The Mediterranean maquis
Italian habitats
Italian Ministry of the Environment and Territory Protection / Ministero dell’Ambiente e della Tutela del Territorio
Friuli Museum of Natural History / Museo Friulano di Storia Naturale · Comune di Udine
Scientific coordinators
Alessandro Minelli · Sandro Ruffo · Fabio Stoch
Editorial commitee
Aldo Cosentino · Alessandro La Posta · Carlo Morandini · Giuseppe Muscio
"The Mediterranean maquis · Evergreen coastal formations"
edited by Alessandro Minelli
Giuseppe Carpaneto · Gaudenzio Paola · Simonetta Peccenini · Margherita Solari
In collaboration with
Luca Lapini · Mirca Zocchi
English translation
Gabriel Walton
Roberto Zanella
Graphic design
Furio Colman
The Mediterranean Maquis
Evergreen coastal formations
Archive INFS (Valter Trocchi) 139 · Archive MFSN (Ettore Tomasi) 36/2, 37, 39, 48/2, 50, 60 ·
Mauro Arzillo 128, 129 · Paolo Audisio 8 · Pietro Baccino 12, 15, 20/1, 22, 24, 32, 35/1, 36/1, 38/1, 38/2,
41/1, 41/2, 42, 44/1, 44/2, 47/2, 51, 64/1, 64/2, 64/3, 69, 78, 132/1, 132/2, 133, 150, 151 ·
Enrico Benussi 116/1 · Eugenio Busetto 18/1, 26, 34 · Giuseppe Carpaneto 11, 33/2, 35/2, 44/3, 47/1, 70,
71, 80, 87, 88/1, 88/2, 89/1, 102, 107, 109, 110/1, 110/2, 111, 112/1, 112/2 · Ulderica Da Pozzo 67, 134 ·
Vitantonio Dall’Orto 74 · Dario Ersetti 19/3, 33/1 · Maurizio Fabbri 93, 95/1 · Luca Facchinelli 10 ·
Gabriele Fiumi 90/1, 90/2, 90/3, 94, 95/3, 96, 97, 98 · Giovanni Gobbi 84, 89/2, 99, 101, 104, 114/1, 126 ·
Gianluca Governatori 18/2, 18/4, 19/2, 82, 148 · Luca Lapini 81, 110/3, 122, 140 ·
Daniele Macale 95/2, 103 · Paolo Maltzeff 90/4 · Maurizio Rizzotto 77 · Ugo Mellone 18/3, 83, 130 ·
Giuseppe Muscio 28 · Michele Panuccio 86, 108, 120 · Gaudenzio Paola 25, 43, 48/1, 54, 56, 62, 142,
144 · Roberto Parodi 40, 113, 114/2, 115/1, 115/2, 116/2, 116/3, 117, 118, 119, 135 ·
Silvia Sebasti 123, 137, 138 · Paola Sergo 19/1 · Pino Sfregola 20/2 · Margherita Solari 147 ·
Mido Traverso 52/1, 52/2, 53/1, 53/2 · Roberto Zucchini 57
© 2003 Museo Friulano di Storia Naturale, Udine, Italy
All rights reserved.
No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or
by any means, without the prior permission in writing of the publishers.
ISBN 88 88192 11 5
Cover photo: low maquis with thyme and ever-lasting flowers (photo by Margherita Solari)
M I N I S T E R O D E L L’ A M B I E N T E E D E L L A T U T E L A D E L T E R R I T O R I O
M U S E O F R I U L A N O D I S T O R I A N AT U R A L E · C O M U N E D I U D I N E
Of the various environmental aspects which may be found in Italian
territory, many are familiar to the traveller's eye, and are commonly
accepted as an expression of naturalness. They are in fact the
result of centuries-long interactions with man, who favoured the
development of secondary formations, variegated and quite often
enriched in several types of flora and fauna.
These interactions - the history and above all the initial conditions
of which are not always easy to reconstruct - have certainly not
spared the Mediterranean maquis, which originally grew along
most of the coastline of the Italian peninsula and its islands and, in
the southernmost regions, extended to the hills inland.
Attentive naturalists will be able to recognize, in the diversity of
maquis formations, those few but precious and beautiful remnants
which are still in a more or less well-preserved state of naturalness.
This volume of the "Italian Habitats" series covers a detailed
itinerary through all the various types of primary and secondary
maquis, reconstructing its ancient dynamics and presenting its
most significant aspects from the botanical and zoological
When attempting to document the historical processes undergone
by the animal and plant populations of the maquis environment so characteristic in a country like Italy, so essentially part of the
Mediterranean world and its life-style - the role played by Natural
History museums is fundamental, since these institutions preserve
and house irreplaceable information on all types of fauna and flora
and their distribution on the territory.
With this series of "Italian Habitats", therefore, the Friuli Museum
of Natural History is proud to contribute to the further knowledge
of these important and indeed unique parts of the environmental
mosaic of Italy.
Carlo Morandini
Director, Friuli Museum of Natural History
Italian habitats
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Giuseppe Carpaneto · Gaudenzio Paola · Simonetta Peccenini
Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Gaudenzio Paola · Simonetta Peccenini
Caves and
Springs and
of the Po
Sand dunes
and beaches
Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Giuseppe Carpaneto
Preservation and management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Giuseppe Carpaneto · Gaudenzio Paola · Simonetta Peccenini
Suggestions for teaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Margherita Solari
Sea cliffs and
coastal lakes
Realms of
snow and ice
Select bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
ponds and
Rocky slopes
and screes
forests of the
“He took to the hills”. How many times have we seen these words in the
adventure stories we read in our youth! Robbers, thieves and bandits, in all
societies, always took to the hills to avoid the legal consequences of their illegal actions, and were often able to hide for long periods of time. In the
Mediterranean context, “taking to the hills” means plunging into the maquis,
that sea of green, which closes above our heads and hides us completely from
the external world. Particularly during the foreign colonization of Southern Italy
and in the confused years following the unification of Italy, brigands and bandits took refuge here and, during the Second World War, so did partisans and
members of the resistance.
