REDEFINING PONTE MOLLE (ROME, CENTRAL ITALY):  
AN IMPORTANT LOCALITY FOR MIDDLE PLEISTOCENE  

MAMMAL ASSEMBLAGES OF EUROPE. 
 

Beniamino Mecozzi1,2, Alessio Iannucci1,2, Marco Mancini3, Raffaele Sardella1,2 
 

1 Dipartimento di Scienze della Terra, Sapienza Università di Roma, Italy 
2 PaleoFactory, Sapienza Università di Roma, Italy 

3 Istituto di Geologia Ambientale e Geoingegneria, CNR, Italy. 
 

Corresponding author: B. Mecozzi <beniamino.mecozzi@uniroma1.it>  

 
ABSTRACT: In this work, the Middle Pleistocene mammal assemblage from Ponte Molle, a historical locality of the urban area of 
Rome, has been revised together with a review of the stratigraphical succession of the deposit. This allows us to reconstruct the 
provenance of the fossil material and to provide chronological constrains trough the correlation with the lithostatigraphic and syn-
themic units of the national geological cartography and the geochronologically-constrained aggradational units of the Paleo-Tiber 
reported in literature. The paleontological study together with the geological and stratigraphical review allow us to redefine the 
Ponte Molle deposit and its Middle Pleistocene faunal assemblage. In its new look, the age of the faunal assemblage from Ponte 
Molle could be referred to a time span ranging from 550 ka to 450 ka.  
 
Keywords: Fossil vertebrates, Biochronology, Galerian, Palaeoecology. 

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

 
 

Alpine and Mediterranean Quaternary, 34 (1), 2021, 131-154 

1. THE URBAN AREA OF ROME 
 
The city of Rome lays in one of the most important 

sedimentary basins of Italy for the paleontological re-
search, considering the very high number of fossilifer-
ous findings. The city is crossed by the Tiber River, 
whose evolution is the result of complex geological pro-
cesses including tectonic, volcanism and glacio-eustatic 
fluctuations (Conato et al., 1980; Milli; 1997; Karner et 
al., 2001; Giordano et al., 2003; Funiciello & Giordano, 
2008a, b). The Tiber River fluvio-deltaic evolution (the 
“Paleo-Tiber” system), which commenced during the 
latest Early-earliest Middle Pleistocene (1.1-0.7 Ma; Milli 
et al., 2016), mainly includes sedimentation linked to 
sea-level rises during the Pleistocene glacial termina-
tions (Pandolfi & Marra, 2015). What follows is a discon-
tinuous stratigraphic record constituted by twelve low 
rank and high frequency (80-100 ka spaced) deposition-
al sequences, complexly stacked within the basin and 
deposited by the start of MIS 32-31 (Marine Isotopic 
Stage) (late Early Pleistocene). In fact, forced by the 
eustatic and relative falls of the sea level, the Tiber Riv-
er cyclically excavated the fluvial valleys and coastal 
plain incisions into the bedrock, later filled during the 
following transgressive and highstand phases with de-
posits often including pyroclastic products. These were 
sourced from the Alban Hills and Sabatini Mounts Vol-
canic Districts, with main activity spanning the 600-250 
ka time interval (De Rita et al., 1993; Marra et al., 2009, 
2014; Funiciello & Giordano, 2010). The presence of 
volcanic material, among which pumice and tephra in-
terbedded with alluvial-deltaic sediments, is of consider-

able interest to obtain 40Ar/39Ar radiometric age dates 
and therefore to provide reliable geo-chronological con-
straints for the sediments (Marra & Florindo, 2014 and 
references therein). What makes this territory important 
for paleontological research is the exceptional number of 
remains recovered over the last 150 years from the allu-
vial deposits of the Paleo-Tiber River and its tributaries 
(e.g., Di Stefano et al., 1998; Petronio & Sardella, 1998, 
1999; Milli et al., 2004). The first findings took place in 
the 1800s, when the fluvial deposits were investigated 
by eminent scientists of the time, such as Giuseppe 
Ponzi and Alessandro Portis (Funiciello & Rosa, 1995; 
Funicello & Giordano, 2008b; Romano et al., 2021). The 
majority of the fossil remains was collected from depos-
its outcropping in the urban area of Rome and its periph-
ery, due to the intense urbanization and quarry activities 
(e.g., Portis, 1893, 1896,1900). Many works focused on 
the description of these fossiliferous localities and their 
remains (e.g., Di Stefano et al., 1998; Petronio & Sar-
della, 1999), even if often most of them lacks exact strat-
igraphic constrains and is not always associated to a 
precise sedimentary level (gravel, sand, clay) (Pandolfi 
& Marra, 2015). Most of the historical collections from 
the area of Rome is stored at the “Museo Universitario di 
Scienze della Terra (MUST)” (Earth Science University 
Museum, Sapienza University of Rome) (including the 
former Museo di Paleontologia di Roma, MPUR). Unfor-
tunately, the paleontological collections endured a series 
of misadventures, as for instance the overflow of the 
Tiber River in 1870 (Portis, 1893) or the bombardment 
that struck the Department of Earth Sciences of Sapien-
za University (Rome) carried out by the American Army 

 

https://doi.org/10.26382/AMQ.2021.09 



   

 

 

 

132 

 

 
Mecozzi B. et al. 

Fig. 1 - Geological sketch map of the Ponte Molle-Tor di Quinto area (from Funiciello & Giordano, 2008a; modified), with trace of the geo-
logical cross section (Fig. 2), the location of the Cava D’Alessandri and boreholes used for stratigraphic analysis (Figs 4, 5). Toponyms 
refer to the historical geological maps by Tellini (1893) and Verri (1915); see also figure 3.  



   

 

 

 

133 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

during the World War II (Fabiani & Maxia, 1953; Mecoz-
zi et al., 2020). These episodes caused the loss and 
damage of many fossil specimens or of their associated 
labels. Nonetheless, the fossil remains recovered from 
the urban area of Rome and its surroundings, also 
known as Campagna Romana (Roman Campaign), 
represent the most abundant Middle Pleistocene fossil 
samples founded in Italy, and thus the area assumes a 
key-role in the study of vertebrate assemblage evolu-
tionary trends and palaeoenvironmental reconstructions. 
The importance of this territory is reflected in the Large 
Land Mammal Biochronological Scale, where two Ro-
man localities are chosen as Faunal Units: Ponte Ga-
leria and Torre in Pietra (Gliozzi et al., 1997).  

In this work, we analyze the fossil sample from 
Ponte Molle (also known as Ponte Milvio) (PM), a histor-
ical locality of the urban area of Rome described by 
Portis (1893). The revision of the stratigraphical succes-
sion of the deposit allows us to reconstruct the prove-
nance of the fossil material and to provide chronological 
constrains trough the correlation with the lithostatigraph-
ic and synthemic units of the national geological cartog-
raphy (CARG Project; Funiciello & Giordano, 2008a, b) 
and the geochronologically-constrained aggradational 
units of the Paleo-Tiber reported in literature (Marra et 
al., 2014). 

 
2. PONTE MOLLE AND ITS TROUBLED HISTORY 

 
The presence of the fossil remains in the Tiber 

alluvial deposit in the Tor di Quinto area was already 
reported in the second half of the 19th century, when 
Ponzi (1867) recognized the occurrence of several spe-
cies from the quarries opened at PM, from the same 
gravels quarried for building material (Tab. 1). The strat-
igraphical succession exposed in the quarry called Cava 
D’Alessandri was firstly described by Portis (1893). In 
this work, he illustrated several sedimentary deposits 
occurred in the city of Rome, with a special focus on the 
northern-western area, where in addition to Cava 
D’Alessandri, the successions exposed at Tor di Quinto 
(=Torretta di Quinto) and Acquatraversa (all located in 
the Tor di Quinto district) were also described (Fig. 1). 
The author pointed out the presence of fossil remains 
referable to the “classical” fauna of PM (implying that 
findings of similar assemblages were already common) 
only from the lowermost level of the stratigraphic suc-
cession (Tab. 1). This level, consisting of tuffaceous 
gravels and sands (i.e. with tuffaceous clasts and miner-
als of volcanic origin), and including a diversified faunal 
assemblage, was the first properly described outcrop of 
the “cosiddette ghiaie diluviali di Ponte Molle” [so-called 
diluvial gravel of Ponte Molle]. Later, Portis (1902) de-
scribed an isolated upper molar belonging to a juvenile 
specimen classified as Elephas antiquus 
(=Palaeoloxodon antiquus) (Tab. 1). Portis (1907) ana-
lyzed the sample of Bovidae from several deposits of 
the urban area of Rome and its surroundings, attributing 
several cranial remains from PM to Bos primigenius 
(Tab. 1). Portis (1909) studied the canid fossil remains 
from the urban area of Rome, among which two frag-
ments of hemimandible and one isolated lower carnassi-
al from the “cosiddette ghiaie di Ponte Molle presso 

