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Ecology / cologie

An ecological map of Europe: why and how?


Paul Ozenda
a
, Jean-Luc Borel
a
*
a
Laboratoire cosystmes et changements environnementaux, centre de biologie alpine, universit Joseph-
Fourier, BP 53, 38041 Grenoble cedex 9, France
Received 27 March 2000; accepted 20 June 2000
Communicated by Paul Ozenda
Abstract After a recall of the recent emergence of the great ecological problems on the
scale of the European continent and of the privileged use of vegetation maps as an
ecological tool, a cartographic synthesis of the main plant formations in Europe is
submitted on the basis of a hierarchized and numerized nomenclature of the vegetation
units. Examples of connections between vegetation and ecology, as suggested or
facilitated by this map, are given: use of the monthly ombrothermic diagrams as a
privileged tool, thermic limits of the boreal zone, indicative value of the Mediterranean
xerothermic area, predictive models of the geographical shifts of the great ecosystems
according to the expected climate changes. 2000 Acadmie des sciences/ditions
scientiques et mdicales Elsevier SAS
plant ecology / phytogeography / vegetation of Europe / phytoecological mapping
Rsum Une carte cologique de lEurope : pourquoi et comment ? Aprs un
rappel de lmergence rcente des grands problmes cologiques lchelle du
continent europen et de lutilisation privilgie des cartes de vgtation comme
instrument danalyse cologique, une synthse cartographique des formations vgtales
de lEurope est prsente sur la base dune nomenclature hirarchise et numrise des
units retenues. Des exemples de liaison entre vgtation et cologie, suggrs ou
facilits par cette carte, sont exposs : utilisation des diagrammes pluviothermiques
mensuels comme outil privilgi, limites thermiques de la zone borale, valeur
indicatrice de la plage xrothermique mditerranenne, modle prdictif du dplace-
ment gographique des grands cosystmes vgtaux en fonction des changements
climatiques attendus. 2000 Acadmie des sciences/ditions scientiques et mdicales
Elsevier SAS
cologie vgtale / phytogographie / vgtation de lEurope / cartographie phytocologique
Version abrge
Lampleur croissante des problmes lis lenviron-
nement, la gestion durable des ressources naturelles
et aux transformations, notamment sous leffet des
changements climatiques en cours ou venir, justie
dintensier les recherches concernant les grandes
rgions cologiques du continent europen. La ralisa-
tion dune carte cologique de lEurope apparat comme
un moyen privilgi de faire progresser rapidement ces
recherches et den exprimer clairement une synthse.
Lutilisation des cartes de vgtation sous leur forme
moderne semble constituer la meilleure approche :
une nouvelle carte gnrale de la vgtation du conti-
nent europen a donc t tablie sur la base des
documents biogographiques les plus rcents. Prsen-
te ici sous un format rduit, elle spare soixante-six
formations vgtales, dont une hirarchie homogne
assortie dun code numrique est propose.
Mais entre la cartographie des facteurs bioclimati-
ques et celle de la vgtation existe encore un hiatus
qui ncessite lintroduction dune mthodologie origi-
* Correspondence and reprints.
E-mail address: borel@lamasig.ujf-grenoble.fr (Jean-Luc Borel).
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C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
2000 Acadmie des sciences/ditions scientiques et mdicales Elsevier SAS. Tous droits rservs
S0764446900012270/FLA
nale pour intgrer ces deux familles de documents en
une reprsentation des grands cosystmes. Lutilisa-
tion des diagrammes pluviothermiques mensuels est
prsente comme un outil privilgi par la densit des
stations renseignes et la diversit des paramtres qui
peuvent tre lus ou dduits, outil dont lexploitation
pourrait tre davantage valorise. Il sagit ensuite
daffecter chaque formation les paramtres cologi-
ques les plus signicatifs, cest--dire de poser les bases
dune cartographie des cosystmes. Trois premiers
exemples de cette dmarche sont prsents.
1. Les limites thermiques de la zone borale, carac-
trise par le climat forestier des tagas. Dans tout le
nord-ouest de lEurope (Finlande et Russie), cette
formation est associe une fourchette de tempratu-
res de 5 C environ ; mais les tempratures aux limites
de cette zone et de ses divisions latitudinales diminuent
douest en est en raison de la continentalit croissante.
Ces limites sont en revanche en bonne corrlation avec
la dure de la priode vgtative au seuil de 10 C.