But what exactly is the maquis? In the general acceptance of this French
word, which cannot really be translated into English, the maquis is a vegetal
formation with prevailing large shrubs or small to medium-sized trees, often
spaced so far apart that light can penetrate well. This allows the development
of a thick, intricate undergrowth formed of shrubs and creepers. These formations are found where the original broad-leaved forest has been felled, and
they generally evolve in order to reconstitute that forest by means of a long
process of self-restoration, called succession. The maquis, in its widest sense,
may be composed of deciduous broad-leaved trees which tend towards the
reconstruction of the original deciduous oak forest, or evergreen shrubs which
tend to recompose the Mediterranean forest, dominated by evergreens like
holm oak and cork oak.
In a strict sense, the word maquis is mainly applied to its Mediterranean version - that formation resulting from the destruction of the evergreen forest
(mainly holm oak, as regards Italy), which grows prevalently along the coastline. As regards its extent, evergreen maquis is currently the main vegetal formation in coastal and subcoastal areas of the Mediterranean, and it thus covers much of the Italian peninsula and its islands. In addition, from the coast,
the maquis often penetrates inland, developing on the warmest, often southfacing flanks of the anti-Appennine and pre-Appennine hills. Even in the heart
of the Appennines and in some Prealpine districts, so-called “xerothermic”
areas may be found, where favourable local climatic conditions allow the
Maquis with rockrose and oleander along littoral of Focene (Latium). Oleander (Nerium oleander) is a
common riverbank species in southern Europe; elsewhere, it is planted by man
conservation of biocenoses associated with hot, arid climates. These biocenoses are relicts from interglacial periods, during which mountain vegetation was forced to retreat due to climatic warming, thus allowing the advance
of biological communities generally found near the coast. This is why, as well
as evergreen plants, xerothermic areas may host a fauna usually associated
with coasts and with limited capacities for dispersion, like some gastropod
molluscs and wingless insects.
The development of the Mediterranean maquis is a very ancient process,
which began in prehistorical times and which reflects the growing impact of
human activities on the environment. At least in Italian territory, evergreen
maquis is the result of man’s direct or indirect intervention on the environment
of coastal belts - burning scrub and woodland, felling trees, and grazing livestock. Initially, tens of thousands of years ago, during the hunter-gatherer
phase of human development, fires were set in vegetation to frighten wild animals and drive them towards points at which they could be ambushed and
trapped. Later, during agricultural and livestock farming phases, the purpose
of setting fire to the surrounding vegetation was to create space for settlements and agricultural exploitation, or to stimulate the growth of forage for
livestock. Thus, with the passing of centuries, the evergreen Mediterranean
maquis became the prevailing landscape of the Italian coastline.
Mediterranean maquis with Euphorbia dendroides on island of Caprara (Tremiti, Apulia)
Pines in Mediterranean maquis along Tuscany coast
■ Introduction
What is the Mediterranean maquis, exactly? Seen from a distance, the maquis
is a sea of changing tones of green, covering hilly slopes near the sea, glowing in spring with the yellow flowers of broom, the white or pink of rockrose,
and the blue of rosemary (Rosmarinus officinalis) and germander (Teucrium
Only if we approach closely can we see the dense intertwinings of the bushes
which compose it: holm oak (Quercus ilex), strawberry tree (Arbutus unedo),
buckthorn (Rhamnus alaternus), broad-leaved phillyrea (Phillyrea latifolia),
laurustine (Viburnum tinus), a representative of the honeysuckle family with
pretty corymbs of white flowers and fruit of a beautiful metallic blue, tree heath
(Erica arborea) and other types of heath (e.g., Erica scoparia), prickly juniper
(Juniperus oxycedrus) and cork oak (Quercus suber), with its characteristic
thick bark which protects it from bad weather and from fire.
The maquis sometimes contains a series of lower shrubs like butcher’s broom
(Ruscus aculeatus) with evergreen leaves broadened to form “cladodes” holding small star-shaped flowers and then its typical red berries; ivy (Hedera
helix), or plaited lianas like asparagus (Asparagus acutifolius), madderwort
(Rubia peregrina), ruby-coloured with evergreen leaves, verticillate, with small
hooked hairs on the edges; Mediterranean honeysuckle (Lonicera implexa),
with highly perfumed flowers and connate leaves, i.e., welded together at the
base round the stem; sarsaparilla (Smilax aspera) and rose (Rosa sempervirens). The last two make the maquis almost impenetrable with their thorns as clearly suggested by the common Italian name for Smilax: “strappabraghe”
(“tears your trousers”)!
In cooler locations, we find laurel (Laurus nobilis), flowering ash (Fraxinus ornus)
with pennate deciduous leaves and highly scented white blossom, terebinth
(Pistacia terebinthus), and in warmer spots many-flowered heath (Erica multiflora), tree euphorbia (Euphorbia dendroides), Phoenician juniper (Juniperus
phoenicea), with scale-shaped leaves, another form of juniper (Juniperus
oxycedrus spp. macrocarpa) with needle-shaped leaves and large brown
berries, colonizing sandy dunes, myrtle (Myrtus communis), wild olive (Olea
Holm oak (Quercus ilex) in flower
europaea var. sylvestis), progenitor and sometimes used to graft cultivated
olives, carob (Ceratonia siliqua), with horny, evergreen leaves, paripennate and
with large brown, sugary pods, thorny oak (Quercus coccifera, Q. calliprinos),
with evergreen leaves with very thorny edges, narrow-leaved phillyrea
(Phillyrea angustifolia), lentisk (Pistacia lentiscus), Jupiter's beard (Anthyllis
barbajovis), a legume with silvery foliage and white flowers, and dwarf palm
(Chamaerops humilis), the only species of palm growing spontaneously in Italy.
