OJS 647 Bertini et al 36,1 91-119 .pub


   

 

PIACENZIAN TO LATE PLEISTOCENE FLORA AND VEGETATION IN ITALY: A MOVING SKETCH 
 

Adele Bertini1,  Nathalie Combourieu-Nebout2 
 

1 Dipartimento di Scienze della Terra, Università di Firenze, Firenze, Italy. 
2 HPNP, UMR7194 CNRS, Dépt. Homme et Environnement, Muséum national d’Histoire naturelle, Institut de Paléontologie Humaine, 

Paris, France. 
 

Corresponding author: Adele Bertini <adele.bertini@unifi.it> 
 

ABSTRACT: The rich late Neogene and Quaternary stratigraphical record for the Mediterranean area is particularly useful for the 
reconstruction of palaeoenvironments and palaeoclimate changes. 66 Italian pollen sites (38 terrestrial and 28 marine) have 
been selected to cover the 3.60 - 0.0117 Ma time-interval. Pollen reveals evidence for the main changes in both flora composi-
tion and vegetation structure, tracing the imprint of increasingly cooler climates on Italian vegetation since the latest Pliocene. 
The role of both altitudinal gradients and physiographic reorganizations as well as of major taxa replacements and their extinc-
tions-times is evaluated by the study of calibrated pollen records. The response of flora and vegetation to glacial/interglacial (G/I) 
cycles is especially well expressed from the beginning of the Pleistocene, and across the Early-Middle Pleistocene climate tran-
sition when a major decrease in temperature, during both G/I phases, occurred. Alternations of Artemisia steppe and thermophi-
lous forest illustrate the overall vegetation changes during G/I cycles. However, different patterns in the G/I vegetational cycle 
have been detected in both northern and southern pollen sites. The Late Pleistocene high-resolution pollen records document 
the vegetation response to rapid climate changes such as the Dansgaard-Oeschger “cycles” and Heinrich stadials, associated 
with millennial to sub-millennial events. 
 
Keywords: Quaternary palynology, palaeoenvironment, palaeoclimate, 3.60 - 0.0117 Ma, Mediterranean.  

Available online http://amq.aiqua.it 
ISSN (print): 2279-7327, ISSN (online): 2279-7335 
 
Alpine and Mediterranean Quaternary, 36 (1), 2023, 91-119 

1. INTRODUCTION  
 

Since the Neogene, the Mediterranean area has 
undergone palaeogeographic and palaeoclimatic chang-
es that have gone hand in hand with significant orogenic 
activity, which led to the uplift of several chains including 
the Apennines, in Italy (e.g. Carminati & Doglioni, 2012). 
The collision of the African and Eurasian plates reduced 
the Tethys domain from the early Miocene onwards. 
Active Atlantic-Mediterranean water exchanges, through 
the Betic and Rifean corridors, assured open marine 
conditions and circulation until ca 7.2 Ma, while the pro-
gressive restriction of the corridors accentuated the 
isolation of the Mediterranean Basin and supported, at a 
regional scale, the development of the so-called Mes-
sinian salinity crisis, from 5.97 Ma (e.g. Clauzon et al., 
1996; Roveri et al., 2014). At 5.33 Ma open marine con-
ditions were restored at the base of the Pliocene (e.g. 
Van Couvering et al., 2000). Significant geomorphic 
modifications linked to the progressive uplift of the Ap-
ennines promoted the formation of numerous eastward-
progressing and younger, marine and continental ba-
sins, extending WSW of the orographic divide (e.g. Mar-
tini & Sagri, 1993; Santangelo et al., 2012; Cosentino et 
al., 2017). At a global scale, the maximum extension of 

the Arctic ice promoted the onset of well-expressed 
glacial/interglacial (G/I) cycles from 2.6 Ma (Berger & 
Loutre, 1991; Lisiecki & Raymo, 2005; Sarnthein et al., 
2009). The set of previous changes gave rise, at a re-
gional scale, to important reorganization and restructur-
ing of the Italian vegetation, also affecting vertebrate 
communities, as attested by significant turnover events 
(e.g. Masini & Sala, 2007; Palombo, 2007, 2014; Magri 
& Palombo, 2013; Strani et al., 2021), and rearrange-
ment of terrestrial ecosystem structures and patterns. 
The high number of Italian Neogene and Quaternary 
deposits has encouraged advances in several areas of 
geo-stratigraphical research, including palynological 
analyses. Terrestrial vegetation can be reconstructed 
from the rich pollen records preserved in marine and 
continental successions, providing a detailed picture of 
the spatial and temporal modifications of flora, vegeta-
tion and climate (e.g. Bertini, 2003, 2010; Sadori et al., 
2013; Combourieu-Nebout et al., 2015; Magri et al., 
2017 and references therein). By integrating previous 
summaries, the present pollen-based review provides 
an overview of Pleistocene Italian flora, vegetation, and 
climate as well as a coherent and comprehensive histo-
ry of the Italian palaeoenvironments, according to a 
solid stratigraphic framework (Fig. 1; Tab. 1). 

https://doi.org/10.26382/AMQ.2023.05 



   

 

Knowledge of past palaeoenvironments and climate 
context is highly important in a geological perspective 
and is crucial for correctly evaluating the present-day 
ecological and biogeographical patterns; appropriately 
applied, this knowledge can contribute to biodiversity 
conservation and landscape restoration.  

 
 
 

2. PRESENT-DAY CLIMATE AND VEGETATION  
 

Italy extends at mid-latitude (between 47°N and 
35°N) into the Mediterranean Sea, between the Balkan 
and Hellenic peninsula to the east, the Iberian Peninsu-
la to the west, North Africa to the south and continental 
Europe, from which it is separated by the Alps, to the 
north. Three major sectors are recognised (Figs. 1, 2): a 
continental sector (northern Italy), between the Alps to 

Fig. 1 - Location map of selected Piacenzian and Pleistocene marine and continental sites. See also Table 1 for the numbering of the 
sites and additional information. Heavy black dotted line traces the 42.5°N isocline.  

 
 

92 

 
 

Bertini A., Combourien-Nebout N. 



   

 

the north and the conventional line 
connecting La Spezia with Rimini to the 
south (Fig. 1); a peninsular sector 
(central and southern Italy, Fig. 1), 
which stretches into the Mediterranean 
in a north-west-south-east direction; 
and an insular sector encompassing 
the two largest islands of the Mediterra-
nean, namely Sardinia and Sicily. Its 
territory is largely occupied by hills (up 
to ca 42 %) followed by mountain 
(35%), and flat areas (23%). Its highest 
point stands at 4811 m (Monte Bianco, 
Aosta) while the lowest point lies at -
3.44 m (Contane, Ferrara). The Italian 
climate includes, according to the Kö-
ppen climate classification, a humid 
temperate climate (Cfa / Cfb) to the 
north and a Mediterranean climate with 
a dry summer period (Csa) in the cen-
tre-south. A complex climatic gradient 
is very evident moving from the south-
ern warm-temperate Mediterranean 
climate, through the cool-temperate 
climate of the Apennines and pre-
Alpine ranges, to the continental cli-
mate of the innermost Alps with its 
marked temperature seasonality. The 
Italian flora comprises more than 7000 
species (Pignatti, 1982) whose distribu-
tion is strongly connected to the com-
plex topographic and climatic features 
of its territory. Plant biodiversity is very 
high, and endemism is widespread. 
The Italian vegetation has been well 
documented in a large number of publi-
cations (e.g. Tomaselli et al., 1973; 
Ozenda, 1975, 1994; Bonin, 1981; 
Pignatti, 1998; Quézel & Médail, 2003; 
Blasi, 2010). The European potential 
natural vegetation zones and the main 
physiognomic-ecological units for the 
Italian peninsula are summarised in 
Fig. 2 (modified from Pignatti, 2011). 

 
3. MATERIALS AND METHODS  
 

Several Italian marine and continental pollen rec-
ords have been selected for the time interval from the 
Piacenzian to Upper Pleistocene. The geographical 
distribution of the pollen sites is shown in Fig. 1. For 
each site, the main stratigraphical and geological infor-
mation is summarized in Tab. 1 (see also Bertini, 2003, 
2010; Combourieu-Nebout et al., 2015; Magri et al., 
2017, and references therein). The most significant 
pollen evidence for selected single sites is summarized 
in section 4. 

 
3.1. Pollen records  

The abundance of pollen records permits the de-
tection of past environmental changes and allows the 
reconstruction of the history of the flora, vegetation and 
climate throughout the Pleistocene in Italy. The Piacen-

zian is also part of this revision, particularly for docu-
menting the last warm period preceding the installation 
of the G/I cycles at 2.6 Ma. The main palaeoenviron-
mental events, acting at both global and Mediterranean 
scales, are taken into account in the reconstructions. 
Each pollen site and its associated record have been 
assigned to either “Northern” or “Southern” Italy, relative 
to latitude 42.5°N, according to the isocline for current 
dry season duration (longer or shorter than 3 months) 
(Fig. 1- heavy black dotted line). Pollen-inferred vegeta-
tion from available well calibrated records (Tab. 1 and 
Figs. 3-7) is used to illustrate the succession of climatic 
and environmental changes within six key stratigraph-
ical intervals during the last 3.6 Myr: Piacenzian (3.6 - 
2.58 Ma), Gelasian (2.58 - 1.80 Ma), Calabrian (1.80 - 
0.774 Ma), Chibanian (0.774 - 0.129 Ma), and Late 
Pleistocene (0.129 - 0.0117 Ma). In the text 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 

 
 

93 

 

Fig. 2 - Present-day vegetation and climate in Italy (modified from Pignatti, 2011). A 
selection of climate (ombrothermic) diagrams (calculated using the NewLocClim soft-
ware, Grieser et al., 2006) illustrates the modern north-south climate gradient. 



   

 

 

 
 

94 

 
 

Bertini A., Combourien-Nebout N. 



   

 

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Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

“Taxodiaceae” is in quotation marks; many genera for-
merly assigned to Taxodiaceae are now grouped in 
subfamilies of Cupressaceae, for example Taxodioide-
ae Endl. ex K. Koch (Taxodium, Glyptostrobus, and 
Cryptomeria) and Sequoioideae (Luerss.) Quinn 
(Sequoia, Sequoiadendron, and Metasequoia) whereas 
the genus Sciadopitys is now generally placed in Sci-
adopityaceae (Brunsfeld et al., 1994; Farjon, 1998, 
2005). In the Italian pollen records, “Taxodiaceae” are 
mainly represented by Sciadopitys, Taxodium type 
(which includes Taxodium cf. distichum and Glyptostro-
bus) and Sequoia type (which includes Sequoiadendron 
giganteum, Sequoia sempervirens, Metasequoia glyp-
tostroboides, Cunninghamia and Cryptomeria). 

A sketchy reconstruction of vegetation distribution 
is proposed for the Piacenzian to Late Pleistocene, 
according to different transects from non-specific areas 
of high relief to coastal areas (Figs. 3 - 7). The arboreal 
plant cover is in accordance with AP (arboreal plants) / 
NAP (non-arboreal plants) percentages values in pollen 
diagrams. However, the density and distribution of veg-
etation are not based on accurate estimates, they only 
provide a snapshot of vegetation in various areas at 
different times. The distribution of main taxa is illustrat-
ed by informal “plant symbols” and is supplemented by 
the evidence from previous works integrating pollen and 
macroflora (Bertini & Martinetto, 2008, 2011; Com-
bourieu-Nebout et al., 2015).  
 
3.2. Chronology  

Age models for the most recent time intervals are 
based on 14C dates, links to Greenland ice-core records 
and tephra dates. For the geologically older records K/
Ar dates, biostratigraphy (foraminifers and nannofos-
sils), tuning to oxygen isotope records from Mediterra-
nean or Atlantic marine sediment cores and astronomi-
cal tuning were variously used (Tab. 1). The chronologi-
cal framework for terrestrial sites was often provided by 
biochronology (mammal fauna and palaeomagnetism; 
e.g. Napoleone et al., 2003). It is undoubted that the 
chronological control of all published age models is 
quite critical, especially for the older sites (e.g. Sarzana 
and Compiano sites; Fig. 1, Tab. 1). It is not part of our 
review to discuss the individual age models, and in Tab. 
1 all sites are noted whether they have a reliable chro-
nology or not. 

 
4. AN OVERVIEW OF THE ITALIAN POLLEN 
      EVIDENCE 
 
4.1. Piacenzian (3.6 - 2.58 Ma): the last subtropical 
forests (Fig. 3)  

During the Piacenzian a transition from relatively 
warm to cooler climates starts to develop until the in-
stauration of high-magnitude G/I oscillations during the 
Quaternary. Short-lived cooling episodes punctuated 
this interval reflecting a progressive decline of tempera-
ture. Just after glacial marine isotope stage M2, at ∼ 3.3 
Ma, during the last climate phase that was warmer than 
today, known as mid-Piacenzian Warm Period (mPWP; 
e.g., Dowsett et al., 2012; Haywood et al., 2013; De 
Schepper et al., 2013; De La Vega et al., 2020, 2021), 

sea surface temperatures increased by up to ∼3 °C 
(Haywood and Valdes 2004), and CO2 concentrations 
reached values of 400 ppm (Pagani et al., 2009; Bartoli 
et al., 2011), similar to the modern anthropogenic val-
ues. The Italian pollen record is quite remarkable for 
these last 700 kyr of the Piacenzian but it lacks continu-
ous data for the interval spanning the Zanclean-
Piacenzian transition; this is the time when modern Eu-
ropean climates begin to develop at ca 3.6 Ma (e.g. 
Zagwijn & Suc, 1984; Suc et al., 1995) in conjunction 
with a cooling event, and the Brunssumian-Reuverian 
transition (Zagwijn, 1960; North West European Stages-
NWES, Cohen & Gibbard, 2022).  

In central Italy, at Santa Barbara (Upper Valdarno 
intermountain basin) a rich and diverse vegetal assem-
blage (e.g.Taxodium/Glyptostrobus type, Nyssa, Engel-
hardia, Itea, Symplocos, Cephalanthus, Myrica, Carya, 
Quercus, Carpinus, Ulmus, Zelkova, Clethraceae, Cyril-
laceae) is documented during the short-lived ‘warm blip’ 
centred at 3.0 Ma, after the MG2 and M2 cold culmina-
tions. A humid forest vegetation, typical of subtropical to 
warm-temperate climate, dominated swamps in a lacus-
trine system, up to about 2.8 Ma when a cooler forest, 
with Picea and Fagus, as main components, became 
widespread. Herbs including steppe taxa formed a very 
minor component (Napoleone et al., 2003; Bertini, 2010, 
2013). The cooling occurred gradually, in agreement 
with isotopic records (e.g. Raymo et al., 1989) that show 
an increase in glacial intensity over several cycles, ra-
ther than an abrupt increase at 2.6 Ma. A permanent/
temporary disappearance and/or the significant reduc-
tion of some taxa (e.g. Symplocaceae, Myrica, Nyssa, 
Cephalanthus, “Taxodiaceae”), points to a progressive 
floristic impoverishment in many sites of both central 
(e.g. Lower Valdarno: Valleri et al., 1990, Benvenuti et 
al., 2007; and Tiberino basin: Pontini & Bertini, 2000, 
Pontini et al., 2002, Martinetto et al., 2014) and northern 
(Villafranca: Francavilla et al., 1970, Lona & Bertoldi, 
1972, Carraro et al., 1996; Stirone: Bertini, 2001; Val 
Marecchia: Rio et al., 1997; M. Falcone-Rio Crevalese: 
Monegatti et al., 2002) Italy. The decrease in tempera-
ture, associated with the forthcoming onset of the North 
Hemisphere Glaciation, resulted in a gradual increase in 
relatively more resilient arboreal plants such as Quercus 
and Carya, which progressively replaced Taxodium/
Glyptostrobus type and Engelhardia (Bertini, 2001; Fau-
quette & Bertini, 2003).  

In southern Italy, the Piacenzian is only represent-
ed by rather short and discontinuous pollen records. At 
3.31 Ma (close to the FAD, first appearance datum, of 
Globorotalia bononiensis) on the site of Bianco, a cool, 
humid climate (with decreased seasonality) has been 
reconstructed during a short-term forest phase dominat-
ed by the Picea-Abies group, while Mediterranean ever-
green virtually disappeared and non-arboreal cover 
declined (Bertoldi et al., 1989). Between 3 and 2.92 Ma, 
Marine Isotope Stage (MIS) 108 to 104 (Combourieu-
Nebout et al., 2004), at Punta Piccola, a well-diversified 
pollen flora suggests an altitudinal zonation of the vege-
tation and several climate fluctuations during precession 
cycles illustrated in laminites/carbonates couplets. Cli-
mate reconstructions from the high resolution pollen 

 
 

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97 

Fig. 3 - The Piacenzian. On the left (top to bottom): location map of the Piacenzian sites (for numbers, see also Tab. 1), list of main taxa 
now extinct in Italy and main vegetation groups during the 21 to 41 kyr cycles in North and South Italy. On the right the sketchy vegeta-
tion profiles for North and South Italy.  

Fig. 4 - The Gelasian. On the left (top to bottom) location map of the Gelasian sites (for numbers, see also Tab. 1), list of main taxa now 
extinct in Italy and main vegetation groups during the 41 kyr cycles in North and South Italy. On the right the sketchy vegetation profiles 
for North and South Italy.  