Cava D’Alessandri” were referred to a small-sized form 
of Canis lupus (Tab. 1). In addition, he noticed that in 
the fossil collection of Ponzi two replica, a fragment of 
maxillary and a fragment of hemimandible, respectively, 
were present, but the original remains were missing. 
These two fossils were also referred to C. lupus. Portis 
(1909) also attributed a complete femur to Meles meles 
(=Meles taxus) and an upper canine to the genus Lynx, 
both collected from the basal level of Cava D’Alessandri 
(Tab. 1). Later, Portis (1920) listed the Cervidae taxa 
identified in the city of Rome and its surroundings, re-
porting the presence of Cervus (Praealces) latifrons 
from PM (Tab. 1). However, due to the intense urbaniza-
tion of the area, the quarries and the other aforemen-
tioned outcrops were destroyed or buried below anthro-
pogenic deposits. Following this, subsequent studies 
were conducted exclusively on the historical fossil col-
lections recovered during the end of 1800s and the first 
decades of 1900. During the 1960s, Ambrosetti & Bo-
nadonna (1967) resumed the paleontological research 
on this locality and correlated the PM fauna with the 
Ponte Galeria formation (sensu Conato et al., 1980) 
exclusively on the basis of the occurrence of some Ga-
lerian taxa, even if no description of the materials was 
performed. After decades, Caloi & Palombo (1986) car-
ried out the revision of the fossiliferous localities of the 
area of Rome and listed the following species from the 
lower level of Ponte Milvio (=Ponte Molle): Cervus 
(Cervus) acoronatus, Cervus (Dama) sp. (=Cervus 
[Dama] quirinus), Megaceros cf. solilhacus (=Cervus 
[Praealces] latifrons) and Hippopotamus incognitus 
(Tab. 1). Even in this case, neither descriptions nor fig-
ures of the fossil sample were included. Di Stefano & 
Petronio (1992) firstly described the mammal assem-
blage from PM, identifying two distinct faunal assem-
blages, as they regarded some mammals as Galerian 
forms (e.g. C. elaphus acoronatus) and others as more 
typical elements of late Middle to Late Pleistocene as-
semblages (e.g. Dama clactoniana) (Tab. 1). Capasso 
Barbato et al. (1998) realized a preliminary revision of 
the deposit, recognizing at least three different assem-
blages of Middle Pleistocene age and also referring a 
few specimens to the Holocene (Tab. 1). Shortly after, 
Di Stefano et al. (1998) further detailed the “two-
assemblages” hypothesis, attributing the first to Isernia 
Faunal Unit (early Middle Pleistocene) and the second 
to Vitinia Faunal Unit (late Middle Pleistocene), mainly 
on the ground of the presence of different fallow deer 
subspecies, respectively Dama clactoniana and Dama 
dama tiberina. 

More recently, an isolated horn-core stored at the 
Monastery of Santa Scolastica in Subiaco (Rome), pre-
viously referred to Bubalus murrensis by Cassoli & Seg-
re (2004), was ascribed to Hemibos galerianus by Mar-
tínez-Navarro & Palombo (2007) (Tab. 1). Palombo 
(2004) figured a partial cranium with nearly complete 
antler classified as C. elaphus acoronatus and a partial 
cranium and complete antler ascribed to D. dama tiber-
ina (=Dama quirinus).  

Billia & Petronio (2009) revised the Rhinocerotidae 
sample, recognizing only the presence of Stephanorhi-
nus kirchbergensis, whereas Pandolfi (2013) and Pan-
dolfi & Marra (2015) identified three different taxa: S. 



   

 

 

 

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Mecozzi B. et al. 



   

 

ly described outcrop (Portis, 1893). Despite several 
specimens from this site have been object of study (e.g., 
Capasso Barbato et al., 1998; Di Stefano et al., 1998), 
an in-depth analysis is still needed to redefine the faunal 
assemblage.  

 
3. GEOLOGICAL SETTING 

 
The study area of PM and Tor di Quinto is located 

on the right bank of the Tiber River and corresponds to a 
well preserved fluvial terrace, with 50-60 m a.s.l. mean 
elevation, laying eastward of the N-S directed Monte 
Mario-Monti della Farnesina ridge, up to 140 m high 
(Fig. 1). The terrace is bounded to the east and south by 
the alluvial plain of the meandering Tiber River, at 16 m 
a.s.l., and to the north by the narrow valley of Fosso 
dell’Acquatraversa.  

The original terrace relief, as deduced by the ob-
servation of historical geological and topographic maps 
(Tellini, 1893; Verri, 1915; IGM, 1936), was character-
ized by a gentle undulating topography interrupted, inter-
nally, by small N-S oriented valleys and at the northern 
and eastern edges by steep scarps, where natural out-
crops were exposed and examined by the authors 
(Portis, 1893; De Stefani, 1904). Nowadays the natural 
landform is strongly modified after a re-shaping from the 
intense urban transformations of the last 120 years, and 
almost no exposures are preserved.  

On the local stratigraphy, the bedrock corresponds 
to the marine clay, silt and sands of the Monte Vaticano 
and Monte Mario Formations, attributed respectively to 
the Zanclean-Gelasian and to the late Santernian 
(Calabrian) (Funiciello & Giordano, 2008a, 2008b). The 
bedrock crops out along the Monte Mario-Monti della 
Farnesina ridge, is found in the subsoil of the fluvial 
terrace (Fig. 2), and is crossed by NW-SE trending nor-
mal faults down-throwing the north-eastern blocks 

kirchbergensis, Stephanorhinus hemitoechus and 
Stephanorhinus hundsheimensis, as well as several 
remains undeterminable at specific level 
(Stephanorhinus sp.) (Tab. 1). Pandolfi & Marra (2015) 
also proposed a reinterpretation of the age of the depos-
it. They noticed that part of the specimens labels report 
that fossils were collected from “gravels and sands” and 
others from “tuffaceous conglomerates”, and supposed 
this as indicative of the presence of two levels of differ-
ent age: the lower one, a gravel and sand level where S. 
hundsheimensis occurred; and the upper one, a tuffa-
ceous conglomerate level which included S. kirchber-
gensis and S. hemitoechus. This upper level was corre-
lated with a tephra detected at 1.5 km along Via Flamin-
ia Nuova, dated at 465±7 ka, whereas the lower level 
was correlated to the base of the Paleo-Tiber 2 cycle at 
810-790 ka (Pandolfi & Marra, 2015). However, from an 
historical perspective, this label-based repartition of the 
assemblage is not supported. In fact, even not accepting 
early researchers’ interpretations, they explicitly regard-
ed the fauna of PM as coming from a single level (Ponzi, 
1867; Portis, 1893). Consequently, they would not have 
reported on the labels a separation that they had not 
observed in the field. Moreover, even though some la-
bels do report “gravels and sands” and others 
“tuffaceous conglomerates”, others again indicate 
“gravels, sands, and tuffaceous conglomerates”, or even 
“so-called gravels of Ponte Molle” or variable combina-
tions of these forms, which clearly were all different 
ways to refer to the same thing: a fluvial formation with 
gravel and sand, rich in fossils and volcanic material. 

The so-called fauna of PM was the typical assem-
blage recovered from the gravel quarries and other spo-
radic findings occurred during construction works in the 
district of Tor di Quinto, in the northern-western area of 
Rome (Ponzi, 1867). Several specimens were recovered 
from Cava D’Alessandri, which was also the first proper-

 

 

135 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

Fig. 2 - Geological cross section of the Ponte Molle area. The trace of the section is represented on figure 1.  



   

 

 

 

136 

 

 
Mecozzi B. et al. 

(Funiciello & Giordano, 2008a; Cosentino et al., 2009).  
The terrace is composed of an approximately 50 m 

thick succession of fluvial deposits with interbedded 
pyroclastic units, which covers with angular unconformi-
ty the bedrock; it is interpreted as an alluvial fill terrace 
(Merrits, 2007; Pazzaglia, 2013). At the base are pre-
sent alternated gravel, sand and silt of the Fosso della 
Crescenza and Santa Cecilia Formations of the latest 
Early Pleistocene-early Middle Pleistocene. They record 
the oldest fluvial sedimentation in the area, up to the 
MIS 16-15, and are covered by the Via Tiberina Unit 
(Tufo Giallo della Via Tiberina Auct) an approximately 
550 ka old and few meters thick ignimbrite (Karner et 
al., 2001), sourced by the Sabatini Mts District and crop-
ping out close to the Torretta site.  