2. La surface de la plage xrothermique sur les
diagrammes mditerranens. Cette surface est une fonc-
tion dcroissante de laltitude et de la latitude et sa
mesure peut servir caractriser les ceintures mditer-
ranennes et leurs subdivisions. Elle peut intgrer aussi
en un indicateur unique les variations de temprature
et de prcipitations occasionnes par les changements
climatiques.
3. La translation gographique des grands cosyst-
mes, dans le cadre des diffrents scnarios prdictifs
des changements climatiques. Ces dplacements pos-
sibles peuvent tre estims sous rserve dun temps
certainement long de mise en quilibre et sous rserve
deffets cophysiologiques complexes et encore contro-
verss. Dans une premire approche, le rchauffement,
suppos agir seul, provoquerait une translation des
formations vers le nord, translation dont lamplitude
probable serait dune ceinture de vgtation mais pour-
rait aller jusqu celle dune zone. Lutilisation conjointe
de la carte cologique et des diagrammes mensuels
pourrait permettre de tenir compte des changements
thermiques et pluviomtriques simultans et de leur
modulation saisonnire.
1. Introduction
The current increasing extent of the problems bound to
the environment and to its always faster transformations
would justify a more important effort to be made so as to
develop an ecological mapping, the principles of which
have been drawn up during the last twenty years and are
likely to lead to applications concerning current problems.
The advantages of cartography broadly speaking in com-
parison with other methods of information processing
(rapidity of perception, superposition and confrontation of
the topics, objectivity) as well as the numerous applica-
tions of ecological mapping in particular have been pre-
viously developed [1, 2].
For example, the possible consequences, on the natural
or modied environments, of a potential climate change
during the next decades might be better assessed if one
could compare the climatic scenarios on a regional scale
with a mapped representation of the main plant ecosys-
tems and of the factors which govern their spatial distribu-
tion; it might be then possible to carry out simulations of
the ecological impacts according to the assumed climate
changes. Questions arise particularly for the interpretation
of the various climatic scenarios now available on the
scale of the European continent [3].
2. Revisiting the methodology
of ecological mapping
Here we shall only deal with maps of wide geographical
areas, on a continental or subcontinental scale, and more
exactly with the attempted ecological mapping of Europe.
For that reason, the anthropogenic inuences on a regional
or local scale (agro-pastoral activities, forester practices,
urbanpollutions, impact of transportation) are not involved
or are far behind the great climatic factors.
These maps may be regrouped into the following two
great families.
Bioclimatic maps. They use data supposed to be
biologically signicant, whether raw (isotherms, isohyets,
etc.) or computed (evapotranspiration, etc.) or even syn-
thetic indices amalgamating some of these data. Among
the oldest published maps, one may mention those by
Cur [4], Emberger et al. [5], and more recently those by
Tuhkanen [6] or Rivas-Martinez [7]. None of them use the
distribution of the vegetation; surprisingly enough, they
often disregard relief and even high mountain chains.
Vegetation maps (for the whole continent [8, 9, 10]).
Reciprocally, they generally represent only the plant cover,
usually only the forest climax. Ecological data are pushed
back into the text accompanying the map (through cap-
tions or notices, but usually a succinct form, reduced to
the essential climatic parameters of the main identied
vegetation units) or else through auxiliary insets on a
smaller scale, set in the margin of the main map.
It is quite surprising that none of these maps of the
European continent attempts to integrate climatic and
botanical data, i.e. to carry out a genuine ecological map.
On the contrary, it has generally been set as an a priori
principle that a climatic map must include only meteoro-
logical data and that, reciprocally, a vegetation map has to
be expurgated of all elements not belonging to this vegeta-
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tion. Here methodological strictness actually hides a com-
fortable withdrawal and denial of the notion of interdisci-
plinarity.
We want to react to this situation by introducing here a
quite different step based on proposals presented on sev-
eral occasions [1, 2]. Our purpose is to use the connec-
tions between vegetation and environmental factors, not
only in order to explain the rst ones by the second ones,
but also in order to use the vegetation as an indicator of
ecological conditions and as a foundation for improve-
ment of their mapping. With this aim in view, it is advis-
able:
on the one hand, to stop looking at the vegetation as an
end in itself and a nal document, but, on the contrary, to
change it into a starting point and an analysis tool emerg-
ing into a second generation of maps: ecological maps;
on the other hand, to replace the common reference to
ecological data supposedly bound to vegetation units by a
real statistical study of these connections, which would
encompass the whole spatial extent of each vegetation
unit, as shown below (The thermic limits of the boreal
zone) for the boreal zone.
