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IX/2/2018
INTERDISCIPLINARIA ARCHAEOLOGICA
NATURAL SCIENCES IN ARCHAEOLOGY
homepage: http://www.iansa.eu
Thematic Review
The “Mediterranean Forest”: A Perspective for Vegetation History Reconstruction
Marta Mariotti Lippi
a
, Anna Maria Mercuri
b*
, Bruno Foggi
a
a
Department of Biology, University of Florence, Via La Pira 4, 50121 Florence, Italy
b
Laboratory of Palynology and Palaeobotany, Department of Life Sciences, University of Modena and Reggio Emilia, Viale Caduti in Guerra 127,
41121 Modena, Italy
1. Introduction
“Mediterranean” and “Mediterranean Forest” are terms
frequently used: either in a too generic manner or in a very
specifc sense. Their meanings difer not only when used
by amateurs but also when they are mentioned in scientifc
papers, depending on the country, from the education and
research felds of scholars. The “Mediterranean diet”,
for instance, is not only an eating pattern of the countries
surrounding the Mediterranean sea, but also a way of
naming the “good food” prepared in several restaurants
of the world. Thus, to reply to what meant by the word
Mediterranean
, we can reply with Braudel (1985 in
Blondel
et al
., 2010): “One thousand of things at a time”.
Even referring to the Mediterranean Sea, the adjective
“Mediterranean” encompasses several diferent concepts:
it may indicate geographical location (Mediterranean
Basin, also pointing to the countries bordering the sea),
climate (Mediterranean climate) or botanical features
(Mediterranean fora and Mediterranean vegetation). This
paper focuses on the use of the term “Mediterranean” and
its interpretation in environmental/palaeoenvironmental
studies with a focus on the botanical/palynological approach.
It is known that the current landscape of plants is a result
of diferent factors, making vegetation history an important
chapter of the environmental sciences. The fora and
vegetation are basically the results of climatic changes that
have occurred over millions of years. Further variations
have been recently added in as a function of the human
interference that has drawn and re-drawn the plant landscape
through the development of diferent cultures during
the Holocene (Mercuri and Sadori, 2014; Sadori
et al.
,
2013). This seems to have been particularly true for the
Mediterranean Region, according to the opinion of Blondel
et al.
, (2010): “Nowhere else more than in the Mediterranean
Region has nature moulded people so much and have people
in turn so deeply infuenced landscapes”. The reason for this
deep interdependence between cultures and environment is
actually visible in the nature of the plant communities around
the Mediterranean Basin, a geographic area which shows
Volume IX ● Issue 2/2018 ● Pages 207
–218
*Corresponding author. E-mail: annamaria.mercuri@unimore.it
ARTICLE INFO
Article history:
Received: 22
nd
September 2018
Accepted: 31
st
December 2018
DOI: http://dx.doi.org/ 10.24916/iansa.2018.2.7
Key words:
foristic studies
palynology
sclerophyllous
Quercus
species
Meso-Mediterranean
ABSTRACT
Starting from the multifaceted meaning of “Mediterranean”, this thematic review wishes to reconnect
the palaeobotanical with the phytogeographical approach in the reconstruction of the Mediterranean
Forest of the past. The use of the term “Mediterranean” is somewhat ambiguous in its common use, and
has not an unequivocal meaning in diferent research felds. In botany, geographical-foristic studies
produce maps based on the distribution of the plant species; foristic-ecological studies, produce maps
that deal with the distribution of the plant communities and their relationships with diferent habitats.
This review reports on the diferent use of the term “Mediterranean” in geographical or foristic
studies, and on the way climate and plant distributions are used to defne the Mediterranean area. The
Mediterranean Forest through the palynological records is then shortly reported on. Pollen analysis
may be employed to reconstruct the Mediterranean Forest of the past but a number of problems make
this a difcult task: low pollen preservation, lack of diagnostic features at low taxonomical level,
and low pollen production of species which form the Mediterranean Forests. Variable images of this
vegetation are visible in diferent landscapes, but the Mediterranean Forest often remains a sort of
“ghost forest” in pollen spectra from the Mediterranean Region.
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a remarkable variety of topographical features, edaphic
conditions, and plant communities.
With regard to vegetation history, the palaeo- and
archaeo-botanical studies deal with the fora and vegetation
changes as evidenced in long-term chronological records.
Analyses of pollen and plant macroremains from sediment
strata and archaeological layers provide lists of plants that
better attends to the fora rather than the vegetation, but
references to plant communities are highly informative for
reconstructing landscapes. Single-case studies are often
limited in time and in space, whereas syntheses of several
sites allow for wide-ranging reconstructions that overcome
specifc local events (see, for example, Mercuri, 2014, for
cultural landscapes reconstructed through pollen analyses).