Most of the fill terrace is represented by the fluvial 
Valle Giulia Formation (MIS 14-13), up to 30 m thick and 
composed of basal channel gravels grading upward into 
cross bedded sands, travertine, tufas and floodplain 
muds. This formation has a relatively high-relief basal 
unconformity carved into the bedrock and older fluvial 
deposits and defines an approximately N-S directed 
paleo-valley infill (Giordano et al., 2003; Funiciello & 
Giordano, 2008a; Marra & Florindo, 2014; Giustini et al., 
2018). The formation is rich in volcanoclastic material, 
re-sedimented in the fluvial environment, and crystals of 
volcanic origin found in the sandy matrix of the gravels. 
North of the Acquatraversa Stream the “Tufi stratificati 
varicolori di Sacrofano” crop out, composed of alternat-
ed ash and lapilli sized fallout beds (510-460 ka old) 
that, along with the Valle Giulia Formation, define the 
Villa Glori Synthem.  

On the top of the terrace there are locally present 
the “Tufo rosso a scorie nere sabatino”, a red massive 
ignimbrite with back scoria (about 449 ka old; Karner et 
al., 2001) and the overlaying Vitinia Formation (late 
Middle Pleistocene, approximately 285-270 ka old; Kar-
ner et al., 2001; Giordano et al., 2003) composed of few 
meters thick pebbly sand and mud, rich in volcanic min-
erals. The Vitinia Formation corresponds in this area to 
the “Formazione fluvio-lacustre” by Ventriglia (1971, 
2002). Finally, the Tiber River plain and Acquatraversa 
valley are composed of basal gravels and overlaying 
channel sands and floodplain mud with interbedded 
peat layers of the Tiber River Synthem (or Tiber Deposi-
tional Sequence sensu Milli et al., 2016), which records 
the last cycle of fluvial incision and sedimentation of 
Late Pleistocene-Holocene (MIS 5d-1). Anthropogenic 
deposits, up to 15 m thick, cover the natural substratum 
in most of the study area.  

 
4. MATERIALS AND METHODS 

 
The osteological analyses of the large mammal 

fauna from the Middle Pleistocene site of PM were car-
ried out, considering remains collected from the end of 
1800 to early decades of 1900. The fossil remains are 
stored at MUST. Taxonomic and skeletal element identi-
fications made in this study are based on the reference 
collection of the PaleoFactory Laboratory, Department 
of Earth Sciences, Sapienza, University of Rome (PF). 
In order to evaluate species abundance, the count of the 
number of remains (NISP) (Grayson, 1984) and the 

estimate of the minimum number of individuals (MNI) 
(Bökönyi, 1970) have been performed (Tab. 2). In addi-
tion, the age at death was also determined.  

We carried out biometric comparisons for a selec-
tion of taxa relevant for their biochronological and pale-
oenvironmental significance: Bos primigenius, medium-
sized deer (Dama spp. and Dama-like), Sus scrofa, and 
Hippopotamus. For B. primigenius and medium-sized 
deer, we measured the maximum meso-distal length 
above the root-crown junction of the lower third molar 

(M3L). For S. scrofa, measurements of the lower third 
molar were taken following von den Driesch (1976). For 
Hippopotamus ssp. measurements of the lower third 
molar were taken following Mazza (1995): OL: outer 
length; AB: anterior breadth. Measurements were taken 
with a digital caliper to the nearest 0.1 mm.  

For Bos primigenius we considered specimens 
stored in the following institutions: Contrada Cozze, 
Casa Minniti (National Archeological Museum of Melfi); 
as well as literature data: Punta Lucero (Gomez-Oliveira 
et al., 2015); Malagrotta (Caloi & Palombo, 1979); Igue 
des Rameaux- amont, Lunel-Viel, Pech de l’Azé II 
(Uzunidis-Boutillier, 2017); Fara Sabina (Angelelli, 
1981); Ilford (Wright, 2013). Subfossil samples of Bos 
taurus from Elvas-Kreuzwiese (Boschin, 2018) and Sil-
ves-lix (Davis et al., 2008) were also included. We se-
lected the M3 since it is the best documented tooth in the 
studied sample. In addition, this tooth is easily distin-
guishable from the other lower molars, which makes 
more reliable the literature data used for comparison. 

Morphometric data of medium-sized deer from 
Pirro Nord and extant Italian specimens of Dama dama 
dama stored at PF have been considered. Literature 
data of several medium-sized deer taxa have been in-
cluded: Pseudodama group: Kalamotó (Dama sp., 
Tsoukala & Chatzopoulou, 2005); Saint-Prest (Dama 
sp., Guérin et al., 2003), Erpfinger Höhle (Dama nestii, 
Lehmann, 1957), Le Vallonnet (Dama vallonnetensis, de 
Lumley et al., 1988), Cueva Victoria (Pseudodama val-
lonnetensis, van der Made, 2012), Atapuerca TD8 
(Dama vallonnetensis, van der Made et al., 2017a); Da-

ma roberti: Pakefield, Westbury, West Runton (Breda & 
Lister, 2013), Contrada Monticelli (Dama cf. roberti; 
Stefanelli et al., 2021), Valdemino (Breda, 2015); Dama 
clactoniana: Fontana Ranuccio, Grays Thurrock, Jar-
wick, Visogliano (Di Stefano, 1994), Notarchirico 
(Cassoli et al., 1999), Atapuerca TD11 (Azanza & 
Sanchez, 1990), Caune de l’Arago (Dama cf. clactoni-
ana, Magniez et al., 2013). We selected the lower third 
molar (M3 ) since it is the best documented tooth in our 
Dama sample. In addition, this tooth can be easily rec-
ognized from the other lower molars, and therefore a 
misleading interpretation can be excluded from the data 
taken from the paleontological literature. 

For Sus scrofa we considered specimens studied 
in the following institutions: Bristie 1 (Civic Museum of 
Natural History, Trieste), Melpignano (PF), and literature 
data for West Runton, Trimingham, Grays Thurrock, 
Oreston, Hutton (Lister et al., 2010); Petralona 
(Tsoukala & Guérin, 2016); Kyparíssia (Atanassiou et 
al., 2018); Gajtan, Lunel-Viel (Fistani, 1996); Cerè 
(Fabiani, 1919); Torre del Pagliaccetto (Caloi & Pal-
ombo, 1978), and extant Italian wild boar (Iannucci et 



   

 

al., 2020b); considering 
that a relationship be-
tween size shifts in S. 
scrofa and environmental 
fluctuations has been 
recently recognized in 
late Middle Pleistocene-
Early Holocene Apulia 
(southern Italy), we also 
included several Late 
Pleistocene Apulian local-
ities in the comparison, 
grouped in “Glacial” (MIS 
4  a n d  2 )  a n d 
“Interglacial” (MIS 5 and 
3) samples (Iannucci et 
al., 2020b). We selected 
M3 as it is widely available 
in our sample and in the 
literature, as well as be-
ing easily identifiable and 
common focus of studies 
on wild boar morphome-
try. 

We also considered 
morphometric literature 
data of genus Hippopota-
mus from Plio-Pleistocene sites of Europe as well as 
extant Hippopotamus amphibius (Mazza, 1995). We 
selected M3 because it is the most numerous tooth in 
our sample.  

For investigating chronological and/or geographical 
biometric variability and possible size variations in the 
selected taxa we used boxplots of M3 length, for B. prim-
igenius, medium-sized deer, and S. scrofa; and plot of 
length vs breadth of M3 for Hippopotamus. 

 
4.1. Cartography and stratigraphic analysis 

Historical and modern geological and topographic 
maps have been analyzed and compared among each 
other (Tellini, 1893; Verri, 1915; IGM, 1936; Ventriglia, 
1971, 2002; Funiciello & Giordano, 2008a), coupled with 
a bibliographic review of scientific articles and historical 
chronicles (Ponzi, 1867; Portis, 1893; De Stefani, 1904; 
Ranzato, 2019), in order: 1) to locate as precisely as 
possible the Cava D’Alessandri, other old toponyms and 
fossil sites of PM, and the described stratigraphic sec-
tions no longer observable; 2) to infer the position and 
areal extension of other old quarries active between the 
last decades of the 19th and first half of the 20th century 
(Fig. 1).  

In particular, the synoptic observation of old maps, 
from the oldest to the youngest, has allowed to individu-
alize changes in the shape of topographic contour lines 
(i.e. from a convex to a concave one) resulting in half-
circular morphologies in plain view, here interpreted as 
ancient quarry fronts and floors (see also Ciotoli et al., 
2015, for the method). In situ observations and 3D 
views from the software Google Earth Im-
age©2019TerraMetrics has permitted, in some cases, to 
recognize steep scarps and differences in heights within 
the urbanized area, strengthening the hypothesis of old 
quarry fronts.  

The stratigraphic section of Cava D’Alessandri has 
been drawn basing on the description by Portis (1893), 
who reported the precise position of the unique verte-
brate-bearing level of the quarry.  