3. A new vegetation map of Europe
At rst, so as to avoid frequent ambiguities, it is indis-
pensable, to emphasize what we mean by vegetation
mapping in its current form. This mapping no longer looks
at the distribution of plant species (or taxa in general) but
takes into account the integrated systems which are the
biocoenoses and the ecosystems. One knows that their
connections with the environment are more accurate and
their indicative values more signicant than those of plant
species.
Experience has fully demonstrated that in practice the
part played by plants, i.e. the phytocoenose (and what is
more the reduction of phytocoenose to the vascular plants
alone) gives an approached but rather satisfactory picture
of the studied ecosystem. Vascular plants and especially
ligneous ones represent by far the essential part of the
biomass in the ecosystem. They work towards creating or
shaping in a determining manner the environment which
houses the other biological components. Owing to their
height and to their longevity, ligneous species are indica-
tors and integrators of the environmental conditions in
time as in space. This explains that the representation of
the forest climaxes has been placed to the fore from the
outset in mapping of vegetation. So, the denition, the
spatial delimitation and afterwards the mapping of the
ecosystems becomes nearly identied with those of the
vascular plants and of the communities they form. But, it is
important not to mistake the considered ecosystems with
the fundamentally physionomical and biologically insuf-
cient typology generally used by the remote sensing
(CORINE Land Cover), nor with the solely oristically
based hierarchization used in phytosociology.
There are not even any recently drawn up and ecologi-
cally workable maps available. The rst vegetation maps
of the European continent were too imprecise and repre-
sented only a compilation of other maps, the principles
and the typology of which were not always compatible.
But, during recent decades, maps have been published
which go as far as representing the dynamics of the veg-
etation (France, 1/200 000, Spain, 1/400 000) as well as
syntheses drawn up within the national territory of the
greatest number of western and central Europe states (list
by scales and assemblage plan: ([11], p. 306) and ([12], p.
61). On these bases, four vegetation maps of wider geo-
graphical areas have been published: Danubian Europe
[13], states members of the Council of Europe [14, 15],
eastern Europe [16, 17], Europe and Turkey [10].
A new synthesis, drawn up from all the documents
currently available, has allowed us to elaborate a general
map on a 1/7 500 000 scale, a map which can only be
reproduced here about two times reduced (gure 1 ). It
takes up and develops two former attempts [11, 12]; the
unit number has been raised from 38 to 66 and a homo-
geneous nomenclature paired with a numerical code is
proposed. The reduced scale has required many generali-
zations and compromises, more particularly in the intri-
cate regions of Rhenish and Danubian Europe or in the
Mediterranean area; in this area, even great formations
have been distinguished only in the Spanish peninsula. Of
course, this reduced map may not be directly used as a
work document and is given here only to illustrate the
units list. The drawing up of a more detailed version on a
1/5 000 000 scale is now in progress. This new version
will be available, in autumn, on the Web site of our
laboratory (http://www.ujf-grenoble.fr/BIO) but subject to
the quotation of the source. An A1 format printed copy
will be sent on request at cost price.
The units are theoretically named from the climacic
forest types. Colours have been chosen according to Gaus-
sen ecological code, well enough known not to require
any further explanations [18].
The vegetation of Europe is usually rst divided into
three great latitudinal zones or biomes: viz. from north to
south, the boreal zone, the nemoral zone and the Medi-
terranean zone. They are not at all equivalent: the nemoral
zone is by far the widest and the most complex. And its
unity is often articially maintained by referring to a unit of
deciduous forests (expressed in phytosociology in a single
class of Querco-Fagetea) thus hiding the geobotanical
diversity.
In our map, the European vegetation (gure 2) is divided
into eight large regions called domains. Two of them
correspond to the boreal (to which the sub-Arctic is
attached) and Mediterranean zones. But the old nemoral
zone has been divided into the following domains.
In the south, a Thermonemoral Domain has been
isolated [19], which includes the great mountain ranges of
mid Europe (Pyrenees, Alps, Carpathian Mountains, etc.)
within which an Alpine Domain would be dened, and to
which in its turn the Caucasus has been added here.
In the north, the nemoral zone (stricto sensu) has been
divided into three domains, relaying from the west to the
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
Figure 1. A new vegetation map of Europe.