Locally, types of common broom may prevail, leguminous with yellow flowers,
like spiny broom (Calicotome spinosa, C. villosa, C. infesta), cytisus (Cytisus
villosus), hairy cytisus (Chamaecytisus hirsutus), European broom (Cytisus
scoparius) and Spanish broom (Spartium junceum). In other places, where
fires are frequent, we find abundant pink rockrose and Cretan rockrose (Cistus
albidus, C. creticus), sage-leaved rockrose (Cistus salvifolius) and Montpellier
rockrose (Cistus monspeliensis).
The Italian “macchia” and the French “maquis” are used throughout the world,
mainly in the scientific literature, to indicate the shrubby, stiff-leaved, evergreen vegetation so typical of the Mediterranean climate. But there are also
many other names: matorral in Spanish-speaking countries (Spain and Chile),
chaparral in California, strandveld and renosterveld (according to dominant
floristic composition) in South Africa, and mallee in Australia.
When increasing summer aridity or heightened pressure of anthropic activities
means that the vegetation becomes scanty and low, without the typical features of maquis, it is called gariga in Italy, garrigue in France, phrygana in
Greece, batha in Israel, jaral in Cile, and coastal sage in California.
Instead, the term fynbos, used in southern Africa, means the Mediterraneantype vegetation growing in those parts: dominated by heath, it is very similar
to the moorland of temperate climates, which develops in soils particuarly
poor in nutrients.
■ The Mediterranean maquis in the world
Just as the term “Alpine” derives from the geographic name for the Alps,
“Mediterranean” refers to the basin of the Mediterranean Sea in a geographic
sense. However, it is also used to indicate the complex of climatic conditions
which occur in this part of the world. This complex may also be observed in
some parts of other continents: in the Americas (California, central Chile),
southern Africa, and south-western and southern Australia.
None of these areas covers an area comparable with that of the Mediterranean
basin, but their vegetation is very similar to the Mediterranean version from the
structural viewpoint and from that of the morphology of the prevailing plant
species. In areas so far removed from each other, floristic diversity is remarkable.
The Mediterranean basin is also an area in which several civilizations followed
each other in ancient times, and man’s relationship with his environment contributed to a marked degree in structuring and consolidating the essential features of the landscape. In spite of its territorial continuity, the Mediterranean
region is in fact highly diversified.
Generally, we can distinguish its northern and southern parts, as aridity
gradually increases from north to south. The western portion is also different
from the eastern one, with rainfall which decreases from west to east. But,
above all, there are great differences in the history of man’s relationship with
the territory, which began in the east about 4,000 years before it did in the
west. Most of the Mediterranean plants of agricultural interest originated in
the Eastern Mediterranean.
Myrtle (Myrtus communis)
■ Ecology of the Mediterranean
The climatic context. Within the
framework of the large-scale climatic zones of our planet, the climate of the Mediterranean basin
represents the transition between
the temperate belt of Europe and
the arid tropical belt of North Africa.
The Mediterranean climate has
autonomous characteristics, which
may be briefly described as follows:
maximum annual precipitation concentrated in cold periods of the year
(autumn, winter); arid conditions
during the warmest months (summer); very variable precipitation
from one year to the next; hot or
very hot summers and cool or cold
winters, but without great differences between day-night temperatures or between the various seasons of continental type, i.e., very
marked; and, lastly, intense solar
radiation, above all in summer.
The size of the Mediterranean basin
and the complex morphology of the
lands which border and sometimes
influence it cause local variations
in the general climate, usually distinguished into four main types,
mainly based on the length of the
arid period.
Thermometric trends during the
course of the year, although kept in
consideration, play a secondary
role. The Mediterranean climate is
thus subdivided into: xero-Mediter-
Areas of world with prevalent
Mediterranean maquis
vegetation (red)
Distribution of olive in
Mediterranean basin. Although
a cultivated species, olive
does approximately indicate
the belt of Mediterranean
ranean (or arid), with no rain for 910 consecutive months; thermoMediterranean (semi-arid), with 7-8
months without rain; mesoMediterranean (sub-humid), with
aridity lasting for 5-6 months, and
sub-Mediterranean (damp), with
“only” 3-4 consecutive months
without rain.
The xero-Mediterranean type of
climate represents the borderline
situation with the desert-type climates of the Sahara; the subMediterranean climate, extensive
in Italy, connects the temperate climates of northern Italy and,
thence, central Europe.
Clearly, therefore, only part of Italy
can be considered strictly Mediterranean. We can draw an ideal line
across the Italian “boot” running
from Liguria (on the Tyrrhenian side)
to the Conero promontory in the
Marches (Adriatic), and thus divide
the country into two climatically different parts: continental Italy, which
includes the Alpine arc and the
whole of the Po Plain, with a temperate climate; and peninsular Italy,
mainly of Mediterranean type.
This approximate subdivision must
be further refined by bearing in
mind Italy’s complex morphology.