To reconstruct the stratigraphic architecture of the 
PM-Tor di Quinto infill terrace, two correlation panels 
among the Cava D’Alessandri section and 15 borehole 
logs have been realized. The latter deriving from litera-
ture and unpublished data (Ventriglia, 2002) and stored 
in the CNR IGAG database of subsoil data of Rome 
(Cavarretta et al., 2005). Correlation was based on the 
application of standard methods of physical stratigraphy 
on core data (Bridge, 2003; Collinson et al., 2006) with 
particular emphasis on the identification of key bounding 
surfaces and architectural elements: main unconformi-
ties, channel bodies, floodplain deposits. This has al-
lowed to put the vertebrate-bearing level of Cava D’Ales-
sandri into the correct physical and chrono-stratigraphic 
frame.  

 
5. RESULTS 

 
5.1. Geography and stratigraphy 

From the compared analysis of cartographic and 
literature data (Portis, 1893; Tellini, 1893; De Stefani, 
1904; Verri, 1915) it appears that Cava D’Alessandri 
was likely sited on the northern flank of via Flaminia 
about 150 m north of the Osteria di Melafumo (Tavern), 
with this tavern located just after PM (Fig. 1). Figure 1 
shows the localities mentioned by the authors and an-
cient toponyms, in some cases no longer in use, plotted 
on a sketch of the modern geological map of Rome 
(Funiciello & Giordano, 2008a).  

On the Tellini’s (1893) geological map a small N-S 
trending topographic depression can be noticed, which 
cuts transversally the southern slope of the PM-Tor di 

 

 

137 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

 

Fig. 3 - Sketches of the historical geological maps by Tellini (1893) on the left, and Verri (1915) on the 
right, showing the area of Ponte Molle and its transformations between the late XIX and the early XX 
century. In the Tellini’s (1893) map there are featured a small N-S oriented topographic depression, just 
north-east of the Osteria (di Melafumo) toponym, in the same area described by Portis (1893), and a 
fossil site (red asterisk; see also figure 1). In the Verri’s (1915) map it is reported the C. d’Alessandri 
toponym just north of the Osteria, and the topographic depression (the former quarry) is replaced by a 
newly built area (V. Trezza toponym).  



   

 

Quinto terraced relief. This place roughly corresponds to 
the C. d’Alessandri toponym, as reported on the later 
Verri’s (1915) geological map (Fig. 3). The topographic 
depression of the Tellini’s (1893) map is interpreted as 
the ancient Cava D’Alessandri (Fig. 1), being it substitut-
ed by a re-covered area with small buildings above it in 
the Verri’s (1915) map. This testifies the very rapid 
changes of the growing city in the PM area. Another 
fossiliferous site was shown by Tellini (1893) further 
north along the Via Flaminia, and is reported on figure 1, 
as other old quarries and the outcrops described by 
Portis (1893): Torretta and Acquatraversa.  

The stratigraphy of Cava D’Alessandri described 
by Portis (1893) is represented on figure 4. The verte-
brate-bearing level was located in the lower cross bed-
ded fluvial gravels, 8.5 m thick, that are replaced above 
by calcareous sand and silt (1.5 m thick) and by a fining-
upward succession (16 m thick) of sandy pebbles, grey 
sand and silt with concretions, plant fragments and 
freshwater and terrestrial mollusks (Corbicula fluminalis, 
Helicidae). From the modern geological map, Cava 
D’Alessandri belongs to the Valle Giulia Formation (Fig. 
1), ascribed to the intermediate portion of Middle Pleis-
tocene and correlated with MIS 14-13 (Funiciello & 
Giordano, 2008b). 

The stratigraphic-sedimentological log of Cava 
D’Alessandri is correlated with nearby borehole logs 
(see also figures 1 and 2 for their location), which allows 
to define the stratigraphic architecture of the PM-Tor di 
Quinto infill terrace. The two produced correlation pan-
els are presented on figure 5. The first panel (Fig. 5a), in 
which the Cava D’Alessandri log is plotted, is NW-SE 
oriented with an across-valley direction; the other (Fig. 
5b) is almost perpendicular in the along-valley direction 
(NE-SW).  

The panels detail the stratigraphic setting already 
shown in the cross section of figure 2, with the marine 
bedrock of the Monte Vaticano and Monte Mario For-
mations, the overlaying Middle Pleistocene formations 
composing the aggradational terrace (Fosso della 
Crescenza and Santa Cecilia Formations, Via Tiberina 
Unit, Valle Giulia Formation, “Tufo rosso a scorie nere 
sabatino”, Vitinia Formation), the late Pleistocene-
Holocene Tiber River Synthem in the plain, the anthro-
pogenic deposits. The main unconformity separating the 
bedrock from the overlaying infill terrace of PM-Tor di 
Quinto results from the enveloping of the singular basal 
unconformities of the fluvial Fosso della Crescenza, 
Santa Cecilia and Valle Giulia Formations, all incised by 
the ancient Tiber into the bedrock in response of recur-
rent sea level falls and lowstands. The infill terrace is 
thus the result of the complex vertical and lateral stack 
of several Middle Pleistocene units, with a discontinuous 
stratigraphic-sedimentologic record occurred throughout 
the approximate 0.80-0.27 Ma time interval.  

The oldest fluvial and pyroclastic units (Fosso della 
Crescenza, Santa Cecilia Formations and Via Tiberina 
Unit) are rarely crossed by the well cores in the eastern 
portion of the terrace. The fluvial sediments are repre-
sented by prevailing beige sand and silt, with intercalat-
ed pebbles and with a reddish-brown paleosol on the 
top (P2313 borehole; Fig 5b).  

Conversely, the panels show for the Valle Giulia 

Formation the classical pattern of infill of the Quaternary 
fluvial incised valleys in the Rome area, i.e. that related 
both to the Tiber River and tributaries (Milli, 1997; Milli et 
al., 2008, 2016; Mancini et al., 2018), with a high relief 
unconformity, basal gravels and overlaying alternated 
sand and fine deposits.  

The basal unconformity of Valle Giulia Formation is 
well incised into the marine bedrock and the older Fosso 
della Crescenza-Santa Cecilia Formations and Via Ti-
berina Unit (550 ka old); it records the sea level fall oc-
curred between the MIS 15 and MIS 14, and the MIS 14 
lowstand of sea level (glacial phase).  

 

 

138 

 

 
Mecozzi B. et al. 

Fig. 4 - Stratigraphic-sedimentologic log of the Cava D’Alessan-
dri; see figure 1 for the location of the log (red triangle).  



   

 

 

 

139 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

Fig. 5 - Correlation panels among borehole logs and the Cava D’Alessandri log detailing the stratigraphic architecture of the Ponte Molle-
Tor di Quinto terrace: a) across-valley panel; b) down valley panel; c) legend. The location of boreholes is on figure 1. 



   

 

 

 

140 

 

 
Mecozzi B. et al. 

Tab. 2 - Mammal fossil remains from Ponte Molle stored at MUST. 



   

 

 

 

141 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

The lower portion of the incised-valley fill corre-
sponds to amalgamated sandy gravels, 10-15 m thick, 
with cross bedded bars and bedforms of the braided 
river environment. The fossiliferous gravels correspond 
to the “cosiddette ghiaie diluviali di Ponte Molle” by Por-
tis (1893) and form a laterally and downstream continu-
ous body with tabular shape. They are related to the 
onset of the sea level rise occurred between MIS 14 and 
MIS 13 (latest lowstand and early transgressive phases 
of filling), at the end of the glacial phase.  

Above the gravels, fluvial pebbly-silty sands alter-
nate with floodplain fine deposits and with travertines 
and freshwater tufas. All these deposits recorded the 
fluvial sedimentation in response to the late rise and 
highstand of sea level during the MIS 13 (transgressive 
and highstand phases) and under interglacial climate 
conditions.  

The sands define at least three vertically stacked 
channel bodies (sensu Gibling, 2006), each 10 m thick, 
lens shaped and with fining upward arrangement of 
facies, from pebbly to silty sand (Fig. 5a). The channel 
sands, likely attributed to a meandering-style fluvial 
system (as compared to other analogues in the Rome 
area; Milli et al., 2016), are laterally confined by planar 
bedded floodplain silt and muds, and by travertines and 

tufas of fluvial and spring environments. Travertines and 
tufas, cropping out at Tor di Quinto, define a down-
stream almost continuous tabular body in the intermedi-
ate portion of the incised valley fill, at 20-30 m a.s.l. ele-
vations (Fig. 5b). These continental carbonates record a 
well-known phase of increasing CaCO3 deposition by 
spring waters, related to a renewal of the local tectonic 
activity during the MIS 13 (Funiciello & Giordano, 
2008b).  