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

record indicate that high temperatures and relatively 
intense precipitation prevailed during the deposition of 
brown laminated sediments; in contrast, arid conditions 
are associated with light carbonates. Still in Sicily, just 
before 2.5 Ma, at the site of Capo Rossello, an expan-
sion of Mediterranean formations alongside the pres-
ence of open vegetation (including steppe and psam-
mophilous taxa), is documented during a general de-
crease in humidity and the establishment of drier condi-
tions during the warm season (Guerrera et al., 1984; 
Bertoldi, 1985a; Bertoldi et al., 1989; Rio et al., 1990).  

At Stirone in northern Italy, Climate Amplitude 
Method (CAM) reconstructions are only available for the 
last 200 ka of the Piacenzian (Fauquette & Bertini, 
2003). They indicate alternating warm/cold phases with 
temperature and precipitation values very close to those 
associated with G/I cycles during the following Gelasian 
(see in 4.2.1).  

 
4.2. The Pleistocene  
4.2.1. Gelasian (2.58-1.80 Ma): the decline of sub-
tropical trees and the onset of G/I cycles at the 
beginning of the Quaternary (Fig. 4) 

The intensification of the Mediterranean outflow 
and the establishment of the Northern Hemisphere ice 
cap from ∼ 3.0 Ma (e.g., Marlow et al., 2000; Lisiecki & 
Raymo, 2007) promoted cooler, more seasonal and in 
some areas dryer conditions in the Mediterranean area 
(Lourens et al., 1996, 2004). In the Italian pollen rec-
ords, 40 kyr obliquity-forced G/I cycles are predominant-
ly characterized by the contrast between forest vegeta-
tion and steppes (e.g. Bertini, 2010; Combourieu-
Nebout et al., 2015 and references therein). This pat-
tern is especially well attested in the Semaforo section 
(Crotone, Calabria; e.g. Combourieu-Nebout & 
Vergnaud-Grazzini, 1991; Combourieu-Nebout, 1993, 
1995; Lourens et al., 1996) by both pollen and oxygen 
isotope data, from 2.46 Ma onwards. Nevertheless, in 
other southern Italian sites, Bertoldi et al. (1989), have 
highlighted a totally different (opposite) response of the 
vegetation to G/I cycles, still on the basis of the biostrat-
igraphic and isotopic evidence. In the Gelasian (-
Calabrian) Le Castella and Gela (Monte San Nicola) 
sites, glacials would have been marked by forests 
(humid conditions), and interglacials by open vegetation 
(dry conditions). A different scenario emerges from the 
Northern Italian pollen records (Castell’Arquato: Lona, 
1962; Lona & Bertoldi, 1972; Stirone: Lona & Bertoldi, 
1972; Bertolani Marchetti et al., 1979; Bertini, 2001; Val 
Marecchia: Rio et al., 1997; M. Falcone-Rio Crevalese: 
Monegatti et al., 2002). Here G/I cycles are apparently 
characterized by contrasts between the subtropical to 
warm temperate forest and higher-elevation coniferous 
forest (mainly Picea) without any significant extension in 
space and time of steppe-like vegetation (minimum 
values of humidity). Herb percentages usually remain 
low although Artemisia, a steppe taxon, can increase 
but not significantly and according to the specific geo-
graphical locations of the above mentioned sites. In the 
Upper Valdarno (central Italy), the location of the Pia-
cenzian Santa Barbara section, we find evidence for the 
first substantial spread of herbs, which include a very 

large component of Artemisia (Bertini, 1994, 2010), 
within a distinctive aeolian-dominated sandy deposit 
(Rena Bianca section), 35 m thick. On the other hand, 
in the Tiberino pollen record, which falls within the Pia-
cenzian-Gelasian transition (Pontini, 1997; Pontini & 
Bertini, 2000; Pontini et al., 2002; Martinetto et al., 
2014), the vegetation response to the G/I cycles is ex-
pressed by the competition between coniferous forests 
dominated by Picea and thermophilous arboreal taxa 
including “Taxodiaceae” (MIS 100 to MIS 82). During 
the Gelasian the flora suffers a very evident and pro-
gressive impoverishment. In northern Italian pollen 
sites, the first notable reduction in “Taxodiaceae” (with 
Taxodium/Glyptostrobus type as the main component) 
as well as an increase of Cathaya are evident from ca. 
2.7 Ma (at Stirone). In the south, at Semaforo, where 
the record starts at 2.48 Ma, the first significant demise 
of “Taxodiaceae” forests (with Sequoia type as main 
component) is recorded from about 2.38 Ma whereas 
the spread of Cathaya occurs, especially from ca. 1.92 
to 1.74 Ma (Combourieu-Nebout & Vergnaud-Grazzini, 
1991; Combourieu-Nebout, 1993; Klotz et al., 2006; Suc 
et al., 2010). 

CAM reconstructions from Stirone (northern Italy) 
(Fauquette & Bertini, 2003) show that during the Gela-
sian warm phases, the climate was similar to that of the 
Zanclean, with high MAT, MAP and available moisture. 
Annual temperatures were almost equal to the modern 
value (around 12-13 °C) during glacials whereas precip-
itation values were between 200 and 500 mm higher 
than today.  

In central Italy, at Poggio Rosso (Upper Valdarno) 
dating to a G/I cycle just before the Gelasian/Calabrian 
boundary, the CAM method was used to calculate mean 
annual temperatures of around 12.5-14 °C and mean 
annual precipitation of around 800 mm, with a minimum 
of 400 mm during the glacial phase and 15/16-19 °C 
and 750 -1200 mm during the interglacial (Bertini et al., 
2010). In southern Italy, at Semaforo, the Probability 
mutual Climatic Spheres (PCS) method reveals mean 
annual temperatures and winter temperatures from ca. 
2.46 Ma to ca. 2.11 Ma (Klotz et al., 2006) that were at 
least 2.8 °C and 2.2 °C warmer, respectively, than to-
day; it also reveals that annual precipitation during inter-
glacials exceeded today’s values by 500 mm. During 
glacials, temperatures were lower than the present-day, 
but precipitation was similar. A trend of reduction in 
annual and winter temperatures (more than 2.3 °C), as 
well as greater seasonality, were also highlighted over 
the course of the consecutive interglacials. A significant 
reduction of at least 1.6 °C in all temperature parame-
ters is also evident during the glacials.  

 
4.2.2. Calabrian (1.80-0.774 Ma): the demise of sub-
tropical ecosystems from the Mediterranean and the 
beginning of the Early-Middle Pleistocene transition 
(Fig. 5) 

From 1.4 Ma, over the course of the Calabrian, a 
major global climatic reorganization starts to affect 
ocean and atmospheric circulation, ice sheets and the 
distribution and evolution of biota, including the ances-
tors of modern humans (Head & Gibbard, 2015). During 

 
 

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this phase, termed the Early-Middle Pleistocene transi-
tion (EMPT) which extends up until 0.4 Ma, low-
amplitude 41-ka obliquity-driven climatic cycles of the 
earlier Pleistocene were progressively superseded by 
100 kyr fluctuations in the later Pleistocene. A succes-
sion of severe glacial episodes (i.e. MIS 36, 24, 22) 
occurred, particularly in the northern hemisphere. A rich 
and detailed palynological record, often very well cali-
brated, is available for southern Italy in particular. In the 
Crotone basin (Calabria) the most significant pollen 
sites are: Vrica (ca. 2.2-1.36 Ma; e.g. Combourieu-
Nebout et al., 1990), Santa Lucia (1.356-1.24 Ma, MIS 
43-40; Joannin et al., 2007), and Valle di Manche (0.87-
0.73 Ma, MIS 22-18.3; Capraro et al., 2005, 2015, 2017, 
2022). Moreover, in Basilicata some 150 km north of the 
Crotone basin, the marine Montalbano Jonico (1.24-0.9 
Ma and MIS 37-MIS 23: Joannin et al., 2008; 0. 82-
0.774 Ma and MIS 20-19 p.p.: Bertini et al., 2015; Toti, 
2018; Toti & Bertini, 2018; Nomade et al., 2019;) and 
the neighbouring continental Sant’Arcangelo (San Lo-
renzo cycle: ca. 1.1-0.8 Ma; Sabato et al., 2005) suc-
cessions were also studied. In Sicily, data for the 1.23-
1.095 Ma interval are available from the Monte San 
Giorgio section (Caltagirone; Dubois, 2001). In the up-
permost part of these sites, G/I fluctuations are well 
illustrated by open vegetation and forest alternations. 
The open vegetation assemblages include a large 
amount of steppe taxa, including Artemisia and Ephed-
ra, and sometimes also thermophilous taxa, such as 
Cistus and Phlomis fruticosa. Warm temperate forest 

assemblages include mainly deciduous taxa, such as 
Quercus, Carya, Carpinus, Pterocarya, Ulmus and 
Zelkova after the progressive reduction and disappear-
ance of the most thermophilous taxa, e.g. 
“Taxodiaceae”. However, according to Capraro et al. 
(e.g. 2005, 2015), in the marine Valle di Manche suc-
cession, the glacial interval MIS 20 (as well MIS 18) is 
characterized by an alpine type forest (Picea, Abies, 
Fagus), whereas a wooded steppe landscape was es-
tablished only later (i.e. at the Matuyama-Brunhes 
boundary), during the Early-Middle Pleistocene climatic 
transition, at the beginning of the deglaciation from MIS 
20 to MIS 19. Another very different scenario is ex-
pressed by the pollen records from northern Italy 
(Lamone, Stirone, Leffe, Pianengo). Here, glacial phas-
es are, as in the Gelasian, often marked by expansions 
of coniferous forests dominated by Picea, although in-
terglacials do not exhibit major expansions of steppe 
vegetation but rather thermophilous forests as de-
scribed in the classic schemes of Combourieu-Nebout 
(1993) and Fusco (2007). In the Lamone marine suc-
cession three distinct forest formations, i.e. mixed-
deciduous forest (with Quercus dominating), Juglan-
daceae forest (with Carya dominating) and mountain 
coniferous forest (with Picea dominating) expanded 
between MIS 64-46 (Fusco, 2007, 2010). A complete 
vegetation cycle only developed when forest dynamics 
were interrupted by the spread of open vegetation, indi-
cating a decrease in humidity (Fusco, 2007). Flora con-
tinued to be depleted as documented, in more detail, in 

 
 

99 

 

Fig. 5 - The Calabrian. On the left (top to bottom), location map of the Calabrian sites (for numbers, see also Tab. 1), list of main taxa 
now extinct in Italy and main vegetation groups during the 41 kyr cycles in North and South Italy. On the right the sketchy vegetation 
profiles for North and South Italy.  

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

both Bertini (2010) and Magri et al. (2017) papers which 
also traces the abundance, reduction, and disappear-
ance of certain key taxa (e.g. “Taxodiaceae”, Cathaya, 
Tsuga, Carya, Pterocarya) in selected sites (e.g. Vrica, 
Saticula, Madonna della Strada, Colle Curti, Formaci di 
Ranica, Pianengo, and Piànico-Sèllere). The disappear-
ance of the subtropical ecosystems and their replace-
ment by deciduous forest taxa was identified in the 
Mediterranean area at ca. 1.2 Ma (Combourieu-Nebout 
& Vergnaud-Grazzini,1991; Combourieu-Nebout, 1993, 
1995) i.e. after the appearance of the large Gephy-
rocapsa a coccolith (at about 1.56 Ma), and the fora-
minifer Hyalinea baltica (at about 1.49 Ma).  

Climate reconstructions, using the pollen data, 
indicate an enhanced cooling between 1.4 and 1.3 Ma, 
with winter temperatures decreasing at both interglacial 
maxima and glacial minima (Combourieu-Nebout et al., 
2015). During the same interval, annual precipitation 
during both interglacial and glacial stages decreased 
marking a progressive increase in aridity, especially in 
glacial times.  
 
4.2.3. The Chibanian (0.774-0.129 Ma): the 
development of temperate broadleaf and mixed 
forests in full EMPT (Fig. 6)  

In the Mediterranean area, the progressive floristic 
impoverishment, according to climate gradients, contin-
ues during the upper portion of the Early-Middle Pleisto-
cene climate transition, which also includes some se-
vere glacial episodes (i.e. MIS 12 and 6). The 41 kyr G/I 
cycles were replaced by 100 kyr cycles, with lengthen-
ing of glacial stages (Tzedakis, 2005, 2007; Leroy, 
2007; Tzedakis et al., 2012). However, from MIS 11 
onwards, an abrupt increase in both interglacial temper-
atures and in reconstructed interglacial CO2 values 
(Masson-Delmotte et al., 2010; Lang and Wolff, 2011; 
McClymont et al., 2013 and references therein) is re-
vealed by marine, terrestrial and ice-core records during 
the so-called Mid-Brunhes Event (MBE).  

Early Chibanian pollen records from southern sites 
attest to widespread alternations between Artemisia 
(plus Ephedra) steppes and temperate to warm-
temperate deciduous forests, during the still dominant 
41 kyr G/I cycles. At Montalbano Jonico, clear expan-
sions of steppe taxa are documented, during both glaci-
als and stadials from MIS 19 and the base of MIS 16 
(e.g. Bertini et al., 2015; Toti, 2018; Toti & Bertini, 2018; 
Nomade et al., 2019). At Valle di Manche, MIS 18 is 
again (as well MIS 20) characterized by an alpine type 
forest (Picea, Abies, Fagus) according to Capraro et al. 
(2005, 2015). At Piànico Sèllere in northern Italy 
(Prealps, Lombardy) pollen data document a long inter-
glacial phase within MIS 19, dated by the K/Ar age of a 
tephra level (779±13 ka; Pinti et al., 2001) and palaeo-
magnetic data (Scardia & Muttoni, 2009) within biogenic 
varved sediments. Here the onset of the interglacial is 
marked by the expansion of broad-leaved deciduous 
thermophilous forests with Quercus, Ulmus, Tilia and 
Fraxinus, followed by the expansion of Abies and 
Carpinus betulus. The interglacial succession is inter-
rupted by a short phase characterized by the almost 
complete disappearance of thermophilous trees and the 

expansion of conifer forests (Picea, Pinus, Larix) and 
herbaceous steppe communities (Ravazzi, 2003; Pini et 
al., 2014). A generalized drop in temperature in both 
steppe and forest phases is indicated by changes in the 
floristic assemblages; for example, Hippophaë rham-
noides expanded during the steppe phases (“cold 
steppes”), whereas thermophilous taxa, such as Cistus 
and Phlomis fruticosa, disappeared. The best calibrated 
pollen sites permitted the tracing of the chronological 
distribution of some taxa, notably of “Taxodiaceae”, 
Carya, Pterocarya and Tsuga, including their LO (last 
occurrence) or LCO (last common occurrence). In north-
ern Italy, Tsuga is virtually absent at Pianengo, from the 
late Early Pleistocene, and only sporadic pollen grains 
(0.2-0.8%) were recovered during the early Middle Pleis-
tocene (Muttoni et al., 2003). In central Italy Tsuga de-
clined during the lower part of the Brunhes (Bertini, 
2000); here, both Carya and Pterocarya are absent. At 
Torre Mucchia (Abruzzi) Tsuga is still present (at >3%) 
during MIS 17 according to Pieruccini et al. (2016). At 
Rignano Flaminio (Latium) in a diatomite deposit at-
tributed to MIS 13, Tsuga is absent whereas Carya and 
Pterocarya are sporadic (Di Rita & Sottili, 2019). At the 
archaeological site of Ceprano, just below the stratum 
including hominin remains, pollen analyses permitted a 
detailed picture of interglacial vegetation development 
during MIS 13 (Margari et al., 2018). Here Tsuga is ab-
sent and Cedrus sporadic, whereas Carya exhibits a 
continuous presence. Moving further south, Tsuga is 
present at Valle di Manche until MIS 18 (ca. 0.73 Ma) 
but with values that never exceed 5%; Taxodium, Carya, 
Pterocarya, Liquidambar and Cedrus are also present in 
low percentages; the LO of Carya, specifically, has been 
recorded between MIS 18 and the successive wooded 
steppe expansion. At Vallo di Diano, between 0.65-0.45 
Ma (MIS 16 to 13) Tsuga is virtually absent (as well as 
at Isernia la Pineta; Lebreton, 2002) whereas Carya is 
constantly present, and Pterocarya is sporadic (Russo 
Ermolli, 1994). At Sessano, in contrast, Tsuga, along 
with Cedrus, Zelkova as well as a few grains of Ptero-
carya and Taxodium type have been recorded between 
ca 0.58 and 0.44 Ma (MIS 15-12) (Russo Ermolli et al., 
2010a). Both Tsuga and Carya are absent in the Mer-
cure Basin (MIS 13; Petrosino et al., 2014b) as well as 
in the lacustrine succession of Acerno, where pollen 
analyses documented an interglacial/glacial cycle corre-
lated with MIS 14 and 12 (Petrosino et al., 2014a). At 
Boiano, Tsuga is absent whereas Carya is present until 
the MIS 9 which, at present, represents its latest occur-
rence in Western Europe (Orain et al., 2013, 2015). 
According to previous data, Tsuga apparently disap-
peared from southern Italy later than in the north. How-
ever, in the southern assemblages its presence is usual-
ly subordinate, at least since the Messinian (Suc & Bes-
sais, 1990; Bertini et al., 1998). Bertini (2000), on the 
one hand, related the disappearance of Tsuga to the 
shift in global aridity as well as the progressive decrease 
in temperature associated with the Middle Pleistocene 
shift from the 41 to 100 ka cyclicity in the Milankovitch 
orbital record (Bertini, 2000). Ravazzi et al. (2005), on 
the other hand, linked the Tsuga decline (at Fornaci di 
Ranica) to catastrophic fires triggered by summer 

 
 

100 

 
 

Bertini A., Combourien-Nebout N. 