First of all, there is the Appennine
chain which, although extending
within the Mediterranean area,
does not present its typical features. The Appennines generally
have a climate tending towards
San Fruttuoso
Island of Ponza
Island of San Pietro
Cilento coast
Approximate subdivision of Italy into climatic
green: temperate-continental; blue: cool sub-Mediterranean; light blue: medium sub-Mediterranean;
yellow: warm sub-Mediterranean; red: meso-Mediterranean
Holm oak
Gaudenzio Paola
Holm oak (Quercus ilex)
Family: Fagaceae.
A medium-sized evergreen tree
(sometimes reaching 20 m) or shrub,
dense, spherical foliage, short trunk,
ascending branches; dark brown bark,
divided into small square plaques.
Simple alternate leaves, very variable
in shape according to their position on
the tree and where it grows, hard,
oval-elliptical in shape (4-8 x 2-4 cm),
shiny above and covered with short
hairs below, margin entire or dentate.
Small, inconspicuous flowers, acorns
2-3 cm long, partly covered by a
cupule with flat, downy scales.
The once extensive holm oak groves,
coppiced, supplied high-quality
firewood and charcoal.The wood is
hard and heavy, suitable for making
objects subjected to great mechanical
stress, e.g., the moving parts of carts.
The bark was once used to tan hides
and the acorns as food for pigs or
even, in times of famine, for humans.
Distribution of holm oak in Italy, approximately
coinciding with meso-Mediterranean
and warm/medium sub-Mediterranean
climatic zones
temperate, at least as far down as Campania. Naturally, altitude plays an
essential role, so that part of the Appennines affected by the sea only rises to
a few hundred metres on the flanks of the central and northern mountains, but
widens gradually and steadily to the south and the islands. The belt with the
most marked and typical Mediterranean features, with summer aridity of 5-6
months (meso-Mediterranean) involves most of Sardinia, the southern coastline of Sicily, Salento, the Lucanian and Apulian coastlines facing the Gulf of
Taranto, and part of the Apulian coast south of the Gargano promontory.
The belt where summer aridity is limited to a period of 3-4 months (subMediterranean) is very wide, and includes the remaining parts of the two
largest islands (excluding their highest reliefs), the Tyrrhenian coast from Calabria northwards to Liguria (excluding those parts of eastern Liguria and northwestern Tuscany where the morphology of the land (with very steep reliefs on
the seaward side and/or conspicuous mountain systems only slightly inland)
reduces the arid period, and the remaining Calabrian and Adriatic coast northwards, more or less as far as the Conero promontory. The internal parts of
peninsular Italy have increasingly less marked Mediterranean features as we
move inland and upwards from the sea.
The arid period steadily becomes shorter until it may fade away completely; at
the same time, winter temperatures fall, so that most of the Appennine chain,
defined as medium and cold sub-Mediterranean, shows more or less marked
features common to continental climates.
There is a small, cold, sub-Mediterranean zone along the Gulf of Trieste, again
associated with the peculiar morphological and pedological characteristics of
the local territory.
The soil. Soil may be defined as that superficial part of the Earth’s crust which
is the result of interactions between physico-chemical alteration of rocks
(weathering) and decomposition of the organic matter which is deposited on
them. Plants send their roots down into the soil, and find the water and minerals necessary to their metabolism in it.
Geologically, Italy is made up of a large variety of rocks, often leading to considerable diversification of soil substrates, even within the same valley or
mountain chain. Two of the main soil parameters which are important for plant
life are the quantity of lime present, and the degree of acidity.
Calcifuge species show signs of distress if they are obliged to live in limestone
soils. Examples are bracken, or brake (Pteridium aquilinum), European broom,
green heath, cork oak, and maritime pine (Pinus pinaster). Instead, calcicole
species prefer soils rich in lime.
This preference is not due to lime in itself, but to the particular conditions of
heating and aridity of the substrate which these warmth- and heat-loving
species find in such soils. The degree of soil acidity is also a factor determining the geographic distribution of many species of plants.
Maquis species which prefer acid substrates include cork oak, strawberry
tree, tree heath, and wild lavender (Lavandula stoechas); those preferring alkaline soils are Spanish broom, many-flowered heath, Aleppo pine, rosemary,
and many types of juniper.
So many calcifuge species are also acid-loving, whereas those preferring alkaline soils often live well on limestone. To summarize, since all plant species
establish their own “personal” relationships with the soil, the maquis in various
parts of Italy has similar structural aspects but diverse floristic composition,
due to the local soils.
Biology and phenology of plants.
The Mediterranean maquis is thus
closely associated with precise local
conditions, mainly as regards climate.
Maquis is limited to those portions of
the globe where annual precipitation is
irregular over the course of the year,
most rainfall being concentrated in late
autumn and winter, and more or less
prolonged aridity in summer.
In areas with temperature trends like
those of the Mediterranean climate,
Wild lavender (Lavandula stoechas)
but where precipitation increases in
quantity and duration over the year,
conditions conducive to tree growth prevail. Here, the vegetation is mainly
composed of true evergreen woods and forest. Instead, in areas where dry
summers are longer, the vegetation becomes impoverished and the arid shrub
typical of sub-desert and desert zones dominates.