Local and regional studies can improve our knowledge on
the cause-and-efect patterns which have determined broad
palaeoenvironmental changes (sharp events or gradual
transformations) under the various climate and anthropic
infuences. Synthesis of the data in a coherent scheme is
needed for reconstruction of the vegetation history of each
region, but the many inhomogeneities in the terminology
concerning various vegetation types, often being referred to
in a generic or ambiguous way, is a major difculty in this
task. In papers on palaeo- and archaeo-botany, the authors
rarely explain to which plant community their results refer
to, even if important exceptions exist (
e.g.
Colombaroli
et al.
, 2009, focusing on the dynamics and history of
fres; Piovesan
et al.
, 2018, studying an application for the
restoration of forest ecosystems). Certainly we can say that
reaching an unequivocal, conclusive defnition of the term
“Mediterranean” is a hard task and is outside and not the
aim of this paper. According to “Conservation International:
Biodiversity Hotspots”
1
, the Mediterranean Basin is one of
the hotspots of plant biodiversity (22,500 species with 52%
of endemic species against more then 6000 species in other
parts of Europe). Due to this wealth of biodiversity, defning
the limits of the Mediterranean biogeographical area is a
topic that is deeply under discussion among bio-geographers.
Therefore, our main purpose is to make the reader aware of
the level of this difculty and try to make less ambiguous the
terminology referring to plant communities, and in particular
the “Mediterranean Forest”.
2. The geographical use of “Mediterranean” referring
to plant communities
Recently, a catastrophic event made it quite clear that in
layman’s terms “Mediterranean Forest” is used to indicate a
forest of the Mediterranean Region in a very generic way. On
March 2015, a terrifc storm scourged the coast of northern
Tuscany. The fall of numerous trees changed the face of
the Versilia coast. After this disaster, the local government
ofcials decided to restore the “natural vegetation”. In their
1
Mediterranean Basin September, 2011: http://www.biodiversityhotspots.
org/xp/hotspots/mediterranean/Pages/default.aspx.
view, the natural, Mediterranean vegetation of the area
consisted of woods dominated by umbrella pine often mixed
with holly oak. The media were speaking about the “wild
Mediterranean Forest”, meaning the woods that people are
used to seeing in the territory. They believed those woods
to be the natural vegetation of the area and in naming them
used a geographical term (Mediterranean) derived from
the proximity to the coast: But, are those woods the wild
Mediterranean vegetation? Is merely growing near the
Mediterranean Sea in itself sufcient to be some part of the
“Mediterranean vegetation”? The reality is more a mosaic
of several vegetation types that form parts of this so-called
“wild forest”, including pine plantations dominated by
Pinus pinea
2
and
Pinus pinaster
, that have been planted as
several reforestation events since Roman times, and were
then intensifed between the “600 and 800” ies (Giacomini,
1968; Mondino and Bernetti, 1998; Arrigoni, 1998), with
an undergrowth of
Quercus ilex
, and a European vegetation
consisting of deciduous trees such as
Quercus robur
,
Alnus
glutinosa
, and
Carpinus betulus
.
The geographical use of the term “Mediterranean Forest”
is also present in scientifc papers belonging to research felds
other than botany but concerning woodlands or bushlands of
the countries facing the Mediterranean Sea (
e.g.
Cosandey
et al.
, 2005, on experimental studies on forest hydrology). In
such papers, the term “Mediterranean Forest” and “macchia”
are often reported as synonyms. Indeed, they are both
vegetation types dominated by evergreen and sclerophyllous
species, but they largely difer in their dominating habitus/
growth forms: there is a prevalence of trees in the forest, and
shrubs in the macchia (Arrigoni, 1996). The name “macchia”
more properly refers to a very intricate, impenetrable plant
community characterized by densely-branched, evergreen,
sclerophyllous shrubs and climbing plants.
3. The use of the term “Mediterranean” in foristic
studies
Narrowing to botany, the specifc literature reveals great
difculty in fnding an unambiguous defnition/use of the
adjective “Mediterranean”. In general, two approaches can
be used to describe the plant resources of a territory: the
former is the geographical-foristic approach based on the
local fora leading to the identifcation of phytochoria,
i.e.
areas with similar compositions of plant species (Takhtajan,
1986); the latter is the foristic-ecological approach based on
the study of the distribution of plant communities and their
relationships (Kent, 2012).
Numerous maps of Italy based on the geographical-
foristic approach were published during the last century
(Fiori, 1923; Arrigoni, 1980, Romagnoli, 2003). They
are syntheses useful for the regionalization of areas on a
geographical (
i.e.
large) scale. In this type of map (Figure 1),
2
Species names according to Euro+Med PlantBase (http://www.
emplantbase.org/).