   

 

 

 
 

101 

Fig. 6 - The Chibanian. On the left (top to bottom), location map of the Chibanian sites (for numbers, see also Tab. 1), list of main taxa 
now extinct in Italy and main vegetation groups during the 41 to 100 kyr cycles in North and South Italy. On the right the sketchy vegeta-
tion profiles for North and South Italy.  

Fig. 7 - The Late Pleistocene. On the left (top to bottom), location map of the Late Pleistocene sites (for numbers, see also Tab. 1), list of 
main taxa now extinct in Italy and main vegetation groups during the 100 kyr cycles in North and South Italy. On the right the sketchy 
vegetation profiles for North and South Italy. 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

drought. However, both hypotheses confirm the global 
shift toward aridity of the Early Middle Pleistocene cli-
mate transition at ca. 0.9 Ma (Ruddiman et al., 1989; 
Raymo et al., 2004). 

Rare climate data from pollen records provide 
evidence for increasing drought during increasingly and 
longer glacials, while the shorter interglacial began to 
become cooler (Combourieu-Nebout et al., 2015). 
 
4.2.4. The Late Pleistocene (~0.129-0.0117 Ma): 
towards the present under the effects of the sub-
Milankovitch-scale climate variability (Fig. 7) 

The reconstruction of millennial and sub-millennial 
fluctuations during the last G/I cycles and the associat-
ed floristic and vegetational changes before the main 
anthropic interferences have mainly been based on the 
pollen records, especially lacustrine ones, from northern 
(Azzano Decimo and Fimon; Pini et al., 2009a,b; 2010, 
2022; Badino et al., 2020a,b; Venice pro parte: Mullen-
ders et al., 1996, Massari et al., 2004, Canali et al., 
2007), central (Valle di Castiglione: Follieri et al., 1988; 
Lagaccione: Magri, 1999; Vico: Leroy et al., 1996, Magri 
& Sadori, 1999) and southern (Laghi di Monticchio: e.g. 
Brauer et al., 2007a) Italy. However, a number of ma-
rine sedimentary successions were also retrieved from 
cores in the Mediterranean area (Figs. 1, 7; Tab. 1). 

Recurrent forest and open vegetation develop-
ments as well as vegetation dynamics, together with 
repetitive changes in the vegetation structure, flora 
composition and plant biomass, are documented by the 
mainly continuous high resolution pollen records. Forest 
phases developed during the Eemian according to a 
dominant vegetation pattern and with the presence of a 
small number of taxa now extinct in Italy (e.g. Pterocar-
ya and Cedrus; Magri et al., 2017). In fact, most of the 
changes in floral composition occurred in earlier times. 
In contrast, steppe phases, usually characterized by the 
same flora (e.g. Artemisia, Poaceae, Amaranthaceae), 
are generally less typical with respect to the interglacial 
ones. 

At Azzano Decimo (north-eastern Italy), during the 
last 0.215 Ma the mixed temperate forests persisted 
during MIS 7a-7c, whereas they were replaced by the 
conifer forests after the local glacio-eustatic regression, 
during early MIS 6 (Pini et al., 2009a). The Azzano Dec-
imo Eemian forest record, though incomplete, indicates 
the absence of Mediterranean evergreen trees, in ac-
cordance with the Lake Fimon sequence. Here, mixed 
oak forests characterized the first part of the Last Inter-
glacial; these were then replaced by oceanic mixed 
forests that persisted for a further 7 ka until the end of 
the Eemian succession (Pini et al., 2010). At Azzano 
Decimo, repeated stadial-interstadials are evident since 
ca 0.115 Ma. A correlation has been proposed between 
major xerophyte peaks and IRD (Ice Rafted Debris) 
maxima during Heinrich events in deep-sea cores off-
shore Iberia and in the North Atlantic. As a conse-
quence, the interstadial phases, marked by Picea 
peaks, were interpreted as being linked to Dansgaard-
Oeschger warm events (Pini et al., 2009a). The disap-
pearance of broad-leaved thermophilous forests from 
the north-eastern plain of Italy was placed at the end of 

the Early Würm with cool-temperate mixed forests per-
sisting in the southern Alpine foothills up until the onset 
of Heinrich stadial 4 as documented in the Lake Fimon 
record (Pini et al., 2010).This high-resolution multiproxy 
record (including pollen and dust flux) from Lake Fimon 
(Pini et al., 2022) provided climate evidence of the MIS 
3-2 interval with a focus on the dynamics of the Alpine 
Last Glacial Maximum and the early Alpine Lateglacial, 
including the expression of Heinrich stadial 2 and 1, on 
land. The onset of the Last Glacial Maximum at 27.3 ka 
was associated with a 4 °C drop in July temperatures, 
likely responsible for the onset of glacial build-up in the 
Southern Alpine piedmont (Pini et al., 2022).  

In central Italy, the integrated pollen records from 
the crater lakes of Valle di Castiglione, Lagaccione and 
Lago di Vico document the history of vegetation cover-
ing the last two interglacials (last 0.3 Ma, from MIS 7e). 
The evidence for floral and vegetation changes allows 
the most important events e.g. Eemian, Saint Germain I 
and II and the Pleniglacial Interstadials (Lazio complex) 
(Follieri et al., 1998) to be recognized. At Valle di 
Castiglione, the Eemian is characterized by the large 
scale expansion of thermophilous and Mediterranean 
vegetation (Follieri et al., 1988, 1998). The Saint Ger-
main I forest phase at both Valle di Castiglione and 
Lagaccione is expressed by the expansion of Fagus 
and Abies. The subsequent steppe period is character-
ized by the almost total absence of trees with the ex-
ception of Pinus. Due to a core gap, the Saint Germain 
II Forest phase, is not recorded at Valle di Castiglione, 
whereas it is well characterized at Lagaccione and Lago 
di Vico by the consistent spread of Abies. During the 
following Glacial and Lateglacial phases, Artemisia, 
Poaceae, and Amaranthaceae alternately dominated. 
These steppe and grassland formations were interrupt-
ed during the pleniglacial (roughly corresponding to MIS 
3) by a number of slight expansions of tree cover, in 
particular of mesophilous and thermophilous taxa; this 
is termed the “Lazio complex” by Follieri et al. (1998) 
and probably corresponds to the Heinrich events found 
in oceanic cores. At Monticchio (southern Italy), varve 
chronology indicates the onset of the sedimentation at 
about 0.133 Ma (MIS 6 to present, Allen & Huntley, 
2009 and references therein). Regional vegetation was 
dominated by herbs with steppe taxa as the main com-
ponents during MIS 6, 4, and 2. During Melisay 1 only, 
the decrease of steppe taxa together with an increase in 
Betula probably suggest that, although cold, this period 
had greater moisture availability than other cold periods. 
Mesic woody taxa reached moderate abundance during 
a large number of MIS 3 interstadials, but dominated 
during the Saint Germain I and II and Eemian. Palaeo-
climatic reconstructions as well comparisons with the 
main European sites reveal significant complexity in the 
geographical and altitudinal environmental patterns 
(Allen & Huntley, 2000, 2009). 

Pollen-based climate reconstructions for Southern 
Italian sites (Combourieu-Nebout et al., 2015) show that 
the Mean Temperature of the Coldest Month (MTCO) 
during marine isotope sub-Stage 5e was nearly as 
warm as during the Early Pleistocene interglacials. Nev-
ertheless, annual precipitation remained low compared 

 
 

102 

 
 

Bertini A., Combourien-Nebout N. 



   

 

to the earlier interglacials. During glacials, both temper-
ature and precipitation values were very low. This is 
probably linked to recurrent Mediterranean cooling, 
along with increased aridity on the continent induced by 
the global climate effects of Heinrich event discharges 
in the North Atlantic, as is also attested by palynological 
studies on the Spanish and Portuguese margins (e.g. 
Combourieu-Nebout et al., 2002; Sánchez Gòni et al., 
2002). 

 
5. DISCUSSION 
 

The present overview of Italian Pleistocene pollen 
records includes a total of 66 selected pollen sites (with 
12 also providing information for the Piacenzian) stud-
ied since 1950 (i.e. Lona, 1950) (Fig. 8). A large number 
of continental (including freshwater terrestrial car-
bonates such as travertine, calcareous tufas and flow-
stone) (38) and marine (28) sedimentary successions 
(sites) are presented, as well as a few sedimentary 
successions from depositional transitional environments 
(Tab. 1). Pollen records are especially abundant for the 
Calabrian and Chibanian. The majority of marine pollen 
records refer to the Gelasian when we exclude the off-
shore sites from the Late Pleistocene; most sites are 
located on the south. Continental records refer especial-
ly to Chibanian; again, southern sites are the most 
abundant. Different vegetational belts, at different alti-
tudes between sea level and 850 m a.s.l. (Colfiorito 
basin: Colle Curti - Cesi composite section; Bertini, 
2000) and between the latitudes 45 and 36 °N, have 
been documented.  

 
5.1. Flora and vegetation  

The Piacenzian and Pleistocene pollen floras in-
clude six main “formation classes”, informal biome/
vegetation units (Bertini, 2001), i.e. Tropical forest (only 
some rare, scarce taxa, especially present in southern 
sites), Subtropical humid forest, Temperate broad-
leaved deciduous forest, Sclerophyll forest, Mid- to high

-elevation forest, Open vegetation. In Tab. 2 these have 
been organized in 13 informal pollen groups. Their main 
components as well as their associated climatic and 
ecological features, are also summarized in Bertini & 
Martinetto (2008). Over the last 3.6 Ma, the composition 
of such “formation classes” has changed due to the 
gradual disappearance of some of their main compo-
nents. The analysis of the stratigraphical distribution of 
taxa has highlighted frequent asynchronous disappear-
ance events determined by the occurrence of climate 
gradients across the Mediterranean (e.g. Bertini, 2010; 
Combourieu-Nebout et al., 2015; Magri et al., 2017). 
Some taxa (e.g. Engelhardia, “Taxodiaceae”, Cathaya, 
Tsuga, Carya, Cedrus, Liquidambar, Symplocos, 
Nyssa, Pterocarya, Embolanthera, Distylium, Parroti-
opsis, Parrotia persica) attained the status of relict un-
der particular microclimatic and (or) edaphic conditions 
before their definitive extinction under the effects of the 
progressive decrease of temperature and changes in 
the amount and distribution of precipitation. The floristic 
and vegetation changes, affecting the late Pliocene and 
Pleistocene Italian terrestrial ecosystems, in particular 
the first significant spread of steppe formation class 
close to 2.6 Ma and the disappearance of the 
“subtropical humid forest” formation class close to 1.2 
Ma, contributed to the establishment of the present 
complex mosaic of different climates/biomes in the 
Mediterranean area (Tab. 3). 
 
5.2. Pleistocene Glacial/Interglacial cycles: an 
overview of the main vegetation patterns at the 
Milankovitch-scale climate variability  

Four (1-4) main vegetal patterns associated with 
G/I cycles have been identified on the basis of the avail-
able Italian Pleistocene pollen record (Bertini, 2010) 
(Fig. 9, Tab. 3).  

Pattern 1 is expressed by fluctuations between 
open vegetation and thermophilous forests. It is usually 
considered as the most typical and representative pat-
tern for the Mediterranean littoral (e.g. Combourieu-
Nebout & Vergnaud-Grazzini, 1991; Suc et al., 1995a). 
During the first Quaternary cycles, it embraces four 
main subzones (a-d in Combourieu-Nebout, 1993): (a) 
mixed temperate forest (transitional phase following a 
glacial, with increasing temperatures but still dry condi-
tions), (b) sub-tropical humid forest (interglacial with 
maximum temperature and precipitation), (c) high-
altitude coniferous forest (transitional phase with cooler 
but still humid conditions), and finally (d) expansion of 
open vegetation, including steppe taxa (full glacial 
phase with cooler and dry conditions). Sometimes the 
first (a) or the last (d) one can be absent, as described 
by Combourieu-Nebout (1993) at Semaforo. During the 
late Pleistocene the pattern shows the following four 
main subzones: (a) pioneer forest, (b) mixed temperate 
forest (interglacial with maximum temperature and pre-
cipitation), (c) high-altitude coniferous forest (transitional 
phase with cooler but still humid conditions), and finally 
(d) expansion of open vegetation. 

Pattern 2 is expressed by fluctuations between 
altitudinal coniferous forests and thermophilous forests; 
it was observed in many records from the Northern Ap-

 
 

103 

Fig. 8 - The selected Piacenzian and Pleistocene pollen rec-
ords summarized according to chronology, geographical loca-
tion (North and South) and type of sedimentary succession 
(marine vs terrestrial).  

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

 

 
 

104 

 
 

Bertini A., Combourien-Nebout N. 

Tab. 2 - Summary information on Piacenzian and Pleistocene pollen flora and vegetation. From the left: informal vegetation groups, main 
floristic taxa, their modern and climate ecology, vegetation images, optical microscope images of selected fossil pollen grains (not to 
scale).  



   

 

 

 
 

105 

  
 
 
 
 
 
 
 
 
 
 
 
 
 
Fig. 9 - Glacial/Interglacial cycles: sum-
mary of vegetation and climate evidence. 
Pattern 1: Gelasian-Calabrian theoretical 
cycle. Pattern 1’: end of Chibanian and 
Late Pleistocene theoretical cycle. Pat-
tern 2: Northern Apennines (e.g. Lona & 
Bertoldi, 1972; Bertini, 2001). Pattern 3a 
(e.g. Bertoldi et al., 1989) and Pattern 3b 
(e.g. Capraro et al., 2005) both described 
for the southern Italian sites. See text for 
detail. 

Tab. 3 - Main floristic, vegetal and climate evidence according to Glacial/Interglacial phases between the late Piacenzian and the Late 
Pleistocene. 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

ennines (e.g. Stirone, Castell’Arquato, Monte Falcone-
Rio Crevalese, Val Marecchia). When comparing the 
theoretical cycle of Semaforo with those described for 
Stirone (Bertini, 2001) and for Lamone (Fusco, 2007), 
pattern 2 seems to correspond to pattern 1 when its 
subzone (d) is lacking or extremely reduced.  

Patterns 3a and 3b are both expressed by fluctua-
tions between thermophilous forest and open vegetation 
describing humid glacials and dry interglacials accord-
ing to the pollen records from some southern sites (e.g. 
Bertoldi et al., 1989; Capraro et al., 2005), as already 
addressed in 4.2.1-4.2.3. According to this evidence, 
steppe (i.e. arid conditions) should not be connected to 
glacial maxima without independent supporting data. 
Consequently, the role of orbital signatures on long-
term vegetation patterns in the Mediterranean needs to 
be attentively evaluated before proposing terrestrial 
chronologies through astronomical tuning of vegetation-
al phases. An explanation for patterns 3a and 3b may 
lie in the interplay between local factors and the effects 
of obliquity related warm/humid-cold/dry “interglacial”- 
“glacial” cycles superimposed by precession related 
warm/dry-cold/humid cycles, as proposed by Klotz et al. 
(2006) and Joannin et al. (2007). 
 
5.3. Pollen and Quaternary stratigraphy 

Italy is very rich in late Neogene and Quaternary 
sites with many of them having a significant role in 
Chronostratigraphy. Fig. 10 shows the GSSP from 3.6 
Ma to 0.0117 ka and the main coeval Italian pollen 

sites; the locations of some of them correspond exactly 
with a type-locality (in bold in the figure, e.g. Vrica). 
Palynology is a powerful tool not only in the area of 
palaeoclimatic and palaeoenvironmental reconstructions 
but also with respect to the stratigraphic issues. Howev-
er, this requires appropriate application of the strati-
graphic principles in the interpretation of the significance 
of pollen zones. The fact that it is almost impossible to 
identify first appearance events, as well as to exactly 
trace the time-transgressive extinction of pollen taxa, 
complicates the use of the pollen content in describing 
biozones (except for assemblage and acme zones) or 
climatostratigraphic units, especially when they lack a 
solid chronostratigraphic framework. The ecobiostrati-
graphic events associated with the Pliocene and Pleis-
tocene pollen zones need to be integrated with time-
diagnostic (e.g. magnetostratigraphy and tephrostratig-
raphy) and/or other biological marine or continental 
evidence in order to render them chronologically signifi-
cant. In the past, palynostratigraphic correlations were 
often established according to the idea that the same or 
similar pollen (assemblage) zones in different sites can 
identify synchronous pollen signals, e.g. those based on 
the disappearance of “Taxodiaceae” (i.e. Tiberian phase 
and boundary; Lona et al., 1969; Lona & Bertoldi, 1972), 
or those based on the Northern European pollen stratig-
raphy (e.g. Zagwijn, 1957, 1959, 1960, 1963, de Jong, 
1988). Such interpretations, which assigned chronos-
tratigraphic significance to ecobiostratigraphic events, 
produced a clear misunderstanding between biostrati-

 
 

106 

 
 

Bertini A., Combourien-Nebout N. 