Maquis is formed of species adapted mainly to summer drought, thanks to
their leaf structure and morphology. Leaves are those parts of plants which are
most sensitive to variations in the availability of water. Stiff-leaved plants, or
sclerophylles (from the Greek scleros, meaning stiff, rigid), have long-lasting,
hard leaves, thick cuticles, and deep stomata protected by hairs, which limit
transpiration. An excellent example is the leaf of the holm oak. Other species,
like tree euphorbia and spiny broom, adopt a different strategy to cope with
Adaptations to drought by
hard-leaved plants
Comparison between an ordinary leaf and one from a hardleaved plant (oleander)
upper epidermis with
cuticle; in oleander,
epidermis has several
layers and cuticle is very
palisade-type chlorophyll
lacunose chlorophyll tissue
lower epidermis
in oleander, stomas are
located in deep pockets
and protected from
excessive transpiration by
a mesh of hairs
ample development of
cuticular layer on upper
side of epidermal cells
leaf margins turned
leaf blade folds on itself
during periods of drought
stoma embedded with
respect to leaf surface
cr stomatic crypt
st stomatic cells
cs substomatic chamber
ramified multi-celled hair in
shielded hair of olive,
seen from above and in
summer drought, losing their leaves as summer approaches (a phenomenon
called “estivation”, very frequent in hot, arid, tropical and equatorial zones).
As well as summer drought, plants living in the Mediterranean environment
may have to face a second critical period during winter, due to low temperatures.
Maquis species do not generally resist cold well, and can only tolerate short
periods of frost. Snow damages Mediterranean vegetation, both due to the
thermic shock it causes, and mechanically, by breaking branches, especially if
high winds accompany low temperatures.
Perennial species begin their vegetative activity with the first autumn rains and
pass through an intense period of assimilation, extending to December; then a
pause intervenes, due to the cold. Vegetative activity starts up again more
intensely in spring and continues until the beginning of the dry period; summer
is a phase of almost complete repose. In southern Italy and the islands, winter
temperatures are sometimes sufficiently high to allow vegetative activity to
continue without interruption; in these cases, summer drought often arrives
early, sometimes already at the end of May.
The seeds of annual plants germinate in autumn or spring. In most cases, the
dormant period is relatively short and seeds already germinate with the first
autumn rains. In particularly short-lived species, dormancy is extended and
Strawberry tree (Arbutus unedo) with fruit
germination is delayed until February
or March of the year after that in which
the seeds were produced: in this case,
the vegetative period is reduced to
only 2-3 months. In the coolest season, growth and flowering are at their
peak, and fructification and dissemination take place with the arrival of the
dry season. These species cope with
Juniper misshapen by prevailing direction of
summer aridity in the form of seed.
Despite seasonal changes, the maquis
almost always looks the same, because the plants which flower every month
generally have small, relatively inconspicuous flowers. But, especially where
the maquis is less dense and less dominated by holm oak, it presents fruits of
all colours: the various tones of red of sarsaparilla, rose and strawberry tree,
the metallic reflections of laurustine, the bluish-black of myrtle, and the fluffy
white festoons of clematis (Clematis flammula). The maquis takes on a much
more varied aspect when clearings open, and the blooming of the so-called
early Mediterranean microflora may be briefly admired in spring.
The influence of wind must not be underestimated. The shapes into which
trees are modelled by the wind reveals its strength and frequency. Sometimes,
isolated shrubs or entire areas of maquis are prostrate, or grow curved in the
direction of the prevailing wind.
Dynamism, primary maquis and secondary maquis. The various shrubby
associations collectively called Mediterranean maquis are part of that evolution of vegetation dominated by holm oak or undergrowth with wild olive and
carob, or a degraded series due to various kinds of disturbance, mainly
anthropic. In practice, the following evolutionary sequence may be established: naked soil - steppe with grasses - garrigue - low maquis - high maquis
- forest.
Not all the Mediterranean maquis has the same origin and the same history; its
origin may be viewed as primary or secondary, and it is not always easy to distinguish between the two.
Interpreting Mediterranean shrubby formations is quite difficult everywhere,
because they are mostly the result of a lengthy history which includes several
kinds of activity on the part of man: deforestation, grazing, agriculture, and
more or less frequent fires.
Primary maquis is the original type, not the result of the involution of preceding
forest vegetation, and is rarer. It has two aspects: 1, it is found in areas where
holm oak forest is not the present-day climax, e.g., at the geographical and
altitude limits of the area covered by holm oak; 2, the maquis grows in situations which limit its expansion, e.g., steep slopes, high contents of salt in the
soil, or the continual drying capacity of wind, which means that woody plants
cannot grow tall. These cases may be interpreted as permanent stages of
vegetation, or “subclimax”.
Thus, in most cases, the heterogeneous aspects of Mediterranean maquis
present intermediate stages in the course of evolution or degradation of vegetation, and are thus to be considered as aspects of secondary maquis.
The most common causes leading to secondary maquis are anthropic in origin: felling, or coppicing, of trees, fires, and grazing, which often follow the first
two and contribute further to making the original vegetation retreat.
Many holm oak forests have been destroyed in historical times, both to create
pasture for livestock, and to grow crops, mainly vines and olives. Following
the abandonment of cultivation, both vegetation and soil progressively undergo degradation, which may even lead to naked rock being exposed. But
sometimes plants do manage to regrow, in the form of evergreen shrubs, very
Goats grazing on garrigue in Sardinia
Simplified sketch of Mediterranean vegetation.
Man’s intervention to create space for agriculture and grazing causes either a reduction in the
structural complexity of hard-leaved Mediterranean formations or their disappearance. Abandon of
cultivated or grazing land results in the development of vegetation which, if no fires occur, leads to
increasingly complex formations in the course of time, until the reconstitution of maquis or holm oak
similar to those which made up the undergrowth of the primitive forest, but
with new, more sun-loving components, extraneous to the original vegetation.
Sometimes maquis deriving from evergreen forests which have since disappeared is almost entirely composed of the undergrowth of those forests, without tree cover. This occurs when the climate has become more continental
and no longer allows evergreen shrubs to reconstitute themselves, so that
many forms of maquis may be considered as the remains of once flourishing
holm oak groves, or when such groves, periodically and continually exploited
by man by means of coppicing, are transformed into tall, dense thickets
according to coppicing method used and the length of time in which trees
were left to grow freely. In the case of holm oak coppicing, the maquis which
appears afterwards is a transitory stage, necessary to create the conditions
suitable for future growth of holm oak seedlings and shoots.