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209
the largest part of the Italian peninsula is included in the
Euro-Siberian Region while only a narrow strip along the
coastline, in addition to the whole of Sicily and Sardinia, is
attributed to the Mediterranean Region.
The foristic-ecological approach focuses on the species
that use the same local resources with diferent type of
interactions, from independence to full interdependence
(Kent, 2012). In Europe, phytosociological schools often
employed this type of study (Gehu and Rivas-Martinez,
1981; Blasi, 2010; Biondi, 2011). It is particularly useful
for studying plant communities on a local scale. The maps
based on the foristic-ecological approach (Rivas-Martinez
et al.
, 2004, modifed for Italy by Blasi and Biondi, 2017)
also limited the Mediterranean Region to the coast of the
Italian peninsula, and included the hills and mountains of
Calabria, Puglia, Sardinia and Sicily. The integration of
both the geographical-foristic and foristic-ecological maps
constitutes a valuable tool for the study of the past fora
and vegetation history at the local scale (see, for example,
Mariotti Lippi
et al.
, 2015, for the reconstruction of the
vegetation surrounding Grotta Paglicci-Apulia).
4. Climate and plant distribution to defne
the Mediterranean area
The Mediterranean climate is a typical, temperate, bi-
seasonal climate with the dry period – summer – coincident
with the season of highest temperatures, and with mild, wet
winters (Koppen, 1936). Climate has been used to defne
the borders of the Mediterranean area. Gaussen (1954)
used temperature as a single parameter; more specifcally,
he considered the average temperature of the coldest month
of the year – January or February – as one of the most
Figure 1.
Geographical-foristic map of
Italy (Arrigoni, 1980; Romagnoli, 2003).
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Table 1.
Most important trees and shrubs and the main syntaxa related to the forests of the Mediterranean area, arranged according to the vegetation zonal belt. Vegetation belt and the distribution of species are
according to Quezel and Medail (2003 modifed); species names follow Euro+Med PlantBase (http://www.emplantbase.org/home.html). Syntaxa are according to Mucina
et al.
(2016).
Mediterranean vegetation beltOro MediterraneanMontaneSupra MediterraneanMeso MediterraneanThermo Mediterranean
Coldest month mean
temperature
– 7 / - 3 °C -3 / 0°C 0 / 3°C 3 / 7°C > 7°C
Main syntaxa
Junipero-Pinetea sylvestrisQuercetalia pubescentisQuercetalia pubescentisQuercetalia ilicisPistacio-Rhamnetalia
Geranio striati-Fagion Rhododendro
pontici-Fagetalia orientalis
Querco-CedretaliaQuercetalia callipriniPistacio-Pinetalia
Querco-CedretaliaQuercetalia callipriniPinetalia halepensis
Quercetalia ilicis
Main trees and shrubs
Abies (A. alba, A. pinsapo, A. maroccana,
A. numidica)
Abies (A. alba subsp. apenninica, A.
pinsapo, A. maroccana, A. nebrodensis, A.
cephalonica, A. borisii-regi, A. numidica,
A. cilicica, A. bornumulleriana, A.
nordmanniana)
Cedrus (C. atlantica, C. libani, C.
deodara)
Cedrus (C. atlantica, C. libani)
Pinus (P. sylvestris, P. nigra complex)Pinus (P. heldreichii, P. nigra complex)Pinus (P. halepensis, P. brutia)
Juniperus (J. excelsa, J. phoenicea,
J. foetidissima, J. hemisphaerica, J.
drupacea, J. thurifera)
Junipers (Juniperus deltoides, J. drupacea,
J. oxycedrus, J. thurifera)
Cupressus sempervirensCupressus atlantica
deciduous
Quercus (Q. brandtii, Q.
petraea)
deciduous
Quercus (Q. petraea)
marcescent
Quercus (Q. pyrenaica, Q.
pubescens, Q. cerris, Q. frainetto, Q.
faginea, Q. infectoria, Q. libani)
marcescent
Quercus (Q. pyrenaica, Q.
pubescens, Q. ithaburensis)
marcescent
Quercus (Q. ithaburensis, Q.
pubescens)
evergreen
Quercus (Q. brandtii, Q.
petraea, Q. ilex, Q. rotundifolia)
evergreen
Quercus (Q. rotundifolia. Q.
faginea, Q. ilex)
evergreen
Quercus (Q. rotundifolia. Q.
faginea, Q. ilex, Q. coccifera, Q. alnifolia,
Q. suber)
evergreen
Quercus (Q. lusitanica, Q.
calliprini)
Berberis (B. aetnensis, B. cretica)
Prunus prostrata
Cytisus (C. oromediterraneaus, C.
balansae, C. spinescens, C. aolicus, etc.)