Fig. 10 - Piacenzian and Pleistocene GSSP. On the right, the stratigraphical distribution of selected pollen sites corresponding or correla-
tive to the GSSP locations. (Modified from Cohen et al., 2013; updated). The ICS International Chronostratigraphic Chart. Episodes 36, 
199-204 (online version 2022/10). 

 



   

 

graphic and chronostratigraphic principles and conse-
quent mistakes in the establishment of correlations, also 
at regional scale, between different sites. For example, 
the “Tiberian boundary (and phase)” was identified in 
the lacustrine Pietrafitta succession with the sudden 
disappearance of Taxodium pollen type directly after an 
acme phase according to Lona et al. (1969) and Lona 
(1971). Previous authors also postulated the coinci-
dence of the Tiberian boundary with the former Plio-
Pleistocene boundary (1.8 Ma). Later, the Tiberian 
boundary was also recognized in the marine section of 
Stirone (Lona et al., 1969, Lona & Bertoldi, 1972) and at 
the marker bed, in the controversial marine section of 
Le Castella (Bertoldi, 1977). In the Upper Valdarno, a 
marked decrease in Taxodium type pollen, after an 
acme phase, was also observed close to 1.8 Ma 
(Napoleone et al., 2003). However, further evidence 
demonstrates: 1. recurrent sudden falls in the pollen 
percentages of “Taxodiaceae” throughout the Gelasian; 
and 2. the persistence of “Taxodiaceae” after 1.8 Ma in 
many Italian sites (e.g. Becker-Platen et al., 1977; Ber-
tolani Marchetti et al., 1979; Combourieu-Nebout, 1993, 
1995; Ravazzi & Rossignol-Strick, 1995; Fusco, 1996, 
2007, 2010; Bertini, 2001). The “Taxodiaceae” strati-
graphical range demonstrates that their disappearance 
cannot be considered as a synchronous and sudden 
event matching exactly the former Plio-Pleistocene 
boundary at 1.8 Ma. It is therefore impossible to use the 
Tiberian boundary to cross-correlate Italian succes-
sions. The establishment of wide-scale correlations is 
even more difficult, on this basis: see, for instance, the 
correlation of the Tiberian boundary with the Reuverian/
Pretiglian transition of The Netherlands (Zagwijn, 1975) 
proposed by Lona (1971). The Reuverian/Pretiglian 
transition corresponds on the basis of chronostrati-
graphical evidence, to the Piacenzian/Gelasian bounda-
ry (at ca. 2.6 Ma) and not to the Gelasian/Calabrian 
transition (at 1.8 Ma). North-western European conti-
nental “stages” (e.g. Zagwijn, 1957, 1959, 1960, 1963; 
de Jong, 1988) summarized in Zagwijn & Hager (1987) 
and Zagwijn (1992) have represented a significant 
standard in continental Neogene stratigraphy. However, 
their validity, and their use by many authors, e.g. for 
long-distance chronostratigraphic correlations or to es-
tablish chronostratigraphical frameworks, have been 
criticized conceptually and methodologically (e.g. Ber-
tini, 2003; Leroy, 2007). Substantial revisions were car-
ried out by Drees (2005), Donders et al. (2007), and 
Kemna & Westerhoff (2007). All previous reflections do 
not reduce the stratigraphical potential of palynology for 
the study of the Late Neogene and Quaternary. On the 
contrary, palynology, when correctly applied, is a very 
powerful tool for stratigraphic and palaeoenvironmental 
reconstructions, all the more so because, in contrast to 
other fossils, palynomorphs, being contained in both 
marine and non-marine sediments, permit direct corre-
lation of onshore and offshore successions (e.g. Com-
bourieu-Nebout & Vergnaud-Grazzini, 1991; Capraro et 
al., 2005; Joannin et al., 2007; Joannin et al., 2008; 
Bertini et al., 2010). 

 
 

5.4. Palaeoclimate change, palaeonvironments and 
human migration  

The great abundance of pollen records allows for a 
coherent spatial and temporal framework of the palaeo-
environmental scenarios during successive Pleistocene 
G/I cycles. This information is also fundamental to un-
derstanding the role of climate in the migration and sub-
sequent colonization of the genus Homo (e.g. Dennell 
et al., 2011; Manzi et al., 2011; Messager et al., 2011 
and references therein). Chronologically well-
constrained pollen records have been included in data-
bases and have often contributed to climate quantifica-
tion too, contributing to the compilation of regional to 
continental-scale palaeogeographic maps and the iden-
tification of the most likely migration routes for hominin 
expansion and settlement (e.g. Muttoni et al., 2010, 
2018; Moncel et al., 2018; Sànchez Goñi, 2022 and 
references therein). The beginning of the EMPT which 
follows several migration attempts can be taken to rep-
resent the first undisputed and well-dated phases of 
human colonization in Europe (e.g. Head & Gibbard, 
2005; Head et al., 2008; Muttoni et al., 2010, 2018; 
O'Regan et al., 2011; Abbate and Sagri, 2012). It in-
cludes the last obliquity-forced climate cycles, when 
hominins spread from Africa to the Eurasian continent 
(e.g. Dennell et al., 2011 and references therein). Ac-
cording to some authors (e.g. Leroy et al., 2011), the 
Early Pleistocene interval could provide the latest best 
narrow windows of opportunity for hominins to disperse 
into Europe during the transition from glacial to intergla-
cial periods, the full glacials being too cold for them and 
the interglacial to glacial transition too forested, alt-
hough this has not yet been demonstrated.  

What is more evident is the increasing imprint of 
Human populations after the Late Pleistocene and up to 
the present-day. The opportunity to better understand 
the behaviour of the natural vegetation and its adapta-
tion to climate changes in earlier periods may help to 
develop strategies for sustainable conservation of vege-
tation in the future. 
 
6. CONCLUSIONS  
 

The overall pollen record allows us to trace the 
main vegetational features linked to the complex mosaic 
of different physiographic and climatic conditions of the 
Italian peninsula between ca 3.6 Ma and 0.117 ka i.e. 
preceding the main Holocene human impact on the 
landscape.  

Pollen evidence is chronologically well constrained 
because it is found on sites that are well calibrated 
through the use of time-diagnostic tools e.g. magneto-
stratigraphy, micropalaeontology, biochronology, radio-
metric dating methods (U/Th, 14 C, …) and varve chro-
nology. This approach allows palynologists to address 
the main scientific stratigraphic and biological issues for 
the Quaternary. Particularly relevant is the documenta-
tion of the flora and vegetation response to both Milan-
kovitch cycles and sub-millennial events. Moreover, this 
set of evidence provides indispensable information for a 
better comprehension of topics which become crucial 
during the Holocene, i.e human versus natural climate 

 
 

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factors, location and extent of the glacial refugia for tem-
perate and Mediterranean species, modern floristic rich-
ness and genetic diversity.  

This summary illustrates the behaviour of Italian 
vegetation in the face of natural climatic changes 
through ancient times and in the absence of any human 
influence; the aim is to contribute to predictions regard-
ing the future of Mediterranean vegetation under ongo-
ing climate changes. 
 
ACKNOWLEDGEMENTS 

Authors thank the University of Firenze and the 
CNRS (Centre National de la Recherche Scientifique) 
for providing financial support for their researches and 
the Erasmus program for the possibilities of exchanges 
between the two laboratories. 

 
REFERENCES 
 
Abbate E., Sagri M. (2012) - Early to Middle Pleistocene 

Homo dispersals from Africa to Eurasia: Geologi-
cal, climatic and environmental constraints. Qua-
ternary International, 267, 3-19. 

Albianelli A. Bertini A., Magi M., Napoleone G., Sagri M. 
(1995) - II bacino Plio- Pleistocenico del Valdarno 
Superiore: eventi deposizionali, paleomagnetici e 
paleoclimatici. II Quaternario (Ital. J. Quat. Sci.), 8 
(1), 11-18. 

Allen J.R.M., Huntley B. (2000) - Weichselian palynolog-
ical records from southern Europe: correlation and 
chronology. Quaternary International, 73 (74), 111-
125. 

Allen J.R.M., Huntley B. (2009) - Last interglacial palae-
ovegetation, palaeoenvironments and chronology: 
a new record from Lago Grande di Monticchio, 
southern Italy. Quaternary Science Review, 28, 
1521-1538. 

Amato V., Aucelli P.P.C., Cesarano M., Pappone G., 
Rosskopf C.M., Russo Ermolli E. (2011) - The 
Sessano intra-montane basin: new multi-proxy 
data for the Quaternary evolution of the Molise 
sector of the Central-Southern Apennines (Italy). 
Geomorphology, 128, 15-31. 

Amorosi A., Colalongo M.L., Fiorini F., Fusco F., Pasini 
G., Vaiani S.C., Sarti G. (2004) - Palaeogeograph-
ic and palaeoclimatic evolution of the Po Plain 
from 150- ky core records. Global and Planetary 
Change, 40 (1-2), 55-78. 

Arias C., Bigazzi G., Bonadonna F.P., Iaccarino S., 
Urban B., Dal Molin M., Dal Monte L., Martolini M. 
(1990) - Valle Ricca Late Neogene stratigraphy 
(Lazio region, Central Italy). Paléobiologie Conti-
nentale, 17, 61-68. 

Aucelli P., Amato V., Cesarano M., Pappone G., 
Rosskopf C.M., Russo Ermolli E., Scarciglia F. 
(2011) - New morphostratigraphic and chronologi-
cal constraints for the Quaternary palaeosurfaces 
of the Molise Apennines (southern Italy). Geologi-
ca Carpathica, 62, 17-26. 

Badino F., Pini R., Bertuletti P., Ravazzi, C., Delmonte 
B., Monegato G., Reimer P., Vallé F., Arrighi S., 
Bortolini E., Figus C., Lugli F., Maggi V., Marciani 

G., Margaritora D., Oxilia G., Silvestrini S., Benaz-
zi S. (2020a) - The fast-acting “pulse” of Heinrich 
Stadial 3 in a mid-latitude boreal ecosystem. 
Scientific Reports, 10(1), 18031. 

Badino F., Pini R., Ravazzi C., Margaritora D., Arrighi 
S., Bortolini E., Figus S., Giaccio B., Lugli F., 
Marciani G., Monegato G., Moroni A., Negrino F., 
Oxili G., Peresani M., Romandini M., Ronchitelli 
A., Spinapolice E., Zerboni A., Benazzi S. (2020b) 
- An overview of Alpine and Mediterranean pa-
laeogeography, terrestrial ecosystems and climate 
history during MIS 3 with focus on the Middle to 
Upper Palaeolithic transition. Quaternary Interna-
tional, 551, 7-28.  

Baggioni M., Suc J.-S., Vernet J.L. (1981) - Le Plio-
Pléistocène de Camerota (Italie méridionale): 
géomorphologie et paléoflores. Geobios 14, 229-
237. 

Bartoli G., Hönisch B., Zeebe R.E. (2011) - Atmospheric 
CO2 decline during the Pliocene intensification of 
Northern Hemisphere glaciations. Paleoceanogra-
phy, 26(4). 

Basilici G. (1997) - Sedimentary facies in an extensional 
and deep lacustrine depositional system: the Plio-
cene Tiberino Basin, Central Italy. Sedimentology, 
109, 73-94. 

Becker-Platen J.D., Benda L., Cepek P., Daniels 
C.H.V., Mengeling H., Meyer K.J., Steffens P., 
Streif H., Vinken R. (1977) - Beiträge zur Plio-
Pleistozän-Grenzziehung in Mittelitalien. Giornale 
di Geologia, 41, 107-114. 

Belluomini G., Manfra L., Palombo M.R. (1997) - A re-
cent aminochronology of Pleistocene Mammals 
from Rome area and Isernia-La Pineta deposits 
(Italy). Geologica Romana, 33, 73-79. 

Benvenuti M., Degli Innocenti D. (2001) - The Pliocene 
deposits in the central-eastern Valdelsa Basin 
(Florence, Italy) revised through facies analysis 
and unconformity-bounded stratigraphic units. 
Rivista Italiana di Paleontologia e Stratigrafia, 107, 
265-286. 

Benvenuti M., Bertini A., Conti C., Dominici S. (2007) - 
Integrated analyses of litho and bio-facies in a 
Pliocene cyclothemic, alluvial to shallow marine 
succession (Tuscany, Italy). Geobios, 40 (2), 143-
158. 

Berger A.M., Loutre M.F. (1991) - Insolation values for 
the climate of the last 10 million of years. Quater-
nary Science Review, 10 (4), 297-317.  

 Doi: 10.1016/0277- 3791(91)90033-Q 
Bertini A. (1994) - Messinian-Zanclean vegetation and 

climate in north-central Italy. Historical Biology, 9, 
3-10. 

Bertini A. (2000) - Pollen record from Colle Curti and 
Cesi: early and middle Pleistocene mammal sites 
in the Umbro-Marchean Apennine mountains 
(central Italy). Journal of Quaternary Sciences, 15, 
825-840. 

Bertini A. (2001) - Pliocene climatic cycles and altitudi-
nal forest development from 2.7 Ma in the north-
ern Apennines (Italy): evidence from the pollen 
record of the Stirone section (~5.1 to ca. 2.2 Ma). 

 
 

108 

 
 

Bertini A., Combourien-Nebout N. 



   

 

Geobios 34, 253-265. 
Bertini A. (2002) - Palynological evidence of late Neo-

gene environments in Italy. Acta Universitatis Car-
olinae, Geologica, 46, 15-25. 

Bertini A. (2003) - Early to Middle Pleistocene changes 
of the Italian flora and vegetation in the light of a 
chronostratigraphic framework. II Quaternario (Ital. 
J. Quat. Sci.) 16, 19-36. 

Bertini A. (2010) - Pliocene to Pleistocene palynoflora 
and vegetation in Italy: state of the art. Quaternary 
International, 225, 5-24. 

Bertini A. (2013) - Climate and vegetation in the Upper 
Valdarno Basin (central Italy) as a response to 
Northern Hemisphere insolation forcing and re-
gional tectonics in the late Pliocene-early Pleisto-
cene. Italian Journal of Geosciences,132, 137-
148. 

Bertini A., Roiron P. (1997) - Evolution de la végétation 
et du climat pendant le Pliocène moyen en Italie 
centrale: apport de la palynologie et des macro-
flores à l'étude du bassin du Valdarno Supérieur 
(coupe de Santa Barbara). Comptes Rendus de 
l'Académie des Sciences Paris, 324, 763-771. 

Bertini A., Martinetto E. (2008) - Messinian to Zanclean 
vegetation and climate of Northern and Central 
Italy. Bollettino della Società Paleontologica Ital-
iana, 47, 105-121. 

Bertini A., Martinetto E. (2011) - Reconstruction of veg-
etation transects for the Messinian-Piacenzian of 
Italy by means of comparative analysis of pollen, 
leaf and carpological records. Palaeogeography 
Palaeoclimatology Palaeoecology, 304, 230-246. 

Bertini A., Londeix L., Maniscalco R., di Stefano A., Suc 
J.P., Clauzon G., Gautier F., Grasso M. (1998) - 
Paleobiological evidence of depositional condi-
tions in the Salt Member, Gessoso-Solfifera for-
mation (Messinian, upper Miocene) of Sicily. Mi-
cropaleontology 44 (4), 413-433. 

Bertini A., Ciaranfi N., Marino M., Palombo M.R. (2010) 
- Proposal for Pliocene and Pleistocene land-sea 
correlation in the Italian area. Quaternary Interna-
tional, 219, 95-108. 

Bertini A., Magri D., Martinetto E., Sadori L., Vassio E. 
(2014) - La flora del Pleistocene dell’Italia centra-
le. The Pleistocene flora of central Italy. In: Kusta-
tscher E., Roghi G., Bertini A., Miola A. (eds.): La 
storia delle piante fossili in Italia. Paleobotany of 
Italy. Pubblicato in occasione del XI Congresso 
Europeo di Paleobotanica, Padova, 26-31 Agosto 
2014. Pubblicazione del Museo di Scienze Natu-
rali dell’Alto Adige n. 9, 308-328. ISBN 
9788887108064 

Bertini A., Toti F., Marino M., Ciaranfi N. (2015) - Vege-
tation and climate across the Early-Middle Pleisto-
cene transition at Montalbano Jonico, southern 
Italy. Quaternary International, 383, 74-88.  

Bertini A., Piccini L., Ricci M., Massini M. (2022) - Paly-
nological evidence of Middle Pleistocene palaeo-
environmental changes from the ‘Buca dell’Onice’ 
flowstone (Alpi Apuane, Central Italy). The Depo-
sitional Record, 8(1), 340-354. 

Bertolani-Marchetti D., Accorsi C.A., Pelosio G., Raffi S. 

(1979) - Palynology and stratigraphy of the Plio-
Pleistocene sequence of the Stirone River 
(northern Italy). Pollen et Spores, 21, 149-167. 

Bertoldi R. (1977) - Studio palinologico della serie di Le 
Castella (Calabria). Atti Accademia Nazionale dei 
Lincei, 62 (4), 535-547. 