The most evolved forms of maquis contain heath, strawberry tree, and holm
oak thickets. The least evolved contain rockrose, associated with garrigue.
As these associations are not fully evolved, each of them is affected by soil
characteristics (pH, amount of lime, etc.) and climate (above all aridity) more
than forest formations.
Mosaic of maquis/garrigue along Cilento coast (Campania)
■ The relationship between fire and maquis
When discussing the effects of fire, a distinction must be made between occasional and repeated outbreaks. Occasional fires allow the vegetation to continue its normal evolutionary course towards forested land, and thus the
maquis which establishes itself represents a transitory phase. However, in
most cases, fires are recurring phenomena and may give rise to cycles called
pyrogenic, i.e., caused by fire itself, so that the vegetation which is burnt is the
result of sometimes many preceding fires.
The interval between one fire and another, and its violence, determine both the
type of starting maquis, and how long it takes to regrow.
For example, the region of Liguria - that part of Italy ranging from Tuscany
along the coast northwards, and then bending west towards France - contains extensive woods of Aleppo pine (Pinus halepensis) or umbrella pine,
with a thick shrub layer. The high degree of combustibility of the pines and
the presence of inflammable shrubs mean that flames can reach even the
tops of the trees. Much of the vegetation is destroyed but, although most of
the pines die, the shrubby layer, formed of species with good capacity for
producing shoots which grow from stumps, re-establishes itself quite quickly.
The grassy layer, which was almost absent even before the fire, increases in
the first post-fire stages, thanks to the new space created, but then immediately decreases, as shrubs take over. If the shrubby layer is composed of
species with poor capacity for producing shoots, regrowth will be slower. In
these cases, renewed growth on the part of pines takes on an important role.
Fire enhances the scatter and germination of pine-nuts and many seedlings
develop, because there is no competition from other species. The result is a
thick population of pines alone - easy prey for later fires which, if they occur
before the young trees have had time to reach reproductive maturity, destroy
them completely.
There are also vast extents of maquis composed mainly of strawberry tree,
tree heath and spiny broom. This is often the result of a succession like that
described above for pine woods. In this type of vegetation, fire spreads rapidly and is totally destructive. If species which can produce shoots easily (like
strawberry tree and heath) predominate, then the shrubby layer regrows
quickly and does not allow the entry of sun-loving or grassy species. Sometimes, in the most mesophilous situations, bracken may prevail in the early
post-fire stages, but is later destined to be supplanted by maquis shrubs. After
8-10 years, if no more fires occur, the shrubby layer returns to its previous
state, as regards ground cover and height.
When the maquis is composed of species with poor capacity for producing
shoots, shrubs grow more slowly and a stage dominated by grassy or suffruticous sun-loving species prevails.
Fires are quite frequent in Mediterranean and sub-Mediterranean vegetation,
and so it is rare to find areas which have never been burnt before. The vegetation is thus already the result of perhaps several preceding fires. As fire has
played an important role ever since prehistory in the evolutionary processes
of man in general and of his environment, it would be true to state that the
development of Mediterranean vegetation has taken place in close relationship with fire and has been considerably influenced by it.
The resulting natural selection of hard-leaved evergreen species over time has
led to great uniformity in response to fires, and most of the perennial maquis
species have good shoot-producing capacity. Regrowth is very rapid after
fires, so that these species quickly colonize the terrain, and they stop, or at
least greatly reduce, the possibility that species extraneous to the pre-fire
populations can establish themselves in the area.
One of the consequences is that, after fires, Mediterranean maquis tends to
re-establish itself quickly. This happens even though the soil may contain vital
seeds belonging to other extraneous species. In conclusion, therefore, single
“fire events” do not excessively change the existing vegetation structure and
floristic composition - or, rather, the more or less accentuated variations which
may be caused by the passage of fire are cancelled in a short, sometimes very
short, time. The maquis, particularly if composed of species which produce
shoots easily, like tree heath and strawberry tree, immediately varies its structure considerably, but can grow again in the course of only a few years.
However, there are cases when such changes are very severe, i.e., when the
tree cover is composed of trees containing resin and the undergrowth is dense
and composed of several layers. But even here, at least shrubs tend to reestablish themselves quite rapidly.
The present-day Mediterranean vegetation is influenced not only by man’s
various activities, but also by the frequency of fires. In places where they are
very frequent, the prevailing vegetation is garrigue or poor, sometimes discontinuous meadowland, occasionally thinly covered with pines. When fires occur
every few years, the maquis is mainly composed of spiny broom, heath and
strawberry tree, with or without pine cover.
The tendency of Mediterranean vegetation, in the absence of fires, to regrow
quite quickly is confirmed in areas where agriculture has been greatly reduced
during the last few decades and where, probably thanks to difficult or impossible access by traffic, fires have not occurred for at least twenty years.
before fire
first post-fire phase
second post-fire phase
before fire
after fire
before fire
first post-fire phase
second post-fire phase
before fire
first post-fire phase
second post-fire phase
before fire
after fire
Effect of passage of fire on various types of Mediterranean vegetation. A: pinewood with abundant
undergrowth; B: pinewood with scarce undergrowth; C: maquis; D: hard-leaved evergreen woodland;
E: mixed hard-leaved evergreen woodland and pines
■ The main aspects of maquis
Cork oak (Quercus suber)
One first, physiognomic, distinction to
be made in maquis is its growth
upwards. There is tall maquis, sometimes 4-5 m high, with prevailing holm
oak, strawberry tree, sometimes cork
oak and, on cooler slopes or at higher
altitudes, deciduous oak such as
pubescent oak (Quercus pubescens)
and cerris or turkey oak (Quercus cerris). Instead, in low maquis, plants
rarely exceed 1.5-2 metres in height,
with lentisk, buckthorn, juniper,
phillyrea, rockrose, etc. But what really
differentiates the maquis is its composition as regards flowers. They exhibit
Wild olive (Olea europea var. sylvestris)
considerable variability from place to
place, according to many ecological
factors and, inevitably, man’s intervention. Often the plants are not stable,
but represent stages of degradation or
regeneration, according to changes
occurring in the environment.