Ostrya carpinifolia
Carpinus (C. betulus, C. orientalis)
Arbutus (A. unedo, A. andrachne)
Erica spp.
Acer sempervirens
Fraxinus ornus
Pistacia atlanticaPistacia (P. lentiscus, P. atlantica)
Olea europaea
Myrtus communis
Phillyrea (P. latifolia, P. angustifolia)
Euphorbia dendroides
Rhamnus (R. alaternus, R. oleoides, R.
lycioides)
Ceratonia siliqua
Cneorum tricoccum
Teucrium fruticans
Globularia alypum
Tetraclinis articularis
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relevant parameters, establishing a mean value >5°C as the
limit for the Mediterranean area. This value allows a line
to be drawn that includes the Mediterranean coastal areas
of Europe, West Asia and North-West Africa, and excludes
part of North-East Africa. In a similar vein, Desfontaines
(in De Montgolfer, 2002) took into consideration only one
factor, precipitation, and paid attention to the length of the dry
season and distribution of rainfall over the year. Considering
the occurrence of 1–3 months of aridity as a benchmark, he
moved the limits of the Mediterranean area to Spain, North
Africa and the Middle East. Northern Italy, which was out
of the Mediterranean area in Gaussen’s map, has hence been
partially included.
Taking into consideration the distribution of plant
communities as an indicator of bioclimatic areas, the
dominance of evergreen, sclerophyllous species was
historically used for defning the Mediterranean area. With
this perspective (Table 1), the Meso-Mediterranean Forest
displays the physiognomy of a forest dominated by
Quercus
species, especially
Q. ilex
and
Q. rotundifolia
. Remarkably,
these oak trees do not tolerate extreme temperatures
and drought conditions and are therefore substituted by
sclerophyllous, tall shrubs in a few areas of southern Italy
and in large areas of northern Africa. Such plant communities
are Thermo-Mediterranean communities, characterized by a
few species of small evergreen trees with non-dense foliage
(“
arboscellus
” according to Arrigoni, 1996) such as
Olea
europaea
and
Ceratonia siliqua
,
Acacia
species and shrubs
that generally give rise to open formations, or forest with
pines (such as
P. halepensis
) .
In the Mediterranean Region (1–5 of Figure 1) the forests
dominated by
Quercus ilex
are difusely spread from the
coastal areas to the mountain.
Q. ilex
can also occur in forests
of the Eurosiberian Region (6–8 of Figure 1) dominated by
deciduous and/or marcescent trees, like
Q. pubescens
and
sometimes
Q. cerris
and
Q. frainetto
(marcescent according
to Quezel and Medail, 2003; Garcia-Mijangos
et al.
, 2015,
i.e.
plants that maintain dried leaves for most part of the
winter and are able to have photosynthetic activity for 1–2
months longer than true deciduous trees, at least for parts of
the leaves).
Within this general scheme, topography, soil features,
temperatures and rainfall have interacted with each other
forming a variety of habitats that defy any resolution of
continuity. In addition, human interference has also afected
fora assemblages, shaping and re-shaping landscapes during
Table 2.
Number of genera and species of the most important families of the Mediterranean woody fora. Number of genera and species are according
to Kubitzki’s Families and Genera of Vascular plants (1990–2011). Data for the Mediterranean area are from Euro+MedPlantBase. Alien species, species
growing out of the Mediterranean area and exclusive of the Atlantic islands (Canary Islands, Madeira and Azores) are excluded. 1: Page, 1992a; 2: Page,
1992 b; 3: Kubitzki, 1993; 4: Rohwer, 1993a; 5: Rohwer, 1993b; 6: Hüber, 1998; 7: Kubitzki and Rudall, 1998; 8: Conran, 1998; 9: Mayo
et al.
, 1998;
10: Dransfeld and Uhl, 1998; 11: Stevens
et al.
, 2004; 12: Medan and Schirarend, 2004; 13: Green, 2004; 14: Köhler, 2007; 15: Wen, 2007; 16: Pell
et al.
,
2011; 17: Wilson, 2011; 18: Kubitzki
et al.
, 2011. a: The PlantList.org.
FamilyTotal n.
genera
Total n.
Species
Euro+med
genera
Euro+med
species
Distribution of the familyPollination agent
1Cupressaceae 20125311WorldwideWind
2Pinaceae 12200325WorldwideWind
3Fagaceae 8
620/750
340
Temperate, Subtropics (Worldwide)
Wind
4Moraceae 3711001
Ficus
4Tropics, Subtropics, Warm Temperate
Insects (Wind ?)
5Lauraceae 502500/35001
Laurus
1Tropics, Subtropics, Temperate
Insects (Wind ?)