Bertoldi R. (1985a) - Palinostratigrafia in sedimenti ma-
rini del Pliocene della Sicilia e considerazioni pa-
leoclimatiche. Atti Accademia Nazionale dei Lin-
cei, 78, 157-164. 

Bertoldi R. (1985b) - Testimonianze palinologiche di età 
“Villafranchiana” nel deposito fluvio-lacustre di 
Compiano (Alta Val di Taro). Acta Naturalia dell’A-
teneo Parmense, 21, 23-30. 

Bertoldi R. (1988) - Una sequenza palinologica di età 
Rusciniana nei sedimenti lacustri basali del bacino 
di Aulla-Olivola. Rivista Italiana di Paleontologia e 
Stratigrafia, 94, 105-138. 

Bertoldi R., Castello F. (1990) - Palinoflora di età Villa-
franchiana inferiore nel deposito continentale di 
Sesta Godano in Val di Vara (Bacino del F. Ma-
gra). Atti della Società Toscana di Scienze Natu-
rali, 97, 31-45. 

Bertoldi R., Rio D., Thunell R. (1989) - Pliocene-
Pleistocene vegetational and climatic evolution of 
the south-central Mediterranean. Palaeogeogra-
phy Palaeoclimatology Palaeoecology, 72, 263-
275. 

Bertoldi R., Binotti A., Castello F. (1994) - Reevesia and 
Itea in the pollen flora of the Upper Neogene con-
tinental deposit at Sarzana (lower Magra valley, 
Northern Italy). Review Palaeobotany and Paly-
nology, 80, 159-172. 

Blasi C. (2010) - La vegetazione d'Italia con carta delle 
Serie di Vegetazione in scala 1:500.000. Palombi 
Editore, Rome, Italy, pp. 536. 

Bonin G. (1981) - L'étagement de la végétation dans 
l'Appenin méridional. Ecologie Mediterranéenne 7, 
79-91. 

Borzi M., Carboni M.G., Di Bella L., Florindo F., Girotti 
O., Piccardi E., Sagnotti L. (1998) - Bio- and ma-
gneto-stratigraphy in the Tiber valley revised. 
Quaternary International 47 (48), 65-72. 

Bossio A., Costantini A., Lazzaretto A., Lotta D., 
Mazzanti R., Mazzei R., Salvatorini G., Sandrelli 
F. (1993) - Rassegna delle conoscenze sulla stra-
tigrafia del Neoautoctono toscano. Memorie della 
Società Geologica Italiana, 49, 17-98. 

Brauer A., Mingram J., Frank U., Gunter C., Schettler 
G., Wulf S., Zolitschka B., Negendank J.F.W. 
(2000) - Abrupt environmental oscillations during 
the early Weichselian recorded at Lago Grande di 
Monticchio, southern Italy. Quaternary Internation-
al, 73 (74), 79-90. 

Brauer A., Allen J.R.M., Mingram J., Dulski P., Wulf S., 
Huntley B. (2007a) - Evidence for last interglacial 
chronology and environmental change from 
Southern Europe. Proceedings of the National 
Academy of Sciences, 104, 450-455. 

Brauer A., Wulf S., Mangili C., Moscariello A. (2007b) - 
Tephrochronological dating of varved interglacial 
lake deposits from Piànico-Sèllere (Southern Alps, 

 
 

109 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

Italy) to around 400 ka. Journal of Quaternary 
Sciences, 22, 85-96. 

Brauer A., Mangili C., Moscariello A., Witt A. (2008) - 
Palaeoclimatic implications from micro-facies data 
of a 5900 varves time series from the Piànico 
interglacial sediment record, Southern Alps. Pa-
laeogeography Palaeoclimatology Palaeoecology, 
259, 121-135. 

Brenac P. (1984) - Végétation et climat de la Campanie 
du Sud (Italie) au Pliocène final d'après l'analyse 
pollinique des dépots de Camerota. Ecologie 
Mediterranéenne, 10, 207-216. 

Brolsma M.J. (1978) - Quantitative foraminiferal analy-
sis and environmental interpretation of the Plio-
cene and topmost Miocene on the South coast of 
Sicily. Utrecht Micropaleontological Bullettins, 18, 
159. 

Brunsfeld S.J., Soltis P.E., Soltis D.E., Gedek P.A., 
Quinn C.J., Strenge D.D., Ranker T.A. (1994) -
Phylogenetic relationships among the genera of 
Taxodiaceae and Cupressaceae: evidence from 
rbcL sequences. Systematic Botany, 19 (2), 253-
262. 

Buccheri G., Capretto G., Di Donato V., Esposito P., 
Ferruzza G., Pescatore T., Russo Ermolli E., 
Senatore M.R., Sprovieri M., Bertoldo M., Carella 
D., Madonia G. (2002) - A high resolution record 
of the last deglaciation in the southern Tyrrhenian 
Sea: environmental and climatic evolution. Marine 
Geology, 115, 1-24. 

Canali G., Capraro L., Donnici S., Rizzetto F., Serandrei
-Barbero R., Tosi L. (2007) - Vegetational and 
environmental changes in the eastern Venetian 
coastal plain (Northern Italy) over the past 80,000 
years. Palaeogeography Palaeoclimatolology 
Palaeoecology, 253, 300-316. 

Capraro L., Asioli A., Backman J., Bertoldi R., Channell 
J.E.T., Massari F., Rio D. (2005) - Climatic pat-
terns revealed by pollen and oxygen isotope rec-
ords across the Matuyama Brunhes Boundary in 
the central Mediterranean (southern Italy). In: 
Head M.D., Gibbard P.L. (Eds.), Early-Middle 
Pleistocene Transition: the Land-Ocean Evidence. 
Geological Society of London Spec. Publ. 247, 
159-182. 

Capraro L., Macrì P., Scarponi D., Rio D. (2015) - The 
lower to Middle Pleistocene Valle di Manche sec-
tion (Calabria, Southern Italy): state of the art and 
current advances. Quaternary International, 383, 
36-46. 

Capraro L., Ferretti P., Macrì P., Scarponi, D., Tateo F., 
Fornaciari E., Bellini G., Dalan G. (2017) - The 
Valle di Manche section (Calabria, Southern Italy): 
a high resolution record of the Early-Middle Pleis-
tocene transition (MIS 21-MIS 19) in the Central 
Mediterranean. Quaternary Science Reviews, 
165, 31-48.  

Capraro L., Bonomo S., Di Stefano A., Ferretti P., 
Fornaciari E., Galeotti S., Incarbona A., Macrì P., 
Raffi I., Sabatino N., Speranza F., Sprovieri M., Di 
Stefano E., Sprovieri R., Rio D. (2022) - The Mon-
te San Nicola section (Sicily) revisited: A potential 
unit-stratotype of the Gelasian Stage. Quaternary 

Science Reviews, 278, 107367. 
Carminati E., Doglioni C. (2012) - Alps vs. Apennines: 

The paradigm of a tectonically asymmetric Earth. 
Earth-Science Reviews, 112(1-2), 67-96.  

Carraro F., Ambrosetti P., Angela-Franchini M., Azzaroli 
A., Basilici G., Belluomini G., Bertoldi R., Boano 
P., Bormioli D., Campanino F., Caramiello R., 
Cerasoli M., Ciangherotti A., Compagnoni R., Esu 
D., Ferrero E., Ficcarelli G., Forno M.G., 
Genovese G., Giardino M., Girotti O., Gliozzi E., 
Lanza R., Mai D., Manfra L., Martinetto E., Masini 
F., Moscariello A., Pavia G., Perotto A., Polino R., 
Sala B., Siniscalco C., Torre D., Violanti D. (1996) 
- Revisione del Villafranchiano nell'area tipo di 
Villafranca d'Asti. Il Quaternario (Ital. J. Geosci.), 
9, 5-120. 

Caruso A. (2004) - Climatic changes during late Plio-
cene and early Pleistocene at Capo Rossello 
(Sicily, Italy): response from planktonic foraminif-
era. In: Coccioni R., Galeotti S., Lirer F. (Eds.), 
Proceedings of the First Italian Meeting on Envi-
ronmental Micropaleontology. Grzybowski Foun-
dation Special Publication 9, pp. 17-36. 

Castradori D., Rio D., Hilgen F.J., Lourens L.J. (1998) - 
The global standard stratotype section and point 
(GSSP) of the Piacenzian Stage (Middle Plio-
cene). Episodes, 21, 88-93. 

Chiarini E., Giardini M., La Posta E., Papasodaro F., 
Sadori L. (2007) - Sedimentology, palynology and 
new geochronological constraints on Quaternary 
deposits of the Corvaro intermontane basin 
(Central Italy). Review of Micropaleontology, 50, 
309-314. 

Chiarini E., Giardini M., Mattei M., Papasodaro F., 
Porreca M., Sadori L. (2009) - Plio-Quaternary 
evolution of the high Salto river valley: the Marano 
de' Marsi unit. Il Quaternario (Ital. J. Quat. Sci.) 22 
(2), 325-344. 

Cita M.B., Gartner S. (1973) - Studi sul Pliocene e sugli 
strati di passaggio dal Miocene al Pliocene. V. 
The stratotype Zanclean. Foraminiferal and nan-
nofossil biostratigraphy. Rivista Italiana di Paleon-
tologia e Stratigrafia, 79, 503-558. 

Cita M.B., Capraro L., Ciaranfi N., Di Stefano E., Lirer 
F., Maiorano P., Marino M., Raffi I., Rio D., 
Sprovieri R., Stefanelli S., Vai G.B. (2008) - The 
Calabrian Stage redefined. Episodes, 31, 408-
419.  

Clauzon G., Suc J. P., Gautier F., Berger A., Loutre M. 
F. (1996) - Alternate interpretation of the Messini-
an salinity crisis: controversy resolved? Geology, 
24(4), 363-366. 

Cohen K.M., Finney S.C., Gibbard P.L., Fan, J.X. 
(2013; updated) - The ICS International Chronos-
tratigraphic Chart. Episodes, 36, 199-204. (Online 
version 2022/10). 

Cohen K., Gibbard P. (2022) - Global chronostratigraph-
ical correlation table for the last 2.7 million years 
v. 2019 (Poster version). Mendeley Data, V5. 

 Doi: 10.17632/dtsn3xn3n6.5 
Coltorti M., Albianelli A., Bertini A., Ficcarelli G., 

Laurenzi M.A., Napoleon G., Torre D. (1998) - The 
Colle Curti Mammal site in the Colfiorito area 

 
 

110 

 
 

Bertini A., Combourien-Nebout N. 



   

 

(Umbria-Marchean Apennine, Italy): geomorpholo-
gy, stratigraphy, paleomagnetism and palynology. 
Quaternary International, 46 (48), 107-116. 

Combourieu-Nebout N. (1993) - Vegetation response to 
upper Pliocene glacial/interglacial cyclicity in the 
central Mediterranean. Quaternary Research, 40, 
228-236. 

Combourieu-Nebout N. (1995) - Réponse de la végéta-
tion de l'Italie méridionale au seuil climatique de la 
fin du Pliocène d'après l'analyse pollinique haute 
résolution de la section de Semaforo (2,46 à 2.1 
Ma). Comptes Rendus de l'Académie des 
Sciences Paris, 321, 659-665. 

Combourieu-Nebout N., Vergnaud-Grazzini C. (1991) - 
Late Pliocene northern hemisphere glaciations: 
the continental and marine responses in the cen-
tral Mediterranean. Quaternary Sciences Review, 
10, 319-334. 

Combourieu-Nebout N., Semah F., Djubiantono T. 
(1990) - La limite Pliocène/Pléistocène: précisions 
magnétostratigraphiques par l'étude sériée de la 
coupe type de Vrica (Crotone, Italie). Comptes 
Rendus de l'Académie des Sciences Paris, 311, 
851-857. 

Combourieu-Nebout N., Siani G., Paterne M., Turon 
J.L. (1998) - A high resolution record of the Last 
Deglaciation in the Central Mediterranean Sea: 
palaeovegetation and palaeohydrological evolu-
tion. Quaternary Sciences Review, 17, 3-4. 

Combourieu-Nebout N., Fauquette S., Quézel P. (2000) 
- What was the Late Pliocene Mediterranean cli-
mate like: a preliminary quantification from vegeta-
tion. Bulletin de la Société Géologique de France, 
171, 271-277. 

Combourieu-Nebout, N., Turon J.L., Zhan R., Capotondi 
L., Londeix L., Pahnke K. (2002) - Enhanced aridi-
ty and atmospheric high-pressure stability over the 
western Mediterranean during the North Atlantic 
cold event of the past 50 ky. Geology, 30, 863-
866. 

Combourieu-Nebout N., Foucault A., Mélières F. (2004) 
- Vegetation markers of palaeoclimate cyclical 
changes in the Pliocene of Punta Piccola (Sicily, 
Italy). Palaeogeography Palaeoclimatolology Pal-
aeoecology, 214 (1-2), 55-66. 

Combourieu-Nebout N., Peyron O., Bout-Roumazeille 
V., Goring S., Dormoy I., Joannin S., Sadori L., 
Siani G., Magny M. (2013) - Holocene vegetation 
and climate changes in central Mediterranean 
inferred from a high-resolution marine pollen rec-
ord (Adriatic Sea). Climate of the Past 9, 2023-
2042. 

Combourieu-Nebout N., Bertini A., Russo Ermolli E.,  
Peyron O., Klotz S., Montade V., Fauquette S., 
Allen J.R.M., Fusco F., Goring S. Huntley B.,  
Joannin S., Lebreton V.,  Magri D.,  Martinetto E.,  
Orain R., Sadori, L. (2015) - Climate changes in 
the central Mediterranean and Italian vegetation 
dynamics since the Pliocene. Review of Palaeo-
botany and Palynology, 218, 127-147. 

Corrado P., Magri D. (2011) - A late Early Pleistocene 
pollen record from Fontana Ranuccio (central 
Italy). II Quaternario (J. Quat. Sci.) 26, 335-344. 

Cosentino D., Asti R., Nocentin M., Gliozzi E., Kotsakis 
T., Mattei M., Esu D., Spadi M., Tallini M., Cifelli 
F., Pennacchioni M., Cavuoto G., Di Fiore, V. 
(2017) - New insights into the onset and evolution 
of the central Apennine extensional intermontane 
basins based on the tectonically active L’Aquila 
Basin (central Italy). GSA Bulletin, 129(9-10), 
1314-1336. 

Cremaschi M. (1983) - La serie pleistocenica di Isernia 
la Pineta (Molise) e la posizione stratigrafica dei 
suoli d'abitato paleolitici in essa inclusi. In: Peretto 
C., Terzan, C., Cremaschi M. (Eds.), Isernia La 
Pineta, un accampamento più antico di 700 000 
anni. Calderini, Bologna, 49-62. 

Cremaschi M., Peretto C. (1988) - Les sols d'habitat du 
site paléo-lithique d'Isernia la Pineta (Molise, Italie 
centrale). L'Anthropologie 92, 1017-1040. 

Cremaschi M., Ravazzi C. (1995) - Nuovi dati stratigrafi-
ci e geocronologici sul Bacino di Leffe (Prealpi 
Lombarde, Bergamo). II Quaternario (J. Quat. 
Sci.), 8(1), 167-182. 

De Beaulieu J.L., Andrieu-Ponel V., Cheddadi R., Guiter 
F., Ravazzi C., Reille M., Rossi S. (2006) - Apport 
des longues séquences lacustres à la connais-
sance des variations des climats et des paysages 
pléistocènes. Comptes Rendus Palevol, 5, 65-72. 

De Jong J. (1988) - Climatic variability during the past 
three million years, as indicated by vegetational 
evolution in northwest Europe and with emphasis 
on data from The Netherlands. Philosophical 
Transactions of the Royal Society of London B 
318, 603-617. 

De La Vega E., Chalk T.B., Wilson P.A., Bysani R.P., 
Foster, G. L. (2020) - Atmospheric CO2 during the 
Mid-Piacenzian Warm Period and the M2 glacia-
tion. Scientific Reports, 10(1), 1-8. 

De la Vega E., Chalk T.B., Wilson P.A., Bysani R.P., 
Foster G. L. (2021) - Author Correction: Atmos-
pheric CO2 during the Mid-Piacenzian Warm Peri-
od and the M2 glaciation. Scientific Reports, 11
(1), 1-4. 

De Schepper S., Groeneveld J., Naafs, B.D.A., Van 
Renterghem C., Hennissen J., Head M.J., Louwye 
S., Fabian K. (2013) - Northern hemisphere glaci-
ation during the globally warm early late Pliocene. 
PloS one, 8(12), e81508. 

Delitala M.C., Fornaseri M., Nicoletti M. (1983) - Data-
zioni argon-potassio sulla serie pleistocenica di 
Isernia La Pineta. In: Peretto, C., Terzani, C., Cre-
maschi, M. (Eds.), Isernia La Pineta, un accapa-
mento più antico di 700.000 anni. Calderini, Bolo-
gna, 65-66. 

Dennell R.W., Martinon Torres M., Bermudez de Castro 
J.M. (2011) - Hominin variability, climatic instability 
and population demography in Middle Pleistocene 
Europe. Quaternary Science Reviews, 30, 1511-
1524. 