Although within the polymorphism of
the maquis, only dynamic types can be
defined, there are more highly evolved
types which grow in cooler spots, like
holm oak maquis (phytosociologically
framed in the order of Quercetalia ilicis
Rosemary (Rosmarinus officinalis)
and in the Quercion ilicis alliance).
Nearer the sea in hot, arid spots, we
find formations of Pistacio-Rhamnetalia alaterni, subdivided into maquis with
strawberry tree and tree heath (Erica arboreae), and maquis with carob, wild
olive (Oleo-Ceratonion) and common juniper (Juniperion turbinatae).
Where the maquis tends towards garrigue, low, discontinuous shrubby cover
occurs, with abundant wild lavender and rockrose (Cisto-Lavanduletea) on
acid soils, and maquis with many-flowered heath and rosemary (RosmarinoEricion, Rosmarinetalia) on limestone and marly soils.
■ Types of true maquis
Maquis with holm oak. High maquis with prevailing holm oak (Quercetum ilicis) represents one of the recently degraded, primary or secondary aspects of
holm oak woods, i.e., very similar to them in composition and physiognomy.
Holm oak maquis has more widespread distribution and thus greater importance with respect to the corresponding residual, fragmentary forest. Holm
oak prevails more or less absolutely, followed in order of importance by strawberry tree, lentisk, phillyrea, buckthorn, and other hard-leaved evergreen
shrubs. It is not a true association, but rather a stage determined and maintained by man’s activities. It is poorest in species at the northern limits of its
distribution. Near Duino (Venezia Giulia) it is composed only of holm oak, terebinth, Osyris alba and sarsaparilla. Further south, it is enriched with wild olive,
prickly juniper and phillyrea, becoming impoverished in evergreen species and
progressively mixed with deciduous trees and shrubs as altitude increases.
There is a variation with particularly abundant heath, which approaches
maquis with strawberry tree and tree heath, as we shall see later on. In Italy,
the distribution of holm oak maquis with altitude varies considerably due to
the great adaptational flexibility of holm oak itself. It ranges from sea level near
Trieste to more than 1000 metres on the slopes of Mt. Procinto (Apuan Alps),
Calabria, Sicily and Sardinia.
Holm oak maquis between Camogli and Punta Chiappa (Liguria)
Buckthorn and lentisk
Buckthorn, stinkwood
(Rhamnus alaternus)
Family: Ramnaceae.
An evergreen shrub, 1-5 m tall; trunk
ramified low down; hairy branchlets;
reddish-grey, finely striated bark; dark
yellow wood; spherical, compact foliage.
Simple leaves, alternate or
subopposed, hard, glabrous, oval,
pointed, margin saw-edged or entire,
3-6 cm long.
Yellowish-green flowers, with an
unpleasant odour, gathered in spherical
axillary racemes, no petals or only one.
Flowers from January to April. The fruit
is a spherical drupe, 0.5 cm across,
dark red when ripe, with three kernels.
The wood, excellent for fine
cabinet-making, is very heavy and
fine-grained, but it emits such a foetid
smell while it is being worked that it is
also known as “legno puzzo”
(stinking wood).
In dyeing, the leaves and fresh twigs
may be used to dye fabrics a fine
orange-yellow shade, and the fruit give
a “vegetal” green.
The drupe was once used medicinally
as a drastic purge.
Gaudenzio Paola
(Pistacia lentiscus)
Family: Anacardiaceae
An evergreen shrub (sometimes a small
tree, up to 5 m tall), dense, highly
ramified, rounded in shape, scaly brown
bark; small, glabrous, reddish-brown
Composite alternate leaves, with an
alate stalk, paripinnate, leaflets with
entire margins, obtuse or briefly
mucronate at the apex, pale green and
shiny above, paler and opaque
underneath, glabrous, with a strong
resinous odour. The flowers are
dioecious, in small, dense axillary
racemes, dark red in colour. Flowers
appears from March to June. The fruit is
a spherical-compressed drupe, reddish
or almost black when ripe.
The species is sun- and heat-loving,
and prefers siliceous, highly fertile soils.
A Sardinian proverb says: “Terra da
chessa, terra trigale” (Lentisk land,
good wheat land). The wood, with
yellow sapwood (alburnum) and reddish
heartwood (duramen), is hard, veined,
with poorly marked rings, but is not
often used owing to the small size of
the shrub. It makes excellent fuel. The
drupes provide oil as fuel and also for
cooking purposes. The resin which
weeps from gashes in the bark supplies
a mastic, which is used as an ingredient
in industrial paints and varnishes.
Heath and strawberry tree
Heath (Erica arborea)
Family: Ericaceae.
An evergreen shrub, with many
branches, 2-4 m tall, with an erect but
often contorted trunk, and rough
reddish-brown bark. Simple leaves, in
whorls of 3-4, linear, slightly stiff, 4-8
mm long, and very narrow.