6
Dioscoreaceae 20
600
1
Dioscorea
6
Tropics, Subtropics, Warm TemperateInsects
7Asparagaceae 1170/3001
Asparagus
33Tropics, Subtropics, MediterraneanInsects/Zoo
8Smilacaceae 33201
Smilax
2Tropics, Subtropics, Mediterranean
Insects/Zoo (Wind ?)
9Araceae1043300757
Tropics (Worldwide)Insects (Wind ?)
10Palmae19020004
6
Tropics, Subtropics, MediterraneanInsects/Zoo
11Ericaecae1244100422Worldwide Temperate, Tropics montaneInsects/Wind
12Rhamnaceae 52925547Tropics, Subtropics, Warm Temperate
Insects (Wind ?)
13
Oleaceae
25
600
917
Tropics, Subtropics (Temperate)
Insects/Wind
14
Buxaceae
51001
Buxus
2
Tropics, Subtropics, Mediterranean
(Wordlwide)
Insects/Zoo(Wind)
15Vitaceae 147503 3Tropics, Subtropics, TemperateInsects/Wind
16
Anacardiaceae 81800417Tropics, Subtropics, Warm Temperate
Insects (Wind)
17Myrtaceae14255003 4Tropics, Subtropics, Mediterranean
Insects/Zoo (Wind ?)
18Rutaceae1542100340Tropics, Subtropics, MediterraneanInsects/Zoo
aApocynaceae402503120
68
Tropics, Subtropics, Warm TemperateInsects/Zoo
aAraliaceae
46
15051
Hedera
12Tropics, Subtropics, Warm Temperate
Insects (Wind ?)
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the development of diferent cultures over the millennia.
According to Naveh and Dan (1973), the Mediterranean
plant landscape was made by “several variants of diferent
states of degradation and regeneration of the original forest”
that gave origin to the distribution in time and space of a
mosaic of vegetation formations.
5. The “Mediterranean Forest” through
palynological records
Reconstruction of long-term fora and vegetation changes
is a feld of research that has been especially developed
by palynology
(
e.g.
de Beaulieu
et al
., 2005; Jalut
et al
.,
2009; Roberts et al., 2011; Magri
et al
., 2015), even in
interdisciplinary cooperations between the human and
natural sciences (Sadori
et al
., 2015; Mariotti Lippi
et al
.,
2015; Mercuri
et al
., 2015; Mercuri and Florenzano in press).
Despite the fundamental role of palynology in vegetation
history, there are limits to delineating the pollen profle of
the Mediterranean Forest. The main problems are low pollen
preservation, lack of features diagnostic at low taxonomical
levels, and the low pollen production of many species that
form the Mediterranean Forest.
Regarding preservation, for example, it is generally
accepted that the pollen wall may be damaged by an alkaline
pH causing the deterioration of exine sporopollenins. As the
“Mediterranean Forest” dominated by
Q. ilex
is known to
prefer calcareous soils, which are characterized by high pH
values, it is likely that in such Mediterranean forest soils
pollen has low preservation.
Regarding identifcation, pollen morphology does not
always allow discrimination to the species, genus, or even
family level. Taking
Juniperus
as an example, its pollen is
commonly attributed to the
Juniperus
type, a group of pollen
grains that includes also
Cupressus
(Moore
et al.
, 1981).
Cupressaceae pollen is known for the fragility of its wall
(Spieksma
et al.
, 1994) and for its peculiar behaviour during
hydration, which provokes the breakage of the thin exine
layer as a consequence of the swelling of the intine (Danti
et al.
, 2011). The low resolution of its pollen morphology
may cause its identifcation to stop at the family level, making
pollen data from this important family useless for foristic
and phytogeographical studies. Other gymnosperms, like
Pinus
, are also hardly identifed at species level in ancient
sediments.
Regarding pollen production, many plant species of the
Mediterranean Forest (Table 2) are pollinated by animals,
insects in particular, and are weak pollen producers
(Blondel and Aronson, 1995). This feature may be related
to the tropical origin of many species, as indicated by the
current distribution of families that belong to, for example,
Asparagaceae, Dioscoreaceae, Moraceae, and Rhamnaceae.
The production of small amounts of pollen makes these
zoophilous plants underrepresented and infrequently
recorded in pollen spectra (Carrión 2002a; Mercuri
et al.
,
2010). Exceptions have been reported which may reveal a
very local presence of fowers/plants, such as in the case of
Arbutus unedo
(a shrub which is up to subdominant in
Q. ilex
wood and dominant in “tall macchia” in Drescher-Schneider
et al.
, 2007). In the case of
Pistacia
, contrasting records
from high (55% in Tinner
et al.
, 2009) to low values (3% in
Florenzano
et al.