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
.
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
Table I. List of the vegetation units.
1. Boreal
11 Sub-Arctic (toundra)
12 Orocaledonian (mountain tundra)
13 Forest tundra (taiga)
14 Northern taiga
142 Mountain facies on Ural western slopes
15 Central taigas
151 Preuralian facies with Siberian species
152 Mountain facies on Ural western slopes
16 Southern taigas
161 Preuralian facies with Siberian species
162 Mountain facies on Ural western slopes
17 Comprehensive Atlantic taiga on the western slope of the Scandes
2. Atlantic
21 Boreo-Atlantic
211 Scottish mountain heaths
212 Blanket bogs
22 Anglo-Irish acidiphilous oak woodlands
23 Anglo-Irish neutrophilous oakash woodlands
24 Channel oakbeechwoods
241 Hyperhygrophilous armorican sector
251 Continental acidiphilous oak woodlands, North Sea sector
252 Continental acidiphilous oak woodlands, Loire sector
253 Continental acidiphilous oak woodlands, Submontainous Limousin sector
254 Continental acidiphilous oak woodlands, Landes sector
26 Neutrophilous mixed oak woodlands
261 Thermophilous aquitanian sector
27 French Central Massif mountainous beechpine complex
28 Ibero-Atlantic acidiphilous oak woodlands
29 Cantabric mountains range beechwoods
3. Centro-European nemoral
31 Scandinavian boreo-nemoral
32 Baltic beechwoods
33 Subcontinental oak-pine woods with xeric enclaves
34 Subatlantic and submountainous beechwoods
35 Hercynian mountainous beechrsprucewoods
4. Sarmatic
41 Boreo-nemoral mixed forest (hemitaiga)
411 Preuralian sector
412 Mountainous facies in southern Ural
42 Eastern Europe nemoral (without beech)
421 Polish-Bielorussian sector (with hornbeam)
422 Pripet marshes subsector
423 Forest enclaves in the following unit (43)
43 Tree steppe
5. Steppic
51 Transitional forbes steppes
52 Stipa central steppes
53 Stipa-Artemisia southern steppes
54 Caspian hemideserts
6. Thermonemoral
61 Po Basin riparian and collinean vegetation complex
62 Submountainous oakhorbeam woodlands
63 Transylvanian thermophilous oak woodlands, tree steppes and riparian formations complex
64 Thermophilous Ungarian oak woodlands
65 Mountainous eastern beechwoods
66 Xeric Pontus hornbeam woodlands
67 Thermohygrophilous Colchidian vegetation
7. Mediterranean (I = Iberic peninsula only)
71 Monti- and alti-Mediterranean vegetation belts
72 Supra-Mediterrannean vegetation belt
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
721 Silicicolous (I)
722 Calcicolous (I)
73 Medio-Mediterranean vegetation belt
731 Silicicolous, with round-leaved oak (I)
732 Calcicolous, with round-leaved oak (I)
733 With cork oak
734 Semi-arid (I)
74 Thermo-Mediterranean vegetation belt
741 Baetic arid sector (I)
8. Alpine
81 Lower and mid mountain: collinean and montane vegetation belts
82 High mountain: subalpine, alpine and nival vegetation belts
83 Inner axis (Alps and Caucasus)
9. Azonal
91 Coastal dunes
92 Polders and river deltas
east according to the increasing continentality: Atlantic,
centro-European (sub-Atlantic) and sarmatic (continental).
The last one is obliquely cut, in its south-eastern part, by a
Steppic Domain.
Some domains are in their turn subdivided into latitudi-
nal belts (except where mountain ranges disturb this state):
for example, the Boreal Domain into three taigas belts
(gures 3 and 4). In the Alpine Domain, these vegetation
belts are identical to the altitudinal vegetation belts.
Divisions of third order, the so-called sectors, have been
introduced in our vegetation map in special cases.
3.1. Numerical list of vegetation units
The vegetation units are generally named according to
their climactic forest types. An overall description, includ-
ing treeless stages, has been given in Ozenda [12] (table I).
4. From the vegetation map
to an ecological map
Strictly speaking, at that time the technical aspects of the
mapping will not be expounded here. Their development
S
t
e
p
p
e
s


z
o
n
e
Thermonemoral
domain
Atlantic
domain
Boreal zone
U
r
a
l
Sarmatic domain
Central
European
domain
Mediterranean
zone
C
a
u
c
a
s
u
s
0
500 km
Figure 2. Main geobotanical divisions of Europe (according to Ozenda, 1994).