Desprat S., Combourieu-Nebout N., Essallami L., Sicre 
M.A., Dormoy I., Peyron O., Siani G., Bout 
Roumazeilles V., Turon J.L. (2013) - Deglacial 
and Holocene vegetation and climatic changes at 
the southernmost tip of the Central Mediterranean 
from a direct land-sea correlation. Climate of the 

 
 

111 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

Past, 9, 767-787. 
Di Bella L., Carbonia M.G., Bergamin L., Iamund F. 

(2005) - The Early Pleistocene in Latium (Central 
Italy): palaeoecology from benthic foraminiferal 
record. Quaternary International, 131, 23-34. 

Di Donato V., Esposito P., Russo Ermolli E., Scarano 
A., Cheddadi R. (2008) - Coupled atmospheric 
and marine palaeoclimatic reconstruction for the 
last 35 ka in the Sele Plain-Gulf of Salerno area 
(southern Italy). Quaternary International, 190, 
146-157. 

Di Rita F., Sottili G. (2019) - Pollen analysis and tephro-
chronology of a MIS 13 lacustrine succession from 
Eastern Sabatini Volcanic District (Rignano 
Flaminio, central Italy). Quaternary Science Re-
views, 204, 78-93.  

Di Rita F., Anzidei A.P., Magri D. (2013) - A Late glacial 
and early Holocene pollen record from Valle di 
Castiglione (Rome): vegetation dynamics and 
climate implications. Quaternary International, 
288, 73-80. 

Donders T.H., Kloosterboer-van Hoeve M.L., 
Westerhoff W., Verreussel R.M.C.H., Lotter A.F. 
(2007) - Late Neogene continental stages in NW 
Europe revisited. Earth-Science Reviews, 85, 161-
186. 

Drees M. (2005) - An evaluation of the early Pleistocene 
chronology of the Netherlands. Vertebrate Palae-
ontology, 1, 1-46. 

 www.palArch.nl 
Dowsett H.J., Robinson M.M., Haywood A.M., Hill D.J., 

Dolan A.M., Stoll D.K., Le Chan W., Abe-Ouchi A., 
Chandler M.A., Rosenbloom N.A., Otto-Bliesner 
B.L., Bragg JF.G., Lunt D.J, Foley K.M., 
Riesselman, C.R. (2012) - Assessing confidence 
in Pliocene sea surface temperatures to evaluate 
predictive models. Nature Climate Change, 2(5), 
365-371.  

Dubois J.M. (2001) - Cycles climatiques et paramètres 
orbitaux vers 1 Ma. Etude de la Coupe de Monte 
San Giorgio (Caltagirone, Sicile): Palynologie, 
Isotope Stables, Calcimétrie. Mémoire DEA. Lyon 
University, Lyon, France. 

Farjon A. (1998) - World Checklist and Bibliography of 
Conifers. Royal Botanical Gardens, Kew, Rich-
mond, U.K. 

Farjon A. (2005) - A Monograph of Cupressaceae and 
Sciadopitys. Royal Botanic Gardens, Kew, UK 
1842460684. 

Fauquette S., Bertini A. (2003) - Quantification of north-
ern Italy Pliocene climate from pollen data: evi-
dences for a very peculiar climate pattern. Boreas, 
32, 361-369. 

Fauquette S., Combourieu-Nebout N. (2013) - Palaeo-
altitude of the Sila Massif (Southern Apennines, 
Italy) and distribution of the vegetation belts at ca. 
2.4 Ma (Early Pleistocene). Review of Palaeobota-
ny and Palynology, 189, 1-7. 

Favaretto S., Asioli A., Miola A., Piva A. (2008) - Pre-
boreal climatic oscillations recorded by pollen and 
foraminifera in the Southern Adriatic Sea. Quater-
nary International, 190, 89-102. 

Federici P.R. (1973) - La tettonica recente dell'Appenni-

no 1. Il bacino Villafranchiano di Sarzana e il suo 
significato nel quadro dei movimenti distensivi a 
nord-ovest delle Alpi Apuane. Bollettino della So-
cietà Geologica Italiana, 92, 287-301. 

Ficcarelli G., Abbazzi L., Albianelli A., Bertini A., Coltorti 
M., Magnatti M., Masini F., Mazza P., Mezzabotta 
C., Napoleone G., Rook L., Rustioni M., Torre D. 
(1997) - Cesi an early Middle Pleistocene site in 
the Colfiorito basin (Umbro-Marchean Apennine), 
central Italy. Journal of Quaternary Science, 12, 
507-518. 

Fidolini F., Ghinassi M., Magi M., Papini M., Sagri M. 
(2013) - The Plio-Pleistocene fluvio-lacustrine 
Upper Valdarno Basin (central Italy): stratigraphy 
and basin fill evolution. Italian Journal of Geosci-
ences, 132(1), 13-32. 

Follieri M. (1977) - Evidence on the Plio-Pleistocene 
palaeofloristic evolution in central Italy. Rivista 
Italiana di Paleontologia e Stratigrafia, 83, 925-
930. 

Follieri M., Magri D., Sadori L. (1988) - 250,000-year 
pollen record from Valle di Castiglione (Roma). 
Pollen et Spores 30, 329-356. 

Follieri M., Magri D., Sadori L. (1989) - Pollen strati-
graphical synthesis from Valle di Castiglione 
(Roma). Quaternary International, 3 (4), 81-84. 

Follieri M., Giardini M., Magri D., Sadori L. (1998) - Pa-
lynostratigraphy of the last glacial period in the 
volcanic region of central Italy. Quaternary Inter-
national 47/48, 3-20. 

Forno M. G., Gattiglio M., Comina C., Barbero D., 
Bertini A., Doglione A., Gianotti F., Irace A., Marti-
netto E., Mottura A., Sala B. (2015) - Stratigraphic 
and tectonic notes on the Villafranca d’Asti suc-
cession in type-area and Castelnuovo don Bosco 
sector (Asti Reliefs, Piedmont). Alpine and Medi-
terranean Quaternary (AMQ), 28 (1), 5-27. 

Francavilla F., Bertolani-Marchetti D., Tomadin L. 
(1970) - Ricerche stratigrafiche, sedimentologiche 
e palinologiche sul Villafranchiano tipo. Giornale di 
Geologia 36 (02), 701-741. 

Fusco F. (1996) - Analisi polliniche nelle successioni 
pleistoceniche inferiori della Valle del Lamone 
(Appennino nord-orientale). Evoluzione paleoveg-
etazionale e paleoclimatica. Unpublished PhD 
Thesis, University of Bologna, Italy), pp. 96. 

Fusco F. (2007) - Vegetation response to early Pleisto-
cene climatic cycles in the Lamone valley 
(Northern Apennines, Italy). Review of Palaeobot-
any and Palynology, 145, 1-23. 

Fusco F. (2010) - Picea + Tsuga pollen record as a 
mirror of oxygen isotope signal? An insight into 
the Italian long pollen series from Pliocene to Ear-
ly Pleistocene. Quaternary International, 225, 58-
74. 

Ge.Mi.Na. (Ed.) (1962) - Ligniti e Torbe dell'Italia Conti-
nentale. Società Geomineraria Nazionale, ILTE 
Ed. Roma, Italy. 

Ghinassi M., Magi M., Sagri M., Singer B.S. (2004) - 
Arid climate 2.5 Ma in the Plio-Pleistocene Valdar-
no Basin (Northern Apennines, Italy). Palaeogeo-
graphy Palaeoclimatolology Palaeoecology, 207, 
37-57. 

 
 

112 

 
 

Bertini A., Combourien-Nebout N. 



   

 

Giunta S., Negri A., Morigi C., Capotondi L., Combou-
rieu-Nebout N., Emeis K.C., Sangiorgi F., Vigliotti 
L. (2003) - Coccolithophorid ecostratigraphy and 
multi-proxy paleoceanographic reconstruction in 
the Southern Adriatic Sea during the last deglacial 
time (Core AD91-17). Palaeogeography Palaeocli-
matolology Palaeoecology, 190, 39-59. 

Grieser J., Gommes R., Bernardi M. (2006) - New 
LocClim-the local climate estimator of FAO. In 
Geophysical research abstracts, 8, No. 08305, p. 
2.  

Guerrera F., Coccioni D., Corradini D., Bertoldi R. 
(1984) - Caratteristiche lito-sedimentologiche e 
micropaleontologiche (Foraminiferi, Dinoflagellati, 
pollini e spore) di successioni “tripolacee” plioceni-
che del bacino di Caltanisetta. Bollettino della 
Società Geologica Italiana, 103, 629-660. 

Haywood A. M., Valdes P. J. (2004) - Modelling Middle 
Pliocene warmth: contribution of atmosphere, 
oceans and cryosphere. Earth Planetary Science 
Letters, 218, 363-377. 

Haywood A.M., Hill D.J., Dolan A.M., Otto-Bliesner B. 
L., Bragg F., Chan W.L., Chandler M.A., Contoux 
C., Dowsett H.J., Jost A., Kamae Y., Lohmann G., 
Lunt D.J., Abe-Ouchi A., Pickering S.J., Ramstein 
G., Rosenbloom N.A., Salzmann U., Sohl L., 
Stepanek C., Ueda H., Yan Q., Zhan Z. (2013) - 
Large-scale features of Pliocene climate: results 
from the Pliocene Model Intercomparison Project. 
Climate of the Past, 9(1), 191-209.  

Head M.J., Gibbard P.L. (2005) - Early-Middle Pleisto-
cene transitions: an overview and recommenda-
tion for the defining boundary. Geological Society, 
London, Special Publications, 247(1), 1-18. 

Head M.J., Gibbard P.J. (2015) - Early-Middle Pleisto-
cene transitions: linking terrestrial and marine 
realms. Quaternary International, 389, 7-46. 

Head M.J., Gibbard P., Salvador A. (2008) - The Qua-
ternary: its character and definition. Episodes 
Journal of International Geoscience, 31(2), 234-
238. 

Hilgen F.J. (1987) - Sedimentary rhythms and high-
resolution chronostratigraphic correlations in the 
Mediterranean Pliocene. Newsletters on Stratigra-
phy, 17, 109-127. 

Hilgen F.J., Langereis C.G. (1988) - The age of the 
Miocene-Pliocene boundary in the Capo Rossello 
area (Sicily). Earth and Planetary Science Letters, 
91, 214-222. 

Howell M.W., Thunell R., Tappa, E., Rio D., Sprovieri R. 
(1988) - Late Neogene laminated and opal-rich 
facies from the Mediterranean region: geochemi-
cal evidence for mechanisms of formation. Pal-
aeogeography Palaeoclimatology Palaeoecology, 
64, 265-286.  

Joannin S., Quillévéré F., Suc J.P., Lécuyer C., 
Martineau F. (2007) - Early Pleistocene climate 
changes in the central Mediterranean region as 
inferred from integrated pollen and foraminiferal 
stable isotope analyses. Quaternary Research, 
67, 264-274. 

Joannin S., Ciaranfi N., Stefanelli S. (2008) - Vegetation 
changes during the late early Pleistocene at Mon-

talbano Jonico (Province of Matera, southern Italy) 
based on pollen analysis. Palaeogeography Pal-
aeoclimatology Palaeoecology, 270, 92-101. 

Karner D. B., Juvigne E., Brancaccio L., Cinque A., 
Russo Ermolli E., Santangelo N., Bernasconi S., 
Lirer, L. (1999) - A potential early middle Pleisto-
cene tephrostratotype for the Mediterranean ba-
sin: the Vallo Di Diano, Campania, Italy. Global 
and Planetary change, 21(1-3), 1-15. 

Kemna H.A., Westerhoff W.E. (2007) - Remarks on the 
palynology-based chronostratigraphic subdivision 
of the Pliocene terrestrial deposits in NW-Europe. 
Quaternary International, 164/165, 184-196. 

Kent D.V., Rio D., Massari F., Kukla G., Lanci L. (2002) 
- Emergence of Venice during the Pleistocene. 
Quaternary Science Review, 21, 1719-1727. 

Klotz S., Fauquette S., Combourieu-Nebout N., Uhl D., 
Suc J.P., Mosbrugger V. (2006) - Seasonality 
intensification and long-term winter cooling as a 
part of the Late Pliocene climate development. 
Earth Planetary Science Letters, 241, 174-187. 

Lang N., Wolff E. W. (2011) - Interglacial and glacial 
variability from the last 800 ka in marine, ice and 
terrestrial archives. Climate of the Past, 7(2), 361-
380. 

Lebreton V. (2002) - Végétation et climat au Pléistocène 
inférieur et moyen à La Pineta (Isernia, Italie). 
Comptes Rendus Palevol, 1, 11-17. 

Leroy S. (2007) - Progress in palynology of the Gela-
sian-Calabrian stages in Europe: ten messages. 
Review Micropaleontology, 50, 293-308. 

Leroy S., Giralt S., Francus P., Seret G. (1996) - The 
high sensitivity of the palynological record in the 
Vico maar lacustrine sequence (Latium, Italy) 
highlights the climatic gradient through Europe for 
the last 90 ka. Quaternary Science Reviews, 15, 
189-201. 

Leroy S.A.G., Arpe K., Mikolajewicz U. (2011) - Vegeta-
tion context and climatic limits of the Early Pleisto-
cene hominin dispersal in Europe. Quaternary 
Science Reviews, 30, 1448-1463. 

Lisiecki L.E., Raymo M.E. (2005) - A Pliocene-
Pleistocene stack of 57 globally distributed benthic 
� 18O records. Paleoceanography, 20, PA1003. 

 Doi: 10.1029/2004PA001071 
Lisiecki L.E., Raymo M.E. (2007) - Plio-Pleistocene 

climate evolution: trends and transitions in glacial 
cycle dynamics. Quaternary Science Reviews, 26
(1-2), 56-69. 

Lona F. (1950) - Contributi alla storia della vegetazione 
e del clima nella Val Padana e analisi pollinica del 
giacimento villafranchiano di Leffe (Bergamo). Atti 
della Società Italiana di Scienze Naturali, 89, 123-
178. 

Lona F. (1962) - Prime analisi pollinologiche sui depositi 
terziari e quaternari di Castell'Arquato: reperti di 
vegetazione da clima freddo sotto le formazioni 
calcaree a Amphistegina. Bollettino Società Geo-
logica Italiana, 81, 1-5. 

Lona F. (1971) - Correlazioni tra alcune sequenze mi-
cropaleobotaniche plio-pleistoceniche continentali 
e marine dell’Italia centro-settentrionale ed Europa 
centro-occidentale con riferimento al Limite Tibe-

 
 

113 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

riano. L’Ateneo Parmense - Acta Naturalia, 6, 145
-157.  

Lona F., Bertoldi R. (1972) - La storia del Plio-
Pleistocene italiano in alcune sequenze vegeta-
zionali lacustri e marine. Atti Accademia Naziona-
le dei Lincei, 11, 1-45. 

Lona F., Follieri M. (1957) - Successione pollinica della 
serie superiore (Günz-Mindel) di Leffe (Bergamo). 
Veröffentlichungen des Geobotanischen Institutes 
der Eidg. Techn. Hochschule, Stiftung Rübel Zü-
rich, 34, 86-98. 

Lona F., Ricciardi E. (1961a) - Studio pollinologico stra-
tigrafico su una serie lacustre pleistocenica dell'I-
talia centrale (Bacino di Gubbio, Perugia). Pollen 
et Spores, 3, 93-100. 

Lona F., Ricciardi E. (1961b) - Reperti pollinologici nei 
depositi pleistocenici del bacino lacustre del Mer-
cure (Italia meridionale, Regione Lucano-
Calabra). Pollen et Spores, 3(1), 85-92.  

Lona F., Bertoldi R., Ricciardi E. (1969) - Plio-
Pleistocenic boundary in Italy based on the Leffian 
and Tiberian vegetational and climatological se-
quences. Verlag nicht ermittelbar. 

Lourens L., Antonarakou F., Hilgen F.J., Van Hoof A.M., 
Vergnaud-Grazzini C., Zacharias W.J. (1996) - 
Evaluation of the Plio-Pleistocene astronomical 
timescale. Paleoceanography, 11, 491-493.  

Lourens L.J., Hilgen F.J., Laskar J., Shackleton N.J., 
Wilson D. (2004) - The Neogene Period. In: Grad-
stein, F.M., Ogg, J.G., Smith, A.G. (Eds.), A Geo-
logic Time Scale 2004. Cambridge University 
Press, 409-440. 

Magri D. (1999) - Late-Quaternary vegetation history at 
Lagaccione near Lago di Bolsena (central Italy). 
Review of Palaeobotany and Palynology, 106, 171
-208. 

Magri D., Sadori L. (1999) - Late Pleistocene and Holo-
cene pollen stratigraphy at Lago di Vico (central 
Italy). Vegetation History and Archaeobotany, 8, 
247-260. 

Magri D., Palombo M.R. (2013) - Early to Middle Pleis-
tocene dynamics of plant and mammal communi-
ties in South West Europe. Quaternary Interna-
tional, 288, 63-72. 

Magri D., Di Rita F., Palombo M.R. (2010) - An early 
Pleistocene interglacial record from an intermon-
tane basin of central Italy (Scoppito, L'Aquila). 
Quaternary International, 225, 106-113. 

Magri D., Di Rita F., Aranbarri J., Fletcher W., González
-Sampériz P. (2017) - Quaternary disappearance 
of tree taxa from Southern Europe: Timing and 
trends. Quaternary Science Reviews, 163, 23-55. 