Small, perfumed, pendent flowers,
gathered in thick racemes, white or
pink bell-shaped corolla, 2-3.5 mm
long, shorter than its stalk. The fruit is
in the form of a small capsule, divided
into 4 locules. Spring flowering.
Heath plays an important role in
maquis formations, but may also be
found high up, among broad-leaved
Where it covers large areas, heath
gives the landscape a characteristic
aspect. It was used in the past to
produce charcoal and to make coarse
brooms. Small bundles of its branches
were used to cover the roofs and walls
of poor people’s homes. The bunches
of dried branches on which silk-worms
were placed when they were ready to
spin their cocoons were often
composed of heath. When coppiced,
the plant produces a swelling at the
height of the collar, used to make the
bowls of pipes. Heath flowers are an
important source of nectar for bees.
Gaudenzio Paola
Strawberry tree (Arbutus unedo)
Family: Ericaceae.
An evergreen shrub, occasionally a
small tree growing to about 10 m, with
reddish-brown bark which flakes off in
thin scales.
Simple leaves, alternate, 4-10 cm long,
elliptical, crenate or saw-edged, dark
green and shiny above, paler
underneath, with short stalks.
Small flowers of 5-7 mm, in terminal
clusters, pendent, very thick, jarshaped corolla, 5-dentate, creamy
white in colour. Fruit in the form of
spherical berries, 1-2 cm in diameter,
granulous-tubercular on the surface,
first yellowish in colour, then orange,
and bright red when ripe, sweet and
edible, containing many seeds. The
fruit requires one year to mature, with
the result that, in autumn, the plant
may show both flowers and ripe fruit
from the previous year, at the same
Like all plants with winter fructification,
strawberry tree is very important for
birds at a time when little other food is
available for them.
Maquis with buckthorn and lentisk.
On limestone or limestone-marly
slopes, the degradation of holm oak
woods or the evolution of garrigue with
heath and rosemary creates very
dense maquis, quite homogeneous
because it is mainly composed of
buckthorn and lentisk, with the typical
lianas of holm oak, like sarsaparilla,
madderwort, Mediterranean honeysuckle and asparagus. This maquis is
frequent along the western Ligurian
riviera. The Pistacio lentisci-Rhamnetum alaterni association represents a
stage in the climacic series of holm
oak, but aridity due to type of soil and
lower rainfall greatly slows its evolution
with respect to maquis with strawberry
tree and tree heath.
Madderwort (Rubia peregrina)
Maquis with strawberry tree and tree heath. This type of maquis, mainly
silicicolous, located in the sub-Mediterranean belt, anticipates holm oak or
represents permanent communities in xerophytic biotopes, and is phytosociologically called Ericion arboreae. It is widespread along all Tyrrhenian
coastal areas. Strawberry tree prevails on siliceous, acid soils rich in humus,
with cool exposures, and is sometimes found quite high. Tree heath appears in
more degraded stages due to further aridity and impoverishment of the soil.
The stages of transition between maquis with strawberry tree, strawberry tree
plus heath, and prevailing heath, can all be identified. As these two main
species are the ones which vegetate first, sometimes only a few weeks after a
fire, their prevalence may be due to the practice of deliberately setting fire to
The association is well characterized by Pulicaria odora, and is constantly
accompanied by spiny broom, myrtle, lentisk, buckthorn, holm oak, sarsaparilla, madderwort, asparagus, sage-leaved rockrose, and pubescent oak.
From the dynamic viewpoint, Erico-Arbutetum is a quite evolved stage, mainly
belonging to the climacic series of holm oak or, further inland and at higher
altitudes, that of pubescent oak.
The evolution of these formations towards holm oak, in the absence of any
Tree euphorbia and prickly juniper
Gaudenzio Paola
Tree euphorbia (Euphorbia dendroides)
Prickly juniper (Juniperus oxycedrus)
Family: Euforbiaceae.
A large deciduous shrub, glabrous,
spherical, up to 3 m tall, with thick
foliage at the tips of branches, bluegreen in colour.
Thick, oblong-lanceolate leaves,
obtuse, with a mucro.
Umbrella-shaped inflorescence,
enclosed by bracts, with 5-8 rays,
rather thick and stubby. The fruit is in
the form of triangular capsules 5-6 mm
across. Flowers from November to May.
Unlike most Italian plants, it presents
the phenomenon of estivation - that is,
it enters a state of repose during the
hot, dry summer period, instead of
during winter. From June to September,
it loses its leaves and looks as if it were
dead, but then it starts vegetating again
and flowers, from the first late summer
rains onwards.
Lke many other species belonging to
the same genus, the trunk of tree
euphorbia contains a poisonous white
milky sap, once used to capture fish,
mainly freshwater ones: large quantities
of the branches of the tree were thrown
into ponds or specially prepared traps,
and the water thus poisoned.
Family: Cupressaceae.
An evergreen shrub 1-5 m tall, rarely a
tree, reaching 12-15 m.
Needle-shaped, very prickly leaves,
arranged in threes around branchlets,
with two lines above.
This is a dioecious species, i.e.,
specimens produce only male flowers
or only female flowers; the flowers are
devoid of involucre and are
inconspicuous. The fruit is a berry,
reddish-brown in colour and pruinous,
8-15 mm in diameter, maturing in the
second year. This plant was known in
ancient times, and was the subject of
many legends and beliefs, one of the
strangest being that its wood, if burnt,
would keep snakes away. The wood of
prickly juniper is very hard, excellent
for producing charcoal, and was also
used for making carved objects. The
berries are considered delicious by
birds. Cade oil or juniper-tar oil,
prepared by dry-distilling the wood,
has been used for thousands of years
to treat skin diseases – a reputation