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
depends on the studied problemand on the purpose of the
ecological map: nature protection, deterioration of the
environment, country planning [2]. This paper only deals
with the ecological analysis of plant formations.
A rst step consists in assigning to each vegetation unit
the most signicant ecological parameters, whichamounts
to transform the plant vegetation map into an ecosystem
map. It is what will be proposed further on concerning the
temperature limits of the boreal taga (see below The
thermic limits of the boreal zone).
Indeed, ecological literature works are full of attempts at
drawing up relationships between species distribution lim-
its and climatic limits (isotherms, isohyets, or composed
indices); but it always concerns comparisons between
maps on a small scale. The limits of plant formations, the
indicative value of which is higher than that of the plant
species, are often discussed but here again according to
documents showing insufficient accuracy. Now the cur-
rent progress of vegetation mapping allows more elabo-
rate comparisons. For example, the eastern limit of the
Atlantic vegetation on the one hand, of the medio-
European beechwoods on the other (as they are stated
precisely by recent maps and their accompanying com-
mentaries) almost coincide, between the 46N and 55N
latitudes, with the continentality lines (range between the
averaged January and July temperatures) of 12 and 24 C,
respectively. These are the climatic, forest and ecological
limits we have taken into account so as to divide the three
domains of the nemoral zone.
Among the numerous climatic data sources, we recom-
mend, just like before, using the tool of seasonal ombro-
thermic diagrams. The principle has been laid down by
Gaussen [20] then developed by Walter and thousands of
diagrams have been gathered in a monumental book, the
Klimadiagramme Weltatlas of Walter and Lieth [21]. This
eminent information, to be updated urgently, still seems
insufficiently exploited in ecological research.
This type of representation consists in comparing month
after month the ratio between precipitations and tempera-
tures. With this aim, the curve of the monthly averaged
temperature and the curve of the monthly sum of rainfall
are drawn up on the same graph, with such a scale that
1 C corresponds to 2 mm of rain. One agrees to call dry
periods those during which the rainfall curve is under the
temperature curve; it most often concerns a dry summer
season in southern Europe which is usually materialized
on the graph with a dotted area. On the contrary, moist
seasons, for which P > 2T, are represented with vertical
hachures; so as to avoid an excessive size of the diagram
when the climate is very wet, the height of the precipita-
tions greater than 100 mm per month are represented in
black. The rules of use of these diagrams are described in
several languages in the introduction of the Walter and
Lieth atlas. A simplied presentation is given in Ozenda
[2, 12].
An advantage of these diagrams lies in the fact that the
use of monthly means is more objective that composite
indices in which the choice of variables and coefficients
may be empirical or arbitrary. However, the objection may
be raised that the climatic data, generally covering 30
years, approximately refer to the middle of the XXth cen-
tury; but, the variations during the last decades (a rather
general increase in temperature of about 0.2 C) do not
invalidate the comparisons between stations the means of
which differ from several degrees.
We shall hereafter present two cases of use of such data
at rst for the boreal forest (section 4.1) and then for the
Mediterranean region (section 4.2), then an example of
application of the ecological mapping to the possible
biogeographical consequences of the climate change (sec-
tion 4.3).
4.1. The thermic limits of the boreal zone
In north-eastern Europe, the distribution of the yearly
averaged temperatures according to latitude is often prac-
tically linear in the absence of relief; vegetation maps
show that the zone of the boreal forests, the so-called
taiga, approximately lls a belt nine degrees wide in
latitude corresponding to an approximative 5 C range
(gures 3 and 4).
In Finland, around the 28th meridian, this temperature
range corresponds to the interval +4 to 1 C. The same
evaluations, deduced from the most recently published
vegetation maps of Russia [16, 17] show that the boreal
taiga zone, the composition and the structure of which are
similar and roughly constant from the west to the east, lls
a belt the geographical limits of which are practically
constant in latitude. The temperature bracket is always
A
B
N
C
S
70 N
65 N
60 N
-1 0C 1 2 3 4
Figure 3. Geographical and thermal limits of the boreal taiga in
Finland.
AB, yearly averaged temperatures according to latitude (each point
corresponds to the average of ve meteorological stations within one
degree of latitude; N, C, S, latitudinal distribution of the three taigas
belts, north, centre and south.