On the top, the Valle Giulia Formation is con-
strained by the 449 ka old “Tufo rosso a scorie nero 
sabatino”, which deposited on a morphologically very 
articulated surface recording the main fluvial incision at 
the MIS 13-12 transition. A following phase of fluvial 
sedimentation is represented by the Via Tiberina Unit, 
that deposited at the end of MIS 9 and MIS 8.5 
(Giordano et al., 2003).  

 
5.2. Faunal assemblage from Ponte Molle 

The fossil sample from PM includes 741 taxonomi-
cally identified elements recovered from the lower part of 
the deposit, chronologically referred to the Middle Pleis-
tocene (Tab. 2) (Fig. 6). The faunal assemblage is large-
ly dominated by Bos primigenius (50.5%), which is main-
ly represented by isolated upper and lower teeth (Tab. 

Fig. 6 - Mammal fossils from Ponte Molle: a - 415, right hemimandible of Bos primigenius in labial view; b - 426, right hemimandible of Bos 
primigenius in labial view; c -1857, right maxillary of Palaeoloxodon antiquus in occlusal view; d - 46, left hemimandible of Hippopotamus ex 
gr. antiquus in lingual view; e - 166, left lower third molar of Sus scrofa in lingual view; f - 718, right hemimandible of Capreolus capreolus in 
labial view; g - 1175, right lower third premolar of Crocuta crocuta in labial view; h - 128, right hemimandible of Castor fiber in lingual view. 
Scale bar 3 cm.  



   

 

 

 

142 

 

 
Mecozzi B. et al. 

2). The cervids (Cervus elaphus eostephanoceros and 
Dama clactoniana) (14.5%), a large equid (Equus 
mosbachensis) (8.1%), the straight tusked elephant 
(Palaeoloxodon antiquus) (7%), rhinos (Stephanorhinus 
hemitoechus, Stephanorhinus kirchbergensis, Stepha-
norhinus sp.) (7.6%) and Hippopotamus ex gr. antiquus 
(=Hippopotamus tiberinus) (6.7%) are well represented, 
whereas Sus scrofa, Equus hydruntinus and Capreolus 
capreolus are quite scarce (Tab. 2). A single antler testi-
fies the presence of Cervus elaphus acoronatus (Fig. 7). 
The middle- to small-sized mammals are represented by 
a lower number of fossils (2.7%), Among these, there 
are 14 remains ascribed to a single Lynx individual.  

A restricted part of the sample is referred to juve-
nile individuals (9.8%), nearly exclusively represented 
by isolated teeth: H. ex gr. antiquus (2 specimens), S. 
hemitoechus (2 specimens), Stephanorhinus sp. (3 
specimens), B. primigenius (37 specimens), C. elaphus 
eostephanoceros (1 specimen), E. mosbachensis (13 
specimens) and E. hydruntinus (1 specimen). An excep-
tion is represented by the remains of Lynx sp., where 14 
juvenile postcranial specimens belonging to the same 

skeleton. The fossils are in a good state of preservation, 
although often enwrapped by a thin crust of sand with 
abundant volcanic material that range from reddish to 
greyish. A few specimens, especially the remains refera-
ble to large herbivores, are partially embedded in peb-
bles whose dimension vary from a few millimeters to 
more than ten centimetres. In the fossil material there is 
no sign of bite marks or rodent gnaw marks, or human 
butchering and exploiting activity. Therefore, it is possi-
ble to suggest that carnivorans or humans did not play 
any key role in the accumulation of the fossil remains. 
Unfortunately, the stratigraphic information written on 
specimen’s labels is quite limited, preventing further 
taphonomic analyses. 

Finally, likely deriving from the upper part of the 
deposit, clearly chronologically referable to the Holocene 
due to their preservation, several species were identi-
fied: B. primigenius (11 specimens), Bubalus sp. (1 
specimen), C. elaphus (1 specimen), Equus ferus (6 
specimens), E. hydruntinus (5 specimens) and M. meles 
(14 specimens; Mecozzi, 2021).  

 
5.3. Taxonomic revision 

The fossil sample from PM was never systemati-
cally studied, with only a preliminary faunal list reported 
by Di Stefano & Petronio (1992) and Capasso Barbato 
et al. (1998) (Tab. 1).  

Here, we revised the fossil material stored at 
MUST. 

The cervid sample is referred to four taxa: C. ela-
phus acoronatus, C. elaphus eostephanoceros, D. clac-
toniana and C. capreolus. The subspecies C. elaphus 
acoronatus is represented by a partial cranium and a 
nearly complete antler (Fig. 7), whereas the rest of the 
red deer sample is classified as C. elaphus eostepha-
noceros (Fig. 7). These two red deer subspecies differ in 
the terminal part of the antler: a simple bifurcation in C. 
elaphus acoronatus and a real crown with at least 5 
tines in C. elaphus eostephanoceros. These two taxa 
were reported by Di Stefano & Petronio (1992, 1993) 
and their presence in the faunal assemblage from PM is 
confirmed.  

Considering the medium-sized deer, no remain can 
be attributed to Euraxis eurygonos (=Axis eurygonos). 
The only fallow deer recognized from PM is D. clactoni-
ana (Fig. 8). Di Stefano & Petronio (1997) proposed a 
new fallow deer subspecies based on the sample of PM, 
D. dama tiberina. The main diagnostic features of this 
taxon are related to the morphology of the antlers, which 
should be different from D. clactoniana in: a posterior 
direction of the spellers on the palm; a more basally 
located brow tine; brow tine and the trez tine less devel-
oped; flatter palm and terminal tines. Nevertheless, the 
morphology of the antler from PM is extremely close to 
that of D. clactoniana from Fontana Ranuccio, Riano 
and Swanscombe (Fig. 8). Considering that the taxono-
my of the Plio-Pleistocene cervid is mainly based on 
antler (e.g., Di Stefano & Petronio, 1993; Breda & Lister, 
2013), the morphological affinity of the antler from PM to 
that of D. clactoniana suggests it belongs to the same 
species, thus invalidating the validity of the subspecies 
D. dama tiberina. The differences proposed by Di Stefa-
no & Petronio (1997) are not enough to separate D. 

Fig. 7 - Antlers of Cervus elaphus from Ponte Molle: a, CE 4 
Cervus elaphus acoronatus in frontal (1) and medial (2) views; 
b, SN151/FS, Cervus elaphus eostephanoceros in lateral (1) 
and medial (2) views. Scale bar 3 cm.  



   

 

 

 

143 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

third molar (see Sala, 1986 for discussion). In postcrani-
al elements, clear diagnostic features are less numerous 
but however significative, as the distally divergence of 
the medial and lateral intercondylar crests in metapodi-
als (see Sala, 1986, for the description of other diagnos-
tic features). The overall morphology of the sample from 
PM is close to that of B. primigenius. 

Equids are the other well represented group in the 
faunal assemblage from PM, with two forms well-
different in size. The taxonomic status of the Middle 
Pleistocene Equus is still a controversial topic, and no 
consensus is reached (van Asperen, 2012; Boulbes & 
van Asperen, 2019). The open issues concern how to 
interpret the biometric variations, if related to intra-
specific variability or ecomorphotypes (e.g., van 
Asperen, 2012; Boulbes & van Asperen, 2019). As such, 
several authors referred the Middle Pleistocene fossils 
to E. mosbachensis, taxon only characterized for its 
large size and a few features of the postcranial elements 
(e.g., presence of the tendon insertion of the anterior 
brachialis muscle on the inner edge of the diaphysis of 
the radius, strong supra-articular tuberosities on meta-
podials) (e.g., Uzunidis-Boutillier, 2017; Boulbes & van 
Asperen, 2019). Conversely, van Asperen (2013) con-
sidered the European Middle Pleistocene sample as a 
single species, E. ferus ssp. The dental remains from 
PM are large-sized, whereas the postcranial elements 
provided no element for a taxonomical allocation. Con-
sidering the dimensions, the specimens are referred to 
the European Middle Pleistocene E. mosbachensis.  

A few teeth of equid from PM can be instead re-
ferred to a small-sized form. Dental diagnostic features 
of E. hydruntinus have been summarized by Boulbes 
(2009): angular parastyle, rounded and narrow normally 
mesostyle, a deep postprotoconal valley, pli caballin 
simplified and short protocone in the upper teeth; 
metastylid longer than metaconid, with a less deep lin-

clactoniana and D. dama tiberina, but rather fall within 
the intraspecific morphological variability of the former. 
By contrast, the overall antler morphology of D. clactoni-
ana largely differs from that of D. dama (see Leonardi & 
Petronio, 1976 for discussion).  

A small-sized deer is also documented at PM, 
attested by a nearly complete hemimandible. Its mor-
phology cannot be distinct from that of the extant speci-
mens of Capreolus capreolus. Another cervid taxon was 
reported at PM, classified as Megaceros cf. solilhacus 
(=Cervus [Praealces] latifrons) (Tab. 1). Despite this, no 
remains of the MUST collection can be referred to a 
megacerine deer, but we cannot exclude that speci-
mens of this taxon are stored in other repositories, as in 
the case of the horn core of Hemibos galerianus.  