Mai D.H. (1994) - Fossile Koniferenreste in der meridio-
nalen Zone Europas. Feddes Repertorium. 105(3-
4), 207-227. 

Mai D.H. (1995) - Paleocarpological investigations in 
the Villafranchian (Pliocene) of Italy. Bollettino del 
Museo Regionale di Scienze Naturali Torino, 13, 
407-437. 

Manzi G., Magri D., Palombo M.R. (2011) - Early-Middle 
Pleistocene environmental changes and human 
evolution in the Italian peninsula. Quaternary Sci-
ence Reviews, 30, 1420-1438. 

Margari V., Roucoux K., Magri D., Manzi G., Tzedakis 
P.C. (2018) - The MIS 13 interglacial at Ceprano, 
Italy, in the context of Middle Pleistocene vegeta-
tion changes in southern Europe. Quaternary Sci-
ence Reviews, 199, 144-158. 

Marlow J. R., Lange C. B., Wefer G., Rosell-Melé A. 
(2000) - Upwelling intensification as part of the 
Pliocene-Pleistocene climate transition. Science, 
290(5500), 2288-2291. 

Martinetto E. (2009) - Palaeoenvironmental significance 
of plant macrofossils from the Piànico Formation, 
Middle Pleistocene of Lombardy, North Italy. Qua-
ternary International, 204, 20-30. 

Martinetto E., Vassio E. (2010) - Reconstructing “Plant 
Community Scenarios” by means of palaeocarpo-
logical data from the CENOFITA database, with 
an example from the Ca' Viettone site (Pliocene, 
Northern Italy). Quaternary International, 225 (1), 
25-36. 

Martinetto E., Scardia G., Varrone D. (2007) - Magneto-
biostratigraphy of the Stura di Lanzo fossil forest 
succession (Piedmont, Italy). Rivista Italiana di 
Paleontologia e Stratigrafia, 113 (1), 109-125. 

Martinetto E., Bertini A., Basilici G., Baldanza A., 
Bizzarri R., Cherin M., Gentili S., Pontini M.R. 
(2014) - The plant record of the Dunarobba and 
Pietrafitta sites in the Plio-Pleistocene palaeoenvi-
ronmental context of central Italy. Alpine Mediter-
ranean Quaternary (AMQ), 27 (1), 29-72. 

Martinetto E., Monegato G., Irace A., Vaiani S.C., 
Vassio E. (2015) - Pliocene and Early Pleistocene 
carpological records of terrestrial plants from the 
southern border of the Po Plain (northern Italy). 
Review of Palaeobotany and Palynology, 218, 148
-166. 

Martini I.P., Sagri M. (1993) - Tectono-sedimentary 
characteristics of Late Miocene-Quaternary exten-
sional basins of the Northern Apennines, Italy. 
Earth-Science Reviews, 34(3), 197-233. 

Masini F., Sala B. (2007) - Large- and small-mammal 
distribution patterns and chronostratigraphic 
boundaries from the Late Pliocene to the Middle 
Pleistocene of the Italian peninsula. Quaternary 
International, 160, 43-56. 

Massari F., Rio D., Sgavetti M., Prosser G., 
D'Alessandro A., Asioli A., Capraro L., Fornaciari 
E., Tateo F. (2002) - Interplay between tectonics 
and glacio-eustasy: Pleistocene succession of the 
Crotone Basin, Calabria (southern Italy). Geologi-
cal Society American Bullettin, 114, 1183-1209. 

Massari F., Rio D., Serandrei Barbero R., Asioli A., 
Capraro L., Fornaciari E., Vergerio P.P. (2004) - 
The environment of Venice area in the past two 
million years. Palaeogeography Palaeoclimato-
lology Palaeoecology, 202, 273-308. 

Masson-Delmotte V., Stenni B., Pol K., Braconnot P., 
Cattani O., Falourd S., Kageyama M., Jouzel J., 
Landais A., Minster B., Barnola J.M., Chappellaz 
J., Krinner G., Johnsen S., Röthlisberger R., 
Hansen J., Mikolajewicz U., Otto-Bliesner B. 
(2010) - EPICA Dome C record of glacial and 
interglacial intensities. Quaternary Science Re-
views, 29(1-2), 113-128. 

 
 

114 

 
 

Bertini A., Combourien-Nebout N. 



   

 

McClymont E.L., Sosdian S.M., Rosell-Melé A., 
Rosenthal Y. (2013) - Pleistocene sea-surface 
temperature evolution: Early cooling, delayed 
glacial intensification, and implications for the mid-
Pleistocene climate transition. Earth-Science Re-
views, 123, 173-193. 

Messager E., Lebreton V., Marquer L., Russo Ermolli, 
E. Orain R., Renault Miskovsky J., Lordkipanidze 
D., Despriee J., Peretto C., Arzarello M. (2011) - 
Palaeoenvironments of early hominins in temper-
ate and Mediterranean Eurasia: new palaeobotan-
ical data from Palaeolithic key-sites and synchro-
nous natural sequences. Quaternary Science 
Reviews, 30, 1439-1447.  

Moncel M.H., Landais A., Lebreton V., Combourieu-
Nebout N., Nomade S., Bazin L. (2018) - Linking 
environmental changes with human occupations 
between 900 and 400 ka in Western Europe. Qua-
ternary International, 480, 78-94. 

Monegatti P., Raffi S., Raineri G. (1997) - The Monte 
Falcone-Rio Riorzo composite section: biostrati-
graphic and ecobiostratigraphic remarks. Bolletti-
no della Società Paleontologica Italiana, 36, 245-
260. 

Monegatti P., Canali G., Bertoldi R., Albianelli A. (2002) 
- The classical late Piacenzian Monte Falcone-Rio 
Crevalese section (Northern Italy): palynological 
evidence and biomagnetostratigraphic constraints 
for climatic cyclicity and local mollusc extinctions. 
Geobios (Mem. Spéc.), 24, 219-227. 

Moscariello A., Ravazzi C., Brauer A., Mangili C., 
Chiesa S., Rossi S., de Beaulieu J.L., Reille M. 
(2000) - A long lacustrine record from the Piànico-
Sèllere basin (Middle-Late Pleistocene, northern 
Italy). Quaternary International, 73 (74), 47-68. 

Mullenders W., Favero V., Coremans M., Dirickx M. 
(1996) - Analyses pollinique de sondages à Ve-
nise (VE-I, VE-I bis, VE-II). Aardkundige 
Mededelingen, 7, 87-116.  

Munno R., Petrosino P., Romano P., Russo Ermolli E., 
Juvigné E. (2001) - A Late Middle Pleistocene 
climatic cycle in southern Italy inferred from pollen 
analysis and tefrostratigraphy of the Acerno lacus-
trine succession. Géographie physique et Quater-
naire, 55, 87-99.  

Muttoni G., Carcano C., Garzanti E., Ghielmi M., Piccin 
A., Pini R., Rogledi S., Sciunnach D. (2003) - On-
set of major Pleistocene glaciations in the Alps. 
Geology, 31, 989-992. 

Muttoni G., Ravazzi C., Breda M., Pini R., Laj C., Kissel 
C., Mazaud A., Garzanti E. (2007) - Magnetostrati-
graphic dating of an intensification of glacial activi-
ty in the southern Italian Alps during Marine Iso-
tope Stage 22. Quaternary Research, 67, 161-
173. 

Muttoni G., Scardia G., Kent D. V. (2010) - Human mi-
gration into Europe during the late Early Pleisto-
cene climate transition. Palaeogeography Palaeo-
climatology Palaeoecology, 296(1-2), 79-93. 

Muttoni G., Scardia G., Kent D.V. (2018) - Early hom-
inins in Europe: The Galerian migration hypothe-
sis. Quaternary Science Reviews, 180, 1-29. 

Napoleone G., Albianelli A., Azzaroli A., Bertini A., Magi 

M., Mazzini M. (2003) - Calibration of the upper 
Valdarno basin to the Plio-Pleistocene for correlat-
ing the Apennine continental sequences. II Qua-
ternario (Ital. J. Quat. Sci.), 16, 131-166. 

Narcisi B., Anselmi B. (1998) - Sedimentological investi-
gations on a Late-Quaternary core from the 
Lagaccione maar (central Italy): palaeoclimatic 
and palaeoenvironmental inferences. Quaternary 
International, 47/48, 21-28. 

Narcisi B., Anselmi B., Catalano F., Dai Pra G., Magri 
G. (1992) - Lithostratigraphy of the 250,000-year 
record of lacustrine sediments from the Valle di 
Castiglione Crater, Roma. Quaternary Science 
Reviews, 11, 353-362. 

Nomade S., Bassinot F., Marino M., Simon Q., Dewilde 
F., Maiorano P., Isguder G., Blamart D., Girone 
A., Scao V., Pereira A., Toti F., Bertini A., 
Combourieu-Nebout N., Peral M., Bourlès, D.L., 
Petrosino P., Gallicchio S., Ciaranfi N. (2019) - 
High-resolution foraminifer stable isotope record 
of MIS 19 at Montalbano Jonico, southern Italy: A 
window into Mediterranean climatic variability 
during a low-eccentricity interglacial. Quaternary 
Science Reviews, 205, 106-125. 

Orain R., Lebreton V., Russo Ermolli E., Combourieu-
Nebout N., Sémah A.-M. (2013) - Carya as marker 
for tree refuges in southern Italy (Boiano basin) at 
the Middle Pleistocene. Palaeogeography Palaeo-
climatolology Palaeoecology, 369, 295-302. 

Orain R., Ermolli E. R., Lebreton V., Di Donato V., 
Bahain J. J., Sémah A.M. (2015) - Vegetation 
sensitivity to local environmental factors and glob-
al climate changes during the Middle Pleistocene 
in southern Italy - A case study from the Molise 
Apennines. Review of Palaeobotany and Palynol-
ogy, 220, 69-77. 

O'Regan H.J., Turner A., Bishop L.C., Elton S., Lamb 
A.L. (2011) - Hominins without fellow travellers? 
First appearances and inferred dispersals of Afro-
Eurasian large mammals in the Plio-Pleistocene. 
Quaternary Science Review, 30, 1343-1352.  

Ozenda P. (1975) - Sur les étages de végétation dans 
les montagnes du bassin méditerranéen. Docu-
ments de cartographie écologique, 16, 1-32. 

Ozenda P., 1994. La végétation du continent européen. 
Delachaux and Nestlé eds, pp. 280. 

Pagani M., Caldeira K., Berner R., Beerling D.J. (2009) 
- The role of terrestrial plants in limiting atmos-
pheric CO2 decline over the past 24 million years. 
Nature, 460 (7251), 85-88. 

Palombo M.R. (2007) - Large mammal guilds on the 
Italian peninsula from the Pliocene to the Late 
Pleistocene: new methodological approaches. 
Courier Forschungsinstitut Senckenberg 259, 13-
46.  

Palombo M.R. (2014) - Deconstructing mammal disper-
sals and faunal dynamics in SW Europe during 
the Quaternary. Quaternary Science Reviews, 96, 
50-71.  

Petrosino P., Jicha B.R., Mazzeo F.C., Russo Ermolli E. 
(2014a) - A high resolution tephrochronological 
record of MIS 14-12 in the Southern Apennines 
(Acerno basin, Italy). Journal of Volcanology and 

 
 

115 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

Geothermal Research, 274, 34-50. 
Petrosino P., Russo Ermolli E., Donato P., Jicha B., 

Robustelli G., Sardella R. (2014b) - Using tephro-
chronology and palynology to date to the MIS 13 
lacustrine sediments of the Mercure Basin 
(Southern Apennine-Italy). Italian Journal of Geo-
sciences, 133(2), 169-186.  

Pieruccini P., Di Celma C., Di Rita F., Magri D., 
Carnevale G., Farabollini P., Ragaini L., Caffau, 
M. (2016) - Sedimentology, faunal content and 
pollen record of Middle Pleistocene palustrine and 
lagoonal sediments from the Peri-Adriatic basin, 
Abruzzi, eastern central Italy. Quaternary Re-
search, 86(3), 359-372.  

Pignatti S. (1982) - Flora d’Italia, 1-3. Bologna. 
Pignatti S. (1998) - I boschi d'Italia. Sinecologia e biodi-

versità. UTET, Torino, Italy (XVI + pp. 677). 
Pignatti G. (2011) - La vegetazione forestale di fronte 

ad alcuni scenari di cambiamento climatico in 
Italia. Forest 8, 1-12. 

Pini R., Ravazzi C., Donegana M. (2009a) - Pollen stra-
tigraphy, vegetation and climate history of the last 
215 ka in the Azzano Decimo core (plain of Friuli, 
north-eastern Italy). Quaternary Sciences Review, 
28, 1268-1290. 

Pini R., Ravazzi C., Monegato G. (2009b) - La storia 
vegetazionale e climatica del Pleistocene Superio-
re nella successione lacustre continua di Fimon 
(Colli Berici, Vicenza). Convegno AIQUA. “La 
variabilità del clima nel Quaternario: la ricerca 
italiana” 18-20, 76-77 (Roma). 

Pini R., Ravazzi C., Reimer P. J. (2010) - The vegeta-
tion and climate history of the last glacial cycle in 
a new pollen record from Lake Fimon (southern 
Alpine foreland, N-Italy). Quaternary Science Re-
views, 29(23-24), 3115-3137.  

Pini R., Bertini A., Martinetto E., Vassio E. (2014) - The 
Pleistocene flora of northern Italy. Palaeobotany 
of Italy. Publication of the Museum of Nature 
South Tyrol, (9), 290-307. 

Pini R., Furlanetto G., Vallé F., Badino F., Wick L., 
Anselmetti F. S., Bertuletti P., Fusi N., Morlock M., 
Delmonte B., Harrison S.P., Maggi V., Ravazzi C. 
(2022) - Linking North Atlantic and Alpine Last 
Glacial Maximum climates via a high-resolution 
pollen-based subarctic forest steppe record. Qua-
ternary Science Reviews, 294, 107759. 

Pinti D.L., Quidelleur X., Chiesa S., Ravazzi C., Gillot 
P.Y. (2001) - K-Ar dating of an early Middle Pleis-
tocene distal tephra in the interglacial varved suc-
cession of Piànico-Sèllere (Southern Alps, Italy). 
Earth Planetary Science Letters, 188, 1-7. 

Pontini M.R. (1997) - Analisi palinologica e interpreta-
zione paleoclimatica dei depositi continentali del 
Bacino Tiberino (Umbria, Italia). Unpublished 
Ph.D Thesis, University of Perugia, Italy, pp. 134. 

Pontini M.R., Bertini A. (2000) - Late Pliocene vegeta-
tion and climate in central Italy: high resolution 
pollen analysis from the Fosso Bianco section 
(Tiberino Basin). Geobios, 33, 519-526. 

Pontini M.R., Albianelli A., Basilici G., Bertini A., 
Napoleone G. (2002) - Pollen record and magne-
tostratigraphy of the Middle-Late Pliocene lacus-

trine sequence in the Tiberino basin, (central Ita-
ly). Bollettino della Società Geologica Italiana, 1, 
467-472. 

Quézel P., Médail F. (2003) - Ecologie et biogéographie 
des forêts du bassin méditerranéen, Elsevier-
Lavoisier eds, Paris, France, pp. 571. 

Raffi I., Rio D. (1980) - Nuove osservazioni sulla distri-
buzione del nannoplancton calcareo nella serie di 
Le Castella (Calabria). Paleontologia, Stratigrafia, 
Evoluzione, 1, 181-186. 

Ravazzi C. (1993) - Variazioni ambientali e climatiche al 
margine meridionale delle Alpi nel Pleistocene 
Inferiore: Stratigrafia e Analisi Pollinica della Serie 
di Leffe. (PhD Thesis). University of Milan, Italy. 

Ravazzi C. (2003) - Gli Antichi Bacini Lacustri e i Fossili 
di Leffe, Ranica e Piànico-Sèllere. CNR Special 
Publications. Quaderni di Geodinamica Alpina e 
Quaternaria, Bergamo, Italy (pp. 176). 

Ravazzi C., Rossignol-Strick M. (1994) - Interglacial-
glacial cycles on the Early Pleistocene of the Leffe 
basin (northern Italy): preliminary report. Historical 
Biology, 9, 11-15. 

Ravazzi C., Rossignol-Strick M. (1995) - Vegetation 
change in a climatic cycle of early Pleistocene age 
in the Leffe basin (Northern Italy). Palaeogeogra-
phy Palaeoclimatology Palaeoecology, 117, 105-
122. 

Ravazzi C. Moscariello A. (1998) - Sedimentation, pal-
aeoenvironmental evolution and time duration of 
earliest Pleistocene climatic cycles in the 24-56 m 
FM core interval (Leffe basin, Northern Italy). In: 
van Kolfschoten, Th., Gibbard, P. (Eds.), Proceed-
ings of the INQUA-SEQS Symposium “The dawn 
of the Quaternary”Meded. Ned. Inst. Toegep. Ge-
owet. TNO 60, 467-490. 

Ravazzi C., Pini R., Breda M., Martinetto E., Muttoni G., 
Chiesa S., Confortini F., Egli R. (2005) -The lacus-
trine deposits of Fornaci di Ranica (late Early 
Pleistocene, Italian Pre-Alps): stratigraphy, palae-
oenvironment and geological evolution. Quater-
nary International, 131, 35-58. 

