Lacombat F.), Phil’ Print, Yssingeaux, 58-59. 

Van der Made J., Moullé P.E. (2005) - Le Pliocène Moy-
en, le Villafranchien inférieur. La faune du Velay 
vers 3 millions d’années. In: Les grands Mam-
mifères fossils du Velay. Les collections paléon-
tologiques du Plio-Pléistocène du mesée Croza-
tier, le Puy-en-Velay (Ed. by Lacombat F.), Phil’ 
Print, Yssingeaux, 56-57. 

Van der Made J., Stefaniak K., Marciszak A. (2014) - 
The Polish fossil record of the wolf Canis and the 
deer Alces, Capreolus, Megaloceros, Dama and 
Cervus in an evolutionary perspective. Quaternary 
International, 326, 406-430. 

Van der Made J., Rosell J., Blasco R. (2017) - Faunas 
from Atapuerca at the Early-Middle Pleistocene 
limit: The ungulates from level TD8 in the context 
of climatic change. Quaternary International, 433, 
296-346. 

Vekua A. (1995) - Die Wirbeltierfauna des Villafranchium 
von Dmanisi und ihre biostratigraphische 
Bedeutung. Jahrbuch des Römisch-Germanischen 
Zentralmuseums Mainz, 42, 77-180. 

Viret J. (1954) - Le loess à bancs durcis de Saint-Vallier 
(Drôme) et sa faune de mammifères vil-
lafranchiens. Nouvelles archives du Muséum 
d’histoire naturelle de Lyon, 4, 1-200. 

Werdelin L. (1999) - Pachycrocuta (hyaenids) from the 
Pliocene of East Africa. Paläontologische 
Zeitschrift, 73, 157-165. 

Werdelin L., Lewis M.E. (2008) - New species of Crocu-
ta from the early Pliocene of Kenya, with an over-
view of early Pliocene hyenas of eastern Africa. 
Journal of Vertebrate Paleontology, 28, 1162-
1170. 

 
 
 

Ms. received: February 10, 2023 Revised: March 18, 2023 
Accepted: March 19, 2023 Available online: March 29, 2023 

 
 

90 

 
 

Iannucci A. et al.