The taxonomical identification of Middle Pleisto-
cene bovids faced considerable difficulties, especially 
for isolated and/or fragment fossils. Nevertheless, the 
sample from PM includes 374 remains, including diag-
nostic elements (Tab. 2). Portis (1907) also described 
several cranial fragments. We were able to recognize 
only a few of these specimens in MUST. There are, 
however, a few crania and cranial fragments in the old 
collection of MUST without information on their prove-
nance and in need of restoration, some of which could 
have been recovered from PM. Several features can be 
detected in the PM sample: horn cores inserted in the 
cranium in a more latero-posterior position, with an out-
ward and backward direction in the proximal part, and 
upward and slightly forward in the terminal part (e.g., 
Portis, 1907, Plate XIII, Fig. 3); well-developed entostyle 
and a more columnar and hypsodont appearance 
(swelling absent) of the upper molars; V-shaped enamel 
around the central cavity of both the upper and lower 
molars; the two main lobes more mesiodistally devel-
oped in the lower molars; presence of a small accessory 
stylid between hypoconid and hypoconulid in the lower 

Fig. 8 - Antlers of fallow deer: a, MPUR 605 from Ponte Molle; b, Riano 6 from Riano (Leonardi & Petronio, 1976); c, 16349 from Swans-
combe (Leonardi & Petronio, 1976); d, FR56539 from Fontana Ranuccio (Cassoli & Segre Naldini, 1993); e, no catalogue number of extant 
Italian specimen of D. dama dama (Di Stefano & Petronio, 1993). Colours: yellow -Dama clactoniana; green - extant Dama dama dama. 
The images are not in scale.  



   

 

gual groove, a deep ectoflexid (on 
molars) and a few marked or ab-
sent pli caballin in the lower teeth. 
These features have been ob-
served in the sample from PM, and 
therefore the presence of E. hy-
druntinus can be confirmed.  

The hippo remains are very 
common in the European Plio-
Pleistocene record, but even in this 
case there are conflicting opinions 
on their systematics and evolution 
(see van der Made et al., 2017b for 
discussion). Three forms are gener-
ally recognized: Hippopotamus 
antiquus, Hippopotamus ex gr. an-
tiquus (= H. tiberinus) and Hippo-
potamus amphibius. Whereas the 
last shows clearly diagnostic mor-
phological characters, well different 
than the other two taxa, H. antiquus 
and H. ex gr. antiquus differ mainly 
for their size (Mazza, 1995; Mazza 
& Bertini, 2013). Following the taxo-
nomical diagnosis proposed by 
Mazza (1995), the morphology of 
the specimens from PM falls in the 
variability of H. antiquus. Based on 
the medium size of the remains, the 
sample from PM is ascribed to H. 
ex gr. antiquus.  

The elephant sample from PM 
includes fragmentary tusks, partial 
maxillaries and hemimandibles, 
isolated upper and lower teeth and 
several postcranial elements (Tab. 
2). Tusks are incomplete and no 
useful taxonomical characters can 
be observed. Dental remains are 
hypsodont and generally possess a 
high number of laminae, a high 
lamellar frequency, reduced enamel 
thickness and less developed ce-
mentum. These features are con-
sidered typical of P. antiquus 
(Palombo, 1986, 1995; Palombo et 
al., 2003).  

One of the groups better in-
vestigated from PM is the Rhinoce-
rotidae. The sample was taxonomi-
cally identified for the first time by 
Capasso Barbato et al. (1998), who 
recognized the presence of S. kirchbergensis and S. 
hemitoechus. Six isolated teeth of S. kirchbergensis 
were carefully described by Billia & Petronio (2009) 
(Tab. 3). Pandolfi (2013) revised these specimens and 
studied other rhino remains from PM (Tab. 3). The au-
thor classified as S. hundsheimensis four teeth previ-
ously attributed by Billia & Petronio (2009) to S. kirch-
bergensis, and referred other three teeth to S. kirchber-
gensis and four to S. hemitoechus. We need to take into 
account that Pandolfi (2013) recognized two distinct 
faunal assemblages, referred to Ponte Galeria and 

Vitinia FUs respectively. Following this chronological 
repartition, the author attributed a few fossils from this 
hypothetical “lower level” to S. hundsheimensis. The 
revision of the material from PM supports the classifica-
tion of Billia & Petronio (2009) for MPUR 1412/8, MPUR 
1417/115, MPUR 1421/107, MPUR 1445/27, MPUR 
1454/117 and MPUR 1454/118 as S. kirchbergensis. 
The revision of MPUR 1420/97 led doubt on the pres-
ence of S. hundsheimensis. In fact, its morphology dif-
fers from the P4 of S. hundsheimensis from Isernia La 
Pineta (Ballatore & Breda, 2013) in the profile of the 

 

 

 

144 

 

 
Mecozzi B. et al. 

Tab. 3 - Stephanorhinus remains identified in previous works 



   

 

reported the presence of Lynx, Canis lupus and Meles 
meles (Tab. 1). According to the author, the lynx sample 
included a partial hemimandible, currently lost. Other 
fossils belong to the same skeleton probably found in 
anatomic connection. A recent study focuses on evolu-
tionary history of European fossil lynxes, which reveals 
as L. pardinus (=Lynx spelaeus/Lynx pardinus spelaeus) 
is the only lynx recognized during the Middle Pleisto-
cene (Mecozzi et al., 2021a). In the PM sample, only 
skeleton remains are documented, which have small 
dimensions. Nevertheless, postcranial diagnostic char-
acters in fossil lynxes are quite unmapped. Considering 
this, we attributed these specimens to Lynx sp., avoiding 
the taxonomical attribution based on chronological 
grounds.  

Canid sample consists of a lower canine and two 
replica of left maxillary and right hemimandible respec-
tively. Portis (1909) noted the lacking of the original spe-
cimens, writing (Pag. 220):“Dove si trovano conservati 
gli originali di questi due modelli?” (Where are the origi-
nal specimens of these two replica stored?). In addition, 
three specimens, two partial hemimandibles and a lower 
first molar described by Portis (1909), are not present in 
the MUST sample. The available material shows a re-
duced size and the protocone of the upper fourth premo-
lar is aligned with the mesial margin of the teeth. A small
-sized Canis, currently lost, was also reported by Portis 
(1909). Thus, we suggest a possible attribution to Canis 
cf. mosbachensis. 

Portis (1909) also described the right complete 
humerus of the European badger. The revision of this 
fossil confirms its attribution to M. meles. 

postfossette and the medisinus. In this scenario, we 
confirm the presence of S. kirchbergensis and S. hemi-
toechus, whereas that of S. hundsheimensis cannot be 
supported.  

Sus scrofa is the only suid species documented in 
the European Middle Pleistocene, it differs from the 
Early Pleistocene Sus strozzii in several anatomical 
features, the most notable on wild boar remains from 
PM are the presence of a “scrofic” cross-section of male 
lower canines and the proportionally narrower teeth 
(Iannucci et al., 2020a). The chronosubspecific reparti-
tion in a large-sized Sus scrofa priscus eventually giving 
rise to a small-sized S. scrofa scrofa (e.g., Fistani, 1996) 
seems a too simple interpretation of a more complex 
pattern, with several dimensional shifts occurred during 
the Pleistocene (Lister et al., 2010; Iannucci et al., 
2020b). 

Two taxa are here identified for the first time: Cas-
tor fiber and Lepus sp. The European beaver is docu-
mented only by a partial hemimandible, which possess-
es features falling in the variability of the extant speci-
mens of C. fiber (Komosa et al., 2007; Nowicki et al., 
2019). In addition, the specimen is similar to those re-
ported from Cuenca-Bescos et al. (2017) from several 
Spain Pleistocene deposits, with a lower first molar 
squared in shape in occlusal view, both mesial and pos-
terior sides planar, less brachyodont chewing teeth and 
lacking fossettids (isolated enamel islands on the occlu-
sal surface). Few fragmentary postcranial elements can 
be attributed to hare. Lack of clear diagnostic features 
prevents a specific attribution.  

Carnivorans at PM are quite scarce. Portis (1909) 

 

 

145 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

Fig. 9 - M3 length (mm) of Bos primigenius from Middle Pleistocene sites of Europe and subfossil specimens of Bos taurus. Colors: Orange 
- early Middle Pleistocene: PL, Punta Lucero; CMi, Casa Minniti; CC, Contrada Cozze; Ma, Malagrotta; Red - PM, Ponte Molle; Light blue - 
late Middle Pleistocene: IdR, Igue des Rameaux-amont; Pa, Payre; LV, Lunel-Viel; FS, Fara Sabina; Il, Ilford; PdA, Pech de l’Azé II; Green 
- subfossil specimens of Bos taurus; EK, Elvas-Kreuzwiese; Sl, Silves-lix. Number of specimens indicated in the brackets.  