Ravazzi C., Pini R., Breda M. (2009) - Reconstructing 
the palaeoenvironments of the early Pleistocene 
mammal faunas from the pollen preserved on 
fossil bones. Quaternary Sciences Review, 28, 
2940-2954.  

Raymo M.E., Ruddiman W.F., Backman J., Clement 
B.M., Martinson D.G. (1989) - Late Pliocene varia-
tion in northern hemisphere ice sheets and north 
Atlantic deep-water circulation. Paleoceanogra-
phy, 4 (4), 413-446. 

Raymo M.E., Oppo D.W., Flower B.P., Hodell D.A., 
McManus J.F., Venz K.A., Kleiven K.F., McIntyre 
K. (2004) - Stability of North Atlantic water masses 
in face of pronounced climate variability during the 
Pleistocene. Paleoceanography 19, PA2008. 

 Doi: 10.1029/2003PA000921 
Ricci M., Bertini A., Capezzuoli E, Horvatinčić N, 

Andrews J.E, Fauquette S, Fedi M. (2015) - Paly-
nological investigation of a Late Quaternary cal-
careous tufa and travertine deposit: the case 
study of Bagnoli in the Valdelsa Basin (Tuscany, 
central Italy). Review of Palaeobotany and Paly-

 
 

116 

 
 

Bertini A., Combourien-Nebout N. 



   

 

nology, 218, 184-197. 
Ricciardi E. (1961) - Analisi pollinologica dei sedimenti 

lacustri lignitiferi di Pietrafitta (Perugia, Italia). 
Pollen et Spores, 3 (2), 325-328. 

Ricciardi E. (1965) - Analisi polliniche di una serie strati-
grafica dei sedimenti lacustri del Pleistocene Infe-
riore nel Bacino di Leonessa (Rieti-Italia Centrale). 
Giornale Botanico Italiano, 72, 62-82. 

Rio D. (1982) - The fossil distribution of Coccolithophore 
Genus Gephyrocapsa Kamptner and related Plio-
Pleistocene chronostratigraphic problems. In: Prell 
W.L., Gardner J.V., et al. (Eds.). Initial Reports of 
the Deep Sea Drilling Project, 68, 325-343. 

Rio D., Thunell R., Sprovieri R., Bukry D., Di Stefano E., 
Howell M., Raffi I., Sancetta C., San Filippo A. 
(1989) - Stratigraphy and depositional history of 
the Pliocene Bianco section, Calabria, southern 
Italy. Palaeogeography Palaeoclimatology Palaeo-
ecology, 76, 85-105. 

Rio D., Raffi I., Villa G. (1990) - Pliocene-Pleistocene 
distribution patterns in the Western Mediterrane-
an. In: Karstetn K.A., Mascle J., et al. (Eds.), Pro-
ceedings of ODP, Scientific Results 107, 513-533. 

Rio D., Channell J.E.T., Bertoldi R., Poli M.S., Vergerio 
P.P., Raffi I., Sprovieri R., Thunell R.C. (1997) - 
Pliocene sapropels in the northern Adriatic area: 
chronology and paleoenvironmental significance. 
Palaeogeography Palaeoclimatology Palaeoecolo-
gy, 135, 1-25. 

Robustelli G., Russo Ermolli E., Petrosino P., Jicha B., 
Sardella R., Donato P. (2014) - Tectonic and cli-
matic control on geomorphological and sedimen-
tary evolution of the Mercure basin, southern Ap-
ennines, Italy. Geomorphology, 214, 423-435. 

Rossi S. (2003) - Etude pollinique de la séquence la-
custre Pléistocène de Piànico-Sèllere (Italie). 
(PhD thesis). Université de Droit, d'Economie et 
des Sciences d'Aix Marseille III. 

Rossignol-Strick M., Planchais N. (1989) - Climate pat-
terns revealed by pollen and oxygen isotope rec-
ords of a Tyrrhenian Sea core. Nature, 342, 413-
416. 

Rossignol-Strick M., Planchais N., Paterne M., Duzert 
D. (1992) - Vegetation dynamics and climate dur-
ing the last deglaciation in the South Adriatic basin 
from a marine record. Quaternary Sciences Re-
view, 11, 415-423. 

Roulleau E., Pinti D., Rouchon V., Quidelleur X., Gillot 
P.Y. (2009) - Tephrochronostratigraphy of the 
lacustrine interglacial record of Piànico, Italian 
Southern Alps: identifying the volcanic sources 
using radiogenic isotopes and trace elements. 
Quaternary International, 204, 31-43. 

Roveri M., Flecker R., Krijgsman W., Lofi J., Lugli S., 
Manzi V.,Sierro F.J., Bertini A., Camerlenghi A., 
De Lange G., Govers R., Hilgen F.J., Hübscher 
C., Meijer P.T., Stoica M. (2014) - The Messinian 
Salinity Crisis: Past and future of a great chal-
lenge for marine sciences. Marine Geology, 352, 
25-58. 

Ruddiman W.F., Raymo M.E., Martinson D.G., Clement 
B.M., Backman J. (1989) - Pleistocene evolution: 
northern hemisphere ice sheets and North Atlantic 

Ocean. Paleoceanography, 4 (4), 353-412. 
Russo Ermolli E. (1994) - Analyse pollinique de la suc-

cession lacustre pléistocène du Vallo di Diano 
(Campanie, Italie). Annales de la Société Géolo-
gique de Belgique, 117, 333-354. 

Russo Ermolli E. (1999) - Vegetation dynamics and 
climate changes at Camerota (Campania, Italy) at 
the Pliocene-Pleistocene boundary. II Quaternario 
(Ital. J. Quat. Sci.), 12, 207-214. 

Russo Ermolli E. (2000) - Pollen analysis of the Acerno 
palaeolacustrine succession (Middle Pleistocene, 
southern Italy). Geological Society of London 
Spec. Publ., 181, 151-159. 

Russo Ermolli E., Cheddadi R. (1997) - Climatic recon-
struction during the middle Pleistocene: a pollen 
record from Vallo di Diano (southern Italy). Geo-
bios, 30 (6), 735-744. 

Russo Ermolli E., di Pasquale G. (2002) - Vegetation 
dynamics of southwestern Italy in the last 20,000 
yr inferred from pollen analysis of a Tyrrhenian 
Sea core. Vegetation History and Archaeobotany, 
11 (3), 211-220. 

Russo Ermolli E., Juvigné E., Bernasconi S., 
Brancaccio L., Cinque A., Lirer, L., Ozer A., 
Santangelo N. (1995) - Le premier stratotype con-
tinental de quatre stades isotopiques successifs 
du Pléistocène moyen pour le bassin méditerra-
néen septentrional: le Vallo di Diano (Campanie, 
Italie). Comptes Rendus de l'Académie des Sci-
ences Paris, 321, 877-884. 

Russo Ermolli E., Aucelli P.P.C., Di Rollo A., Mattei M., 
Petrosino P., Porreca M., Rosskopf C.M. (2010a) - 
An integrated stratigraphical approach to the Mid-
dle Pleistocene succession of the Sessano basin 
(Molise, Italy). Quaternary International, 225, 114-
127. 

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

Sabato L., Bertini A., Masini F., Albianelli A., Napoleone 
G., Pieri P. (2005) - The lower and middle Pleisto-
cene geological record of the San Lorenzo lacus-
trine sequence in Sant'Arcangelo basin. (Southern 
Apennines, Italy). Quaternary International, 131, 
59-69. 

Sadori L., Giardini M., Chiarini E., Mattei M., 
Papasodaro F., Porreca M. (2010) - Pollen and 
macrofossil analyses of Pliocene lacustrine sedi-
ments (Salto river valley, central Italy). Quaternary 
International, 225, 44-57. 

Sadori L., Bertini A., Combourieu-Nebout N., Kouli K., 
Mariotti Lippi M., Roberts N., Mercuri A.M. (2013) - 
Palynology and Mediterranean vegetation history. 
Flora Mediterranea, 23, 141-156. 

Sánchez Goñi M.F., Cacho I., Turon J.L., Guiot J., 
Sierro F.J., Peypouquet J.P., Grimalt J.O., 
Shackleton N.J. (2002) - Synchroneity between 
marine and terrestrial responses to millenial scale 
climatic variability during the last glacial period in 
the Mediterranean region. Climate Dynamics, 19, 
95-105. 

 
 

117 

 
 
Piacenzian to Pleistocene flora and vegetation in Italy 



   

 

Sánchez Goñi M.F. (2022) - Pollen: A Key Tool for Un-
derstanding Climate, Vegetation, and Human Evo-
lution. In: Progress in Botany. Springer, Berlin, 
Heidelberg.  

 Doi: 10.1007/124_2022_63 
Sangiorgi L., Capotondi L., Combourieu-Nebout N., 

Vigliotti F., Brinkhuis H., Giunta S., Lotter A.F., 
Morigi C., Negri A., Reichart G.J. (2003) - Holo-
cene seasonal sea surface temperature variations 
in the South Adriatic Sea inferred from a multi-
proxy approach. Journal of Quaternary Science, 
18, 723-732. 

Santangelo N., Di Donato V., Lebreton V., Romano P., 
Russo Ermolli E. (2012) - Palaeolandscapes of 
Southern Apennines during the late Early and the 
Middle Pleistocene. Quaternary International, 267, 
20-29. 

Sarnthein M., Bartoli G., Prange M., Schmittner A., 
Schneider B., Weinelt M., Andersen N., 
Garbe-Schonberg D. (2009) - Mid-Pliocene shifts 
in ocean over-turning circulation and the onset of 
Quaternary-style climates. Climate of the Past, 5, 
269-283.  

Scardia G., Muttoni G. (2009) - Paleomagnetic investi-
gations on the Pleistocene lacustrine sequence of 
Piànico-Sèllere (northern Italy). Quaternary Inter-
national, 204(1-2), 44-53. 

Strani F., Sardella R., Mecozzi B. (2021) - An overview 
of the Middle Pleistocene in the north Mediterra-
nean region. Alpine and Mediterranean Quaterna-
ry (AMQ), 34(1), 5-16. 

Suc J.P., Bessais E. (1990) - Pérennité d'un climat ther-
mo-xérique en Sicile avant, pendant, après la 
crise de salinité messinnienne. Comptes Rendus 
de l'Académie des Sciences Paris, 294, 1003-
1008. 

Suc J.P., Bertini A., Combourieu-Nebout N., Diniz, F., 
Leroy S., Russo Ermolli E., Zheng Z., Bessais E., 
Ferrier J. (1995) - Structure of west Mediterranean 
and climate since 5.3 Ma. Acta Zoologica Craco-
viensa, 38, 3-16. 

Suc J.P., Combourieu-Nebout N., Seret G., Popescu S. 
M., Klotz S., Gautier F., Clauzon G., Westgate J., 
Insinga D., Sandhu A.S. (2010) - The Crotone 
series: a synthesis and new data. Quaternary 
International, 219(1-2), 121-133. 

Suc J.P., Popescu S.M. (2005) - Pollen records and 
climatic cycles in the North Mediterranean region 
since 2.7 Ma. Geological Society, London, Special 
Publications, 247(1), 147-158. 

Tomaselli R., Baldazzi A., Filipello S. (1973) - Carta 
Bioclimatica d'Italia. La vegetazione forestale d'Ita-
lia. Ministero Agricoltura e Foreste, Roma, 5-24. 

Toti F. (2018) - A Mediterranean perspective on the 
Early-Middle Pleistocene transition with emphasis 
on marine isotope stage 19. Unpublished PhD 
thesis, Università degli Studi di Firenze, pp. 388.  

Toti F., Bertini A. (2018) - Mediterranean pollen stratig-
raphy across the Early-Middle Pleistocene bound-
ary. Alpine and Mediterranean Quaternary (AMQ), 
31 (1), 199-203. 

Tzedakis P.C. (2005) - Towards an understanding of 
the response of southern European vegetation to 

orbital and suborbital climate variability. Quater-
nary Sciences Review, 25, 1585-1599. 

Tzedakis P.C. (2007) - Seven ambiguities in the Medi-
terranean palaeoenvironmental narrative. Quater-
nary Sciences Review, 26, 2042-2066. 

Tzedakis P.C., Channell J.E.T., Hodell D.A., Kleiven 
H.F., Skinner L.C. (2012) - Determining the natu-
ral length of the current interglacial. Nature Geo-
science, 5, 138-141. 

Urban B., Arias C., Bigazzi G., Bonadonna F. (1983) - 
Early Pleistocene palynostratigraphy of Fornace 
Tini, Valle Ricca (Central Italy). Palaeogeography 
Palaeoclimatology Palaeoecology, 41, 153-164. 

Vaiani S.C., Venezia P. (1999) - La sezione pleistoceni-
ca del Lamone (Appennino Romagnolo): associa-
zioni a foraminiferi ed evoluzione paleoambienta-
le. Bollettino della Società Paleontologica Italiana, 
38, 39-57. 

Valleri G., Bertoldi R., Bertini A. (1990) - Foraminifer 
and pollen assemblages from the Pliocene of Pon-
te an Elsa (Valdarno inferiore, Tuscany). Bollettino 
della Società Paleontologica Italiana, 29, 321-333. 

Van Couvering J. A., Castradori D., Cita M. B., Hilgen F. 
J., Rio D. (2000) - The base of the Zanclean Sta-
ge and of the Pliocene Series. Episodes Journal 
of International Geoscience, 23(3), 179-187. 

van Otterloo R.H., Sevink J. (1983) - The Quaternary 
Evolution of the Upper Volturno Basin. In: Isernia 
la Pineta (Ed.), Calderini, Bologna, 35-39. 

von Koenigswald W., Kolfschoften T. (1996) - The My-
momys arvicola boundary and the enamel thick-
ness quotient (SDQ) of Arvicola as stratigraphic 
markers in the Middle Pleistocene. In: Turner, C. 
(Ed.), The Early Middle Pleistocene in Europe. 
Balkema, Rotterdam, 211-226. 

Watts W.A., Allen J.R.M., Huntley B. (1996) - Vegeta-
tion history and palaeoclimate of the Last Glacial 
Period at Lago Grande di Monticchio, southern 
Italy. Quaternary Science Review, 15, 113-132. 

Watts W.A., Allen J.R.M., Huntley B. (2000) - Palaeoe-
cology of three interstadial events during oxygen-
isotope Stages 3 and 4: a lacustrine record from 
Lago Grande di Monticchio, southern Italy. Pal-
aeogeography Palaeoclimatology Palaeoecology, 
155, 83-93. 

Wutke K., Wulf S., Tomlinson E.L., Hardiman M., Dulski 
P., Luterbacher J., Brauer A. (2015) - Geochemi-
cal properties and environmental impacts of seven 
Campanian tephra layers deposited between 40 
and 38 ka BP in the varved lake sediments of 
Lago Grande di Monticchio, southern Italy. Qua-
ternary Science Reviews, 118, 67-83. 

Zagwijn W.H. (1957) - Vegetation, climate and time-
correlations in the early Pleistocene of Europe. 
Geologie en Mijnbouw 19, 233-244.  

Zagwijn W.H. (1959) - Zur stratigrafischen und polle-
nanalytischen Gliederung der pliozenen Ablager-
ungen im Roertal-Graben und Venloer Graben der 
Niederlande. Fortschritte in der Geologie von 
Rheinland und Westfalen 4, 5-26.  

Zagwijn W.H. (1960) - Aspects of the Pliocene and early 
Pleistocene vegetation in the Netherlands. 
Mededelingen van de Geologische Stichting C III, 

 
 

118 

 
 

Bertini A., Combourien-Nebout N. 



   

 

1-5. 1-78.  
Zagwijn W.H. (1963) - Pollen-analytical investigations in 

the Tiglian of The Netherlands. Mededelingen van 
de Geologische Stichting, 16, 49-71.  

Zagwijn W.H. (1975) - Variations in climate as shown by 
pollen analysis, especially in the Lower Pleisto-
cene of Europe. In: Wright A.E., Moseley F. 
(Eds.), Ice Ages: Ancient and Modern, 137-152.  

Zagwijn W.H. (1992) - The beginning of the ice age in 
Europe and its major subdivisions. Quaternary 
Science Reviews, 11, 583-591.  

Zagwijn W.H., Suc J.P. (1984) - Palynostratigraphie du 
Plio-Pléistocène d’Europe et de Méditerranée 
nord-occidentales: correlations chronostratigra-
phiques, histoire de la végétation et du climat. 
Paléobiologie Continentale, 14, 475-483. 

Zagwijn W.H., Hager H. (1987) - Correlations of conti-
nental and marine Neogene deposits in the south-
eastern Netherlands and the Lower Rhine District. 
Mededelingen van de Werkgroep voor Tertiaire en 
Kwartaire Geologie, 24 (1-2), 59-78.  

Zonneveld K.A.F. (1996) - Palaeoclimatic reconstruction 
of the last deglaciation (18 ka BP-8 ka BP) in the 
Adriatic Sea region: a land-sea correlation based 
on palynological evidence. Palaeogeography, 
Palaeoclimatology, Palaeoecology, 122(1-4), 89-
106. 

 

Ms. received: December 12, 2022 Revised: April 15, 2023 
Accepted: April 20, 2023  Avaiable online: May 10, 2023 

 
 

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