, 2013) suggest that these plants may have
a rather variable image in the pollen spectra from southern
Italy.
With its nature of anemophilous tree,
Quercus ilex
is
generally the best represented plant of the Mediterranean
Forest in pollen spectra from sites located inside (
e.g.
Gruger
and Thulin, 1998) or near (
e.g.
Colombaroli
et al.
, 2007;
Magri and Sadori, 1999; Sadori and Narcisi, 2001) to the
border of the Mediterranean Region (Figure 2). Studies on
the current difusion of
Q. ilex
pollen in Tuscany (Mariotti
Lippi
et al.
, 2000) has given evidence that, at the same site,
Q. ilex
has similar values of pollen percentages in the spectra
as it has dominance in the plant community
3
. As
Quercus
ilex
is not exclusive to the Meso-Mediterranean Forest and
also grows in the Supra-Mediterranean Forest (according to
EUNIS classifcation: G2.122 Supra-Mediterranean Quercus
ilex Forests; 9340 Directive Habitat: Lauvel
et al.
, 2013), we
must take into account that its pollen can belong to plants
growing in both the Meso-Mediterranean Forest and the
Supra-Mediterranean Forest communities. To identify the
Mediterranean Forest from a palynological point of view,
it is thus necessary to look for pollen grains produced by
other species of this plant community. The deciduous and
marcescent species
of
Quercus
, growing in association
with
Q. ilex
(Table 1), do not give good hints because
their
pollen is rarely identifable to species level. Despite
the low amount of pollen produced, suitable plants for
identifying the Mediterranean Forest are
Arbutus
,
Pistacia
and
Fraxinus ornus
(the latter is also in SubMediterranean
forests, and its well-preserved pollen grains are discernible
from those of other ash species: Moore
et al.
, 1991).
Myrtus
,
Phillyrea
and
Rhamnus
may be added to the list
of plants suitable for identifying the Mediterranean Forest,
more properly belonging to the Thermo-Mediterranean
community. Pollen from climbers, as
Smilax
, or herbaceous
plants, such as
Asparagus
, may also support an attribution
to the Mediterranean Forest. The
Asparagus
inclusion
may be controversial: in fact,
Asparagus acutifolius
grows
in the
Quercus ilex
forest (
Quercion ilicis
or
Fraxino-
Quercion ilicis
) and in the SubMediterranean forest
(
Quercetalia pubescentis
), and
A. ofcinalis
is grown in a
variety of habitats and is not typical of the Mediterranean
Forests. However, the presence of this rare pollen, which
unfortunately cannot be easily identifed at species level,
may be taken into consideration to support the attribution of
certain pollen spectra to a Mediterranean Forest.
Indeed,
Ostrya carpinifolia
and
Carpinus orientalis
(often
combined in one single pollen type), together with
C. betulus
,
are suitable for identifying the Supra-Mediterranean Forest
(Tables 1 and 2).
3
Dominance is the cover value of the species in the community.
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As reported above, the Mediterranean Forest is a plant
community dominated by evergreen sclerophyllous (high and
low) trees and shrubs. These communities largely consist of
Quercus ilex
in the central part of the Mediterranean Basin,
Q. rotundifolia
to the western side,
Q. coccifera
subsp.
coccifera
in both, and
Q. coccifera
subsp.
calliprinos
to the
eastern side.
Based on pollen morphology, the discrimination of oaks
is quite problematic: the
Q. ilex
type includes the greater
part of the evergreen oaks, though not
Q. suber
, and is
discriminated by the semideciduous and deciduous oak types
(Smit, 1973). They are high pollen-producers because they
are anemophilous plants, as well as the gymnosperms such
as
Pinus
and Cupressaceae. Pollen from these plants largely
overwhelms those of the other components of the same plant
community. They often seem to be prevalent in the woody
fora of the Mediterranean, but this is a mere efect of their
masking efect over the entomophilous species which actually
prevail in the Mediterranean contexts. All these conditions
make the Mediterranean Forest poorly identifable in the
pollen spectra from the Mediterranean Region.
6. Postglacial history of the Mediterranean Forest
An enormous set of biological archives are available to
provide data on the changes in the fora and vegetation of
the Mediterranean regions. These changes have occurred
not only in the history of the far past but also in recent
times, together determining the shape of the present-day
Mediterranean landscape (Sadori
et al.
, 2013).
Moving on to the tracks of the Mediterranean Forest as
seen through the pollen spectra, we must take into account
the oscillations of the pollen sum of all the species living
in the Mediterranean Forest. As reported above, the pollen
does not allow the discrimination of plants growing in the
Meso-Mediterranean, Supra-Mediterranean or Thermo-
Mediterranean communities. However, the occurrence
of pollen belonging to elements typical of the Meso-
Mediterranean community (Table 1) seems to have been more
frequent in the pollen spectra, and confrms the occurrence
(but not its extension) of this plant formation in the past.