   

 

 

 

146 

 

 
Mecozzi B. et al. 

The last carnivoran species, reported only by Ca-
passo Barbato et al. (1998) (Tab. 1), is Crocuta crocuta. 
An upper third premolar and a replica of coprolite can be 
assigned to the spotted hyaena. The P3 is squared in 
shape in occlusal view, its labial margin is quite straight 
and the lingual cingulum is absent; distal accessory 
cusp is connected to the paracone with a weak crest. 
These features resemble those of C. crocuta, and differ 
from those of Hyaena prisca (=Pliocrocuta perrieri).  

Two additional carnivorans, an ursid and a large 
felid, were reported by previous works (Tab. 1). Never-
theless, no fossils of the MUST sample can be attribut-
ed to these taxa, thus their presence at PM cannot be 
confirmed. Similarly, to large-sized deer, we cannot 
exclude that additional fossils could be stored in other 
repository, as the Scientific Cabinet of high schools of 
Rome or the Monastery of Santa Scolastica in Subiaco 
(Rome). Finally, considering the misadventures suffered 
by the paleontological collection of MUST during the 
1900 century, we cannot exclude that part of the sample 
from PM has been destroyed.  

 
5.4. Biometry of selected taxa 

The biometric comparison of the M3 of B. primige-

nius from Middle Pleistocene European sites highlights 
no chronological or geographical trend (Fig. 9). In partic-
ular, the rich sample from PM (21 specimens) shows a 
large variability, which encompasses that of the fossil 
specimens from Middle Pleistocene sites of Europe. 
Subfossil samples of B. taurus include specimens with 
shorter M3.  

The length of M3 of the fossil samples belonging to 
the fallow deer lineage shows a large variability (Fig. 
10). The sample from Pirro Nord displays the smallest 
size among the considered samples, well different from 
the other contemporaneous materials (late Early Pleisto-
cene). No great difference can be detected among the 
fossil taxa, which generally possess a longer M3 than the 
extant Italian specimens of D. dama dama. It is notewor-
thy that the two smallest samples in figure 10 are from 
Pirro Nord and Contrada Monticelli sites, both geograph-
ically located in the Apulian Peninsula (southern Italy), 
which represent the southern margin of their geograph-
ical range.  

The biometric comparison of S. scrofa M3 length 
also shows a large variability, with no clear chronological 
or geographical trend identifiable (Fig. 11). Middle Pleis-
tocene specimens are on average larger than the extant 

Fig. 10 - M3 length (mm) of medium-sized deer from late Early Pleistocene to late Middle Pleistocene sites of Europe and extant speci-
mens. Colors: Orange - Pseudodama group: PR, Pirro Nord; Ka, Kalamotó; SP, Saint-Prest; EH, Erpfinger Höhle; Va, Le Vallonnet; CV, 
Cueva Victoria; AT8, Atapuerca TD8; Light blue - Dama roberti: Pa, Pakefield; We, Westbury; WR, West Runton; CM, Contrada Monticelli
(Dama cf. roberti); Vl, Valdemino; red - PM, Ponte Molle; Purple: Dama clactoniana; CA, Caune de l’Arago (Dama cf. clactoniana); FR, 
Fontana Ranuccio; Vi, Visogliano; No, Notarchirico; GT, Grays Thurrock; Ja, Jarwick; AT11, Atapuerca TD11; Green - Ex, extant speci-
mens of Dama dama dama. Number of specimens indicated in the brackets. 



   

 

 

 

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Ponte Molle: a Middle Pleistocene mammal assemblage  

Italian populations, and the sample of Late Pleistocene 
MIS 4 and MIS 2 Apulian sites is the only one markedly 
smaller. Remains from PM are among the largest of the 
European Middle Pleistocene. 

The plot of the M3 of fossil and extant hippos re-
veals differences between the groups: H. antiquus 
which generally has longer teeth. H. ex gr. antiquus, 
fossil and extant H. amphibius are characterized by 
reduced dimensions (Fig. 12).  

 
6. DISCUSSION 

 
6.1. The age of the deposit  

From the stratigraphic analysis, the fauna from 
Cava D’Alessandri can be attributed to the lower gravel-
ly level of the Valle Giulia Formation. This formation is 
chronologically constrained by two radiometrically well 
dated ignimbrites, i.e. at the base by the 550 ka old Via 
Tiberina Unit and on the top by the 449 ka “Tufo rosso a 
scorie nere sabatino”. On the basis of the detailed sedi-
mentological reconstruction of the internal architecture 
of the PM-Tor di Quinto fill terrace (sensu Pazzaglia, 
2013), it is excluded that the basal gravels of PM 
(“cosiddette ghiaie diluviali di Ponte Molle”) could be 
attributed to other fluvial formations older than the Valle 
Giulia Formation. In fact, the Fosso della Crescenza and 
Santa Cecilia Formations in the study area are very 
different from the basal gravels, being represented by 
prevailing sand and silt.  

This suggests, for this restricted area, the occur-
rence of different facies for the different formations (or 
portions of them) to be correlated to diverse depositional 
environments: gravelly-dominated braided river environ-
ment for the basal Valle Giulia Formation; sand-
dominated meandering-style river environment for the 
Fosso della Crescenza-Santa Cecilia Formations (Fig. 
5). 

 
6.2. Biochronological implications 

The revision of the geological and stratigraphic 
data indicates a time deposition between 540 and 460 
ka for the mammal faunal assemblage from PM, which 
confirms a Middle Pleistocene age for the deposit. 
Among mammals identified in the MUST sample, sever-
al species provide further biochronological insights.  

The first group considered is Rhinocerotidae, with 
two species recorded: Stephanorhinus kirchbergensis 
and S. hemitoechus. Stephanorhinus kirchbergensis is 
poorly documented in Italy, but in addition to PM this 
species was recorded from the Middle Pleistocene sites 
of Visogliano (MIS 13-10) (Pandolfi, 2013) and Tor di 
Quinto (MIS 13) (Pandolfi & Marra, 2015). Stephanorhi-

nus hemitoechus can be considered an important mark-
er as its earliest occurrence in Europe is from an unde-
fined site of Campagna Romana, approximately dated at 
about 0.5 Ma (age estimated from the correlation of the 
encrusted pumice after texture and the geochemical 
analyses; Pandolfi et al., 2013), and from the Caune de 

Fig. 11 - M3 length (mm) of Sus scrofa from Middle Pleistocene sites of Europe and extant specimens.  
Colors: Light blue - Sus scrofa: WR, West Runton; Tr, Trimingham; Ky, Kyparissia; Ga, Gajtan; Br, Bristie 1; Ce, Cerè; LV, Lunel-Viel; GT, 
Grays Thurrock; Or, Oreston; Hu, Hutton; Me, Melpignano; TP, Torre del Pagliaccetto; Pe, Petralona; AI, Apulia Interglacial (MIS 5 and 
MIS 3 Apulian sites); AG, Apulia Glacial (MIS 4 and MIS 2 Apulian sites); Red - PM, Ponte Molle; Green - Ex, extant specimens of Sus 
scrofa. Number of specimens indicated in brackets. 



   

 

 

 

148 

 

 
Mecozzi B. et al. 

l’Arago, from levels chronologically referred to MIS 14 
(Moigne et al., 2006).  

The second group with strong chronological impli-
cations is Cervidae. The presence of D. clactoniana and 
C. elaphus eostephanoceros clearly indicates a Middle 
Pleistocene age. The Clacton fallow deer (D. clactoni-
ana) appears in several European localities during MIS 
11 (Breda et al., 2013) but it is possible that its dispersal 
predates this age. Indeed, in central Italy an earlier oc-
currence may be that from the levels alfa and a of No-
tarchirico (Cassoli et al., 1999), whose interbedded re-
worked volcanic minerals have been recently dated 
between 658±9 Ka and 612±5 Ka (Moncel et al., 2020; 
Mecozzi et al., 2021b). The other cervid group, the red 
deer, is represented by two different subspecies: C. 
elaphus acoronatus and C. elaphus eostephanoceros. 
The first subspecies is represented by an isolated partial 
cranium and complete antler (Fig. 7). In Italy, the crown-
less red deer (C. elaphus acoronatus) is recorded from 
Slivia to Isernia FUs (ca 850-550 ka) (e.g., Palombo et 
al., 2001; Petronio et al., 2011). Its presence at PM is 
quite surprising, since it is the only taxon that would 
suggest an early Middle Pleistocene age. Despite this, 
we need to consider that these fossils were collected 
during the end of 1800s and early decades of 1900s, 
during quarrying activity. Moreover, this is one of those 
remains that are not accompanied by a historical label, 
and the possibility that it may have been recovered from 
an older deposit outcropping in the surrounding area 
should be taken into account. The other red deer, most 
common in the studied sample, is the eostephanoceros 
red deer (C. elaphus eostephanoceros) (Fig. 7). This 
taxon was instituted by Di Stefano & Petronio (1993) 
mainly considering the antler discovered at Cava Nera 
Molinario (Rome, central Italy) and Fontana Ranuccio, 
localities chronologically ranging between about 500 - 
400 Ka (Marra et al., 2018; Strani et al., 2018). Howev-
er, considering the chronostratigraphic revision carried 
out by Marra et al. (2014, 2018), it seems that both sub-
species are documented from deposits of the area of 

Rome referred to MIS 13, namely C. elaphus acoronatus 
from Via Flaminia Km 8.2, and C. elaphus eostephanoc-
eros from Cava Nera Molinario) (see Iannucci et al., 
2021 for discussion).  