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
about 5 C but its limits are different and increasingly cold
eastwards: +3.3 to 1.6 C along the 40th meridian,
+2.2 to 2.3 C along the 50th meridian. In fact, if the
geographical limits of these three taiga belts are roughly
constant in latitude, they forman angle of more than half a
radian with the isotherms which are very oblique owing to
the increasing continentality from west to east (at equal
latitude, the yearly averaged temperature is lower towards
the east).
It is well known that under cold climates, vegetation is
much less correlated to the yearly means than to the
summer temperatures. For this purpose, Dahl [22], for
northern Europe, uses the average of the temperatures of
the three warmest months (tritherm), Tuhkanen [6] the
sumof the averages of these three months. The time during
which the averaged daily temperature is higher than a
stated threshold might be another thermic indicator of this
summer period. This threshold is often set to 5 C; we
propose to use the threshold of 10 C which presents the
advantage that the duration of the corresponding period
can be directly read on the Walter and Lieth diagrams. At
equal yearly averaged temperature, this duration increases
from west to east (gure 4); but one may note that the
equal duration lines of the period are then roughly par-
allel to the geographical limits of the taiga belts and
become an ecological characteristic of the latter.
Figure 4. Geographical situation of the three boreal taiga belts in northern Russia.
Thick lines represent rough limits of the three taiga belts (N, Cand S) and of the so-called hemitaiga mixed forest, H. The ne oblique lines represent
the yearly isotherms; dashes, the equal duration lines of the summer period (number of days on which the daily thermal average is above 10 C).
The points represent the meteorological stations whose climatic data have been used. The circles (at the top, on the right) represent the Ural ridge.
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
4.2. The Mediterranean xerothermic area
The careful scrutiny of the existing vegetation maps
already allows the limits of the domain or of the vegetation
belts to be connected with raw climatic data more pre-
cisely thanpreviously. So, since the thermo-Mediterranean
belt has been mapped in detail, it appears that its northern
limit almost exactly coincides with the yearly 15 C iso-
therm in the western Mediterranean and 16 C in the
eastern basin [23].
But, a chief ecological factor of the Mediterranean
climate lies in the existence of a more or less intensive and
more or less long summer dryness which is remarkably
expressed on the ombrothermic diagrams by an area mate-
rialized by the passing of the temperature curve over the
rainfall curve (with the classical scale conventions of these
diagrams, as mentioned above). The surface of this area
may be used as a simple indicator integrating the thermic
and the hydric aspects: so, in a Tunisian mountainous
sector, the Djebel Mansour, it has been proved [24] that it
is a quasi linear decreasing function of altitude (conse-
quences of orogenic rains) and of the latitude (change
from the sub-Saharian regime to the steppic regime north-
wards).
The European state where the Mediterranean region
presents the maximum extent and diversity is obviously
Spain. The vegetation map on a 1/400 000 scale in 29
sheets and a compilation of bioclimatic data [25] cover
the whole country. From this large documentation, dia-
grams typical for a part of the vegetation belts have been
selected (gure 5) : the surface of the summer area appears
to be a good indicator at once discriminating and easy to
use. Measurements carried out on the diagrams from
about 30 stations of Mediterranean Spain give table II.
If the validity of this method is conrmed, it will have to
be handledwithcare however inthe comparisons between
distant regions. Already on the northern borders of the
Mediterranean domain, the dryness area is smaller and
disappears as soon as one leaves the Mediterranean belt
[3, 26] and, in the steppic domain, it appears in the units
53 and 54 of our map only in countries in which the
rainfall becomes lower than 350 mm per year [16, 17].
4.3. The geographical shifts of the ecosystems
as a consequence of the climate change
It is now generally admitted that the deep and long-
lasting modications of the atmospheric chemical compo-
sition, induced by the continually increasing emissions of
the so-called greenhouse gases and aerosols, change the
radiative budget of the terrestrial systemand foreshadowa
notable warming of the climate from the outset of the next
decades [27]. It is difficult or untimely to state that the
averaged increase of 0.5 to 0.7 C observed on the world
surface during the last 100 years has been caused by these
emissions but serious presumptions exist [28]. And as the
industrial activities seem unavoidable to amplify, one
considers as possible an increase of 3 1.5 C during the
XXIst century, also involving considerable changes in the
amount and in the regime of precipitations [3, 29].