In the western part of the Mediterranean Basin, at the
beginning of the postglacial period, the Mediterranean
Figure 2.
Pollen percentages of the plants growing in the Mediterranean Forest (
Quercus ilex
,
Arbutus
,
Pistacia
,
Myrtus
,
Rhamnus
,
Asparagus
) inside or
close to the Mediterranean Ecoregion as indicated by pollen analyses of selected sites. Data from Poviglio and Montale (located in northern Italy) are added
to include comparison with the Euro-Siberian Region.
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elements are low and discontinuous in pollen records
(Carrión, 2002b; Gil-Romera
et al.
, 2009). Indeed, in the early
part of the Holocene, the vegetation cover was dominated by
deciduous broadleaved trees (Jalut
et al.
, 2009). In Spain,
Mediterranean communities spread in the mid-Holocene,
in concomitance with the increasing aridity, with maximum
values at around 5900 cal. BP
4
.
Pistacia
is well represented,
as are the evergreen
Quercus
,
Phillyrea
and
Rhamnus
. A
similar scenario also resulted from the study of a core from
Sète, in France, where
Pistacia
, mainly
P. terebinthus
, was
already present in the 6
th
millennium BP and continued to
spread until the end of the 5
th
millennium (Court-Picon
et al.
,
2010). Thus, the difusion of the Mediterranean Forest in the
western part of the Mediterranean Basin may be placed to
the beginning of the mid-Holocene, a phase correlated with
lake-level lowering in the Jura (eastern France) and glacier
retreat in the Alps (Magny
et al.
, 2002; Jalut
et al.
, 2009).
In the central part of the Mediterranean Basin, the
Mediterranean Forest is clearly visible in Holocene spectra
of the central and southern Italian regions (
e.g.
Magri and
Sadori, 1999; Sadori and Narcisi, 2001; Sadori
et al.
, 2013).
In Sicily,
Q. ilex
pollen is well recorded along the Holocene
stratigraphy, showing greater values in the upper part of the
sequence.
Fraxinus ornus
and
Pistacia
were intermittently
present at the beginning of the Holocene, but their amount
increased upwards with peak of
Pistacia
pollen occurring in
samples dated around 7200 BP, when other Mediterranean
elements were also abundant (Sadori and Narcisi, 2001). In
northern Italy, the occurrence of the Mediterranean Forest is
sporadically recorded along the Tyrrhenian coasts, because of
the occurrence of coastal wetlands (Bellini
et al.
, 2009) and
the presence of an extensive alluvial plain in north-eastern
Italy. Where it was found,
Q. ilex
pollen shows increasing
values from around the 6
th
millennium (6000–5000 cal
BP), sporadically accompanied by
Pistacia
(Arobba
et al.
,
2004; Colombaroli
et al.
, 2007; Montanari
et al.
, 2015).
In Dalmatia, evergreen vegetation, mainly
Juniperus
and
Phillyrea
, replaced the deciduous oak forests in the 8
th
millennium (about 7600 BP), while evergreen oaks spread
one millennium later (about 6100 BP; Sadori
et al.
, 2011).
In the western part of the Mediterranean Basin, at Delfnos
in Crete, Greece,
Quercus coccifera
-type pollen is recorded
from about 8400 BP and presents an increasing trend up
to about 6000 BP.
Pistacia
is discontinuous and peaks in
coincidence with
Fraxinus ornus
or
Phillyrea
(Bottema and
Sarpaki, 2003). Forests with
Q. calliprinos
as the dominant
tree in association with
Pistacia palaestina
were widespread
in the Mediterranean area of Israel from 6000 BP to 400 BP
(Liphschitz and Biger, 1990). A mild early Holocene in the
eastern part of the Mediterranean Basin is supported by the
record of
Pistacia
pollen in terrestrial sites and marine cores
(Rossignol-Strick, 1999).
Altogether, pollen records suggest that the Mediterranean
Forest was already developed in the eastern part of the
Mediterranean Basin at the beginning of the Holocene,
4
The chronologies of the paragraph are according to the original papers.
and then spread to the western and central parts of the
Mediterranean Basin in coincidence with the decline of the
deciduous broadleaved forest during the mid-Holocene.
7. Mediterranean Forest and human activities
The mid-Holocene was not only a period of climate changes
and the difusion of the Mediterranean Forest in the central
and western Mediterranean basin, but it also represents a
crucial period in human history. The socio-cultural changes
are the starting point towards new human-environment
relationships, deeply afecting the territory surrounding
human settlements. Therefore, it is not always easy to
distinguish the changes that have been induced by climate,
by human practices, or by both these causes as they coupled
and overlapped (Roberts
et al.