Among the equids, of considerable interest is the 
presence of E. hydruntinus, whose evolutionary history 
was recently redefined by Boulbes & van Asperen 
(2019). The authors suggested a clear presence of this 
taxon in the European record since the late Middle Pleis-
tocene (MIS 11), with a possible first dispersal during 
MIS 15. In Italy, this taxon was considered as a marker 
of the Aurelian large mammal faunal assemblage (late 
Middle Pleistocene, Gliozzi et al., 1997). However, new 
findings from Vallparadís Section (level 11, EVT3, dated 
later than 600 ka) (Aurell-Garrido et al., 2010; Martínez 
et al., 2014) and quarry Carpentier d’Abbeville (dated 
and correlated with MIS 15) (Antoine et al., 2016) sug-
gest an earlier dispersal of the Regalìa Ass (E. hydrunti-
nus) in Europe.  

The wild boar, S. scrofa, is scarcely represented 
during early Middle Pleistocene glacial stages, and the 
relatively large size of the remains recovered from PM is 
close to that of other findings referred to MIS 13, MIS 11 
or MIS 9 interglacials (Fig. 11; Iannucci et al., 2020b).  

Other mammal taxa identified in the PM faunal 
assemblage indicate a Middle Pleistocene age, but with 
no clear chronological definition, among which H. ex gr. 
antiquus, E. mosbachensis, and C. cf. mosbachensis. 
Finally, several species from PM provide no constraints, 
since they are characterized by a long chronological 
distribution (Middle to Late Pleistocene or Holocene), as 
P. antiquus, B. primigenius, S. scrofa, C. capreolus, C. 
fiber, C. crocuta, Lynx sp. and M. meles. In this scenar-
io, the faunal assemblage of PM could be referred to 
MIS 13.  

 
6.3. Paleoenvironmental implications 

Redefining the PM faunal assemblage provides 
important paleoenvironmental and paleoecological 
clues. Indeed, the species represented constitute a fau-

Fig. 12 - Plot of outer length (OL) vs. anterior breadth (AB) of M3 of fossil and extant hippos.  



   

 

 

 

149 

 

 

Ponte Molle: a Middle Pleistocene mammal assemblage  

nal assemblage typical of Middle Pleistocene interglaci-
als of Italy (Strani et al., 2018). The presence and rela-
tive abundance of P. antiquus, H. ex gr. antiquus, C. 
elaphus eostephanoceros, D. clactoniana, S. scrofa and 
B. primigenius indicate warm and humid climatic condi-
tions. This could be also true for Hemibos galerianus, 
ancestral to the Bubalus lineage (Petronio & Sardella, 
1998; Martinez Navarro et al., 2011), considering the 
occurrences and plausible ecological preferences of the 
extinct buffalo (Koenigswald et al., 2019). Nonetheless, 
the mix of taxa adapted to different environments sug-
gests the presence of a mosaic of open spaces (e.g. 
rhinos and equids) and woodlands (e.g. cervids and S. 
scrofa). 

Other ecological and paleoenvironmental infor-
mation can be inferred from the biometric comparisons 
of S. scrofa and D. clactoniana. In fact, even though 
these species are greatly chronologically and geograph-
ically variable (Lister et al., 2010; van der Made et al., 
2014; Di Stefano et al., 2015; Iannucci et al., 2020a; 
Stefanelli & Mecozzi, 2020), shifts in size can be influ-
enced by ecological factors (Geist, 1971, 1987; Lind-
stedt & Boyce, 1985; Weinstock, 1997).  

Focusing on a temporal and chronologically well-
constrained sample, Iannucci et al. (2020b) recognized 
several size shifts experienced by S. scrofa during the 
late Middle Pleistocene to Early Holocene of Apulia in 
southern Italy, with smaller population occurring during 
glacial stages. They suggest an indirect triggering role 
of climate, in reducing the availability of trophic re-
sources. Even though more data are needed to sub-
stantiate this hypothesis and elucidate its geographical 
constraints, S. scrofa remains from the Middle Pleisto-
cene of the area of Rome are scanty but usually large-
sized (Fig. 11), and thus fit well with what expected for 
interglacial stages in the Mediterranean area. 

The very small size of the fallow deer of PM is also 
interesting though puzzling to interpret. Indeed, accord-
ing to Weinstock (1997) the more severe winters of gla-
cial stages should cause a strong decrease of the food 
supply and lead to a higher mortality rate. This leads to 
a decrease of the intraspecific competition during the 
following growth season (Guthrie, 1984), which implies 
a higher quality and quantity of food resources 
(Weinstock, 1997). Therefore, glacial and interglacial 
size fluctuations in cervids should be observed. Never-
theless, the dimensions of the late Early to Middle Pleis-
tocene European samples do not support this separa-
tion (Fig. 10). It is possible that this reflects a different 
ecological adaptation in the species, which would be 
worth to be investigated in future research. 

 
7. CONCLUSIONS 

 
The large mammal faunal assemblage from Ponte 

Molle is dominated by Bos primigenius, with Palaeoloxo-
don antiquus, Hippopotamus ex gr. antiquus, Cervus 

elaphus eostephanoceros and Equus mosbachensis 
well represented (Tab. 1). The carnivorans are docu-
mented by a few taxa, all represented by a limited sam-
ple.  

Aside from a few spurious Holocene specimens, 
most of the faunal elements are consistent with an at-

tribution to a single assemblage, which best fits within 
the Fontana Ranuccio FU. The historical background 
and geological constraints also support the view that the 
recovery of fossil remains from the area of PM was con-
centrated in the Valle Giulia Formation, which is dated 
between 540 ka and 460 ka. Even though we cannot 
exclude that a few specimens may have been misreport-
ed or mixed from different layers, an age older than 550 
Ky seems highly unlikely, as no formation of such an old 
age is documented in the area. 

Moreover, the presence or relative abundance of 
species usually widespread during warm periods, such 
as B. primigenius, P. antiquus, H. ex gr. antiquus, C. 
elaphus eostephanoceros and D. clactoniana, is also 
suggestive of an interglacial environment. 

The reassessment of the classical fauna from PM, 
and the re-description of the sections of Cava D’Ales-
sandri, Torretta di Quinto, and Acquatraversa will serve 
as a basis for reinterpreting other historical and recent 
collections of the area of Rome, as well as providing 
new data for our understanding of Middle Pleistocene 
mammal assemblages of Europe. 

 
ACKNOWLEDGEMENTS 

The study has been supported by Sapienza Re-
search grants, “Fondi Grandi Scavi” 2020 
(SA120172B2C05E68) (Project leader: Prof. R. Sar-
della) and by the CNR DTA.AD003.316. Project “Sistemi 
terrazzati e valli incise: ricostruzioni stratigrafiche e loro 
implicazioni per la valutazione dei geohazard” (Project 
leader: Dr M. Mancini). 

We wish to thank to L. Riti and M. Macri for the 
access to the fossil collection of the “Museo Universitar-
io di Scienze della Terra” (MUST), Department of Earth 
Sciences, Sapienza, University of Rome, and support 
during its study. We are also grateful to all curators and 
staff members of the other institutions visited during this 
research, D. Arbulla (Civic Museum of Natural History, 
Trieste), R. Rocca (responsible of the project for the 
study of the archeological and paleontological material 
from Cimitero di Atella deposit, Potenza) (National Ar-
cheological Museum of Melfi). The authors wish to thank 
L. Bellucci, F. Bona, J. Conti, D.A. Iurino, I. Mazzini and 
F. Strani for their useful suggestions and support. 

Finally, we wish to thank the Editor, Marco Pere-
sani, and two anonymous referees for comments and 
suggestions that greatly improved the manuscript. 

 
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Ms. received: November 24, 2020 Revised: January 25, 2021 
Accepted: May 16. 2021  Avaiable online: April 14, 2021 

https://doi.org/10.32774/FosRepPal.2020.0613