Many scientic research programmes, both theoretical
and experimental, have been undertaken during the last
decades so as to assess the impacts on the different biota of
a considerable and long-lasting climatic change over a
century. One of the main aims lies in the possible assess-
ment of the geographical shifts of the main plant species
and even of the ecosystems entailed by these future cli-
matic modications. But the study of the ecophysiological
effects (the so-called direct effects) is very complex,
becomes continuously more complicated and may not be
developed here [30]. Besides, it is highly unlikely that
ecosystems should shift as made unities. On the contrary,
it is more likely that plant species will individually react
just like during the holocene recolonization. Neverthe-
less, one always has to keep in mind that some plant
species are more important (the so-called key-species),
particularly the main tree species which form almost all of
the biomass and in which the greatest part of the energy
exchanges take place. One should also mention the tree
species which are now associated in a given forest group,
having close ecological requirements, and which conse-
quently will probably shift together. Here, we shall only
deal with the applications of the ecological mapping and
more precisely of the possible shifts of the ecosystems as
consequences of the climate modications, just as hap-
pened on a much greater scale during the Pleistocene
glaciations and during the Late Glacial and the Holocene.
These shifts, whichdonot exclude the physiological effects
but add to them, provide some problems as for:
Figure 5. Characteristic ombrothermic diagrams of Mediterranean
belts and sub-belts in Spain: progressive reduction of the summer
xerothermic area according to the increasing altitude (according to
Rivas-Martinez, 1987, modied).
A, thermo-Mediterranean belt (Sevilla, 10 m); B, meso-Mediterranean
belt (Madrid, 660 m); C, supra-Mediterranean belt (Avila, 1 130 m);
D, oro-Mediterranean belt (Punta Navacerrada, 1 860 m).
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P. Ozenda, J .L. Borel / C.R. Acad. Sci. Paris, Sciences de la vie / Life Sciences 323 (2000) 983994
their direction;
their amplitude;
their chronology: latency time, then duration;
the process of their development;
last and above all, their consequences for humanity.
A rst step may be limited to the thermic effect, suppos-
edly acting alone. Warming would involve a translation of
the vegetation zones northwards. Simulations have been
tempted for North America and Europe, but some have
been based on the ancient classication of Holdridge the
foundations of which are too theoretical and give only
very approximative and small scale gures [31]. The use of
mid scale ecological maps carried out with recent vegeta-
tion maps, as recommended here, constitutes a exible
predictive tool, adapted to the various available climatic
scenarios.
The amplitude of the shift must be evaluated by means
of temperature brackets really allocated to the plant for-
mations. If, for example, the expected warming amounts
to 3.5 C and the width of the zone of the boreal taiga is
4.5 C, this latter would almost completely give way to the
zone of the nemoral forests or at the very least to the
so-called hemitaiga mixed forests.
Already now, the warming of approximately 0.7 C
recorded during the XXth century might, once the equilib-
rium climatevegetation is reached, correspond to a shift
northwards of about 120 km. The rst signs of evidence
are already discernible in the herbaceous vegetation [32]
as well as in the bird populations [33]. In mountainous
areas, the limits of the vegetation belts might raise upwards
[3, 29], as during the Holocene.
But these vegetation shifts suppose a rather long balanc-
ing time: inertia of oceans in contrast to the atmosphere,
inertia of the ecosystems and of the soils; one knows that
maturation of the forest ecosystems may require several
centuries [34].
The anthropogenic increase of the greenhouse effect
will involve immediate modications of temperatures and
of precipitations and many scenarios have been worked
out, particularly for northern Europe [35]. Ecological maps
may be used for the estimation of these two climatic
factors and their common action: so, the quantitative
assessment of the Mediterranean dryness area and of its
expected variations may be a more handy tool that the
habitually used formula.
On the other hand, climatic scenarios show that the
changes would differ greatly according to season: in this
view, the use of the ombrothermic diagrams taking into
account the monthly values is highly commendable.
Acknowledgements. We wishtowarmly thank G. Borel
for her infectious enthusiasm and her ceaseless avail-
ability during the ticklish, tiring and repetitive work of
numerical acquisition and processing of the different
versions of this phytoecological map. The gures have
been carefully drawn up by J.-P. Guichard. A.-L. Tissut
greatly improved the English. We are especially
indebted to the two anonymous referees who provided
many constructive comments on the manuscript which
were greatly appreciated by the authors.
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