, 2011; Sadori
et al.
, 2011;
Marignani
et al.
, 2017), with an increasing importance of
the human impact with time, as is also happening currently
(Barbero
et al.
, 1990). In general, the spread of agrarian
systems, including feld cultivation and pastures, caused
a dramatic decrease in woodlands, especially visible in
continental and Mediterranean Italy since the Bronze Age
(
e.g.
, in Lago di Mezzano: Mercuri and Sadori, 2012; Sadori,
2018). Moreover, the pastoral economy has been largely
responsible for the limitation of forests and the spreading of
shrubby Mediterranean vegetation formations in southern
Italy (
e.g.
, Florenzano, 2019).
Slash and burn are among the most ancient activities
documented in human history. Past fre activity has been
recorded all over the world during the whole Holocene.
Studies of the history of fre indicate that changes in fre
regimes are directly related to climatic changes (Mouillot
and Field, 2005; Vannière
et al.
, 2011). Either way, a direct
connection between the incidence of fre and the presence
of human populations appears during the middle to the late
Holocene (Vannière
et al.
, 2011), just in coincidence with the
development of the Mediterranean cultures. Fires in forested
areas may have been voluntarily set to attain open areas for
pastures. Traces of ancient fre activity are also recorded in
areas where palaeonvironmental reconstruction indicates the
occurrence of Meso-Mediterranean communities (Vannière
et al.
, 2011).
Besides fre, cutting was a common practice for
implementing crop cultivation. By way of example, fre
seems to have prevented the spread of
Q. ilex
during the mid-
Holocene in northern Tuscany, in Italy (Colombaroli
et al.
,
2007); a decline in deciduous and evergreen oaks has been
recorded in Crete, Greece, in coincidence with an increasing
amount of
Olea
pollen at about 5700 BP, suggesting the
planting of olive groves from the foothills towards the sea
(Bottema and Sarpaki, 2003). Indeed, olive groves deeply
impacted the Mediterranean landscape, often replacing pre-
existing forests (Moriondo
et al.
, 2013), and despite the
variable pollen production of the trees, the rise of the
Olea
pollen curve in pollen spectra signals the development of
cultural landscapes and the arboriculture of the last three
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215
millennia in many Mediterranean countries (Mercuri
et al.
,
2013; Florenzano
et al.
, 2017).
With respects to the exploitation of natural resources,
wood-cutting and coppicing have impacted the forests around
the Mediterranean Basin for millennia. Among the main
trees of the Meso-Mediterranean Forest, evergreen oaks have
been used as a source of timber.
Q. ilex
did not seem to have
been selected by humans, as it happened to
Q. suber
(Toumi
and Lumaret, 1998).
Q. ilex
wood was largely used as fuel or
employed for the construction of elements of ship structures
(
e.g.
Giachi
et al.
, 2003). According to Reille (1992), human
action may also be responsible for the establishment of
Q.
ilex
, as indicated through pollen analysis in Corsica (France)
during the Subatlantic, where its difusion seems to have
been a consequence of the deforestation that mainly afected
Pinus
and
Erica arborea
.
Even if the typical elements of the Meso-Mediterranean
forest have been exploited by humans, few of them have
been subjected to cultivation, for example,
Q. suber
and
Q.
coccifera
; the former was cultivated for cork production, the
latter for the production of the kermes dye, obtained from
the insect
Kermes vermilio
Planchon, 1864, Hemiptera.
Cultivation possibly also favoured the difusion of these oaks
beyond their original geographic areas.
8. Concluding remarks
In literature, the term “Mediterranean Forest” is often used
to indicate the three diferent plant communities: Thermo-,
Meso- and Supra-Mediterranean communities. The Thermo-
Mediterranean community generally does not display the
true physiognomy of a forest. Despite the fact it includes the
emblem Mediterranean species
Olea europaea
, this is not
the most appropriate plant association for a Mediterranean
Forest. Moreover, the cultivation of
Olea
for a long time has
also favoured the spread of this plant beyond their natural
geographic areas. Other plant communities,
i.e.
the Pine
forests, have to be considered a variety of the “Mediterranean
Forest”, but they are hardly identifed in pollen analyses.
According to these facts, in the palaeoenvironmental
reconstruction, the “Mediterranean forest”
sensu stricto
can be considered the forest dominated by evergreen,
sclerophyllous
Quercus
species (Figure 3). This is probably
the best Mediterranean Forest detectable by pollen analysis,
but unfortunately often remains a sort of “ghost forest” in
pollen spectra from the Mediterranean Region.
Acknowledgments
The authors wish to thank the referees for the suggestions
useful for improving the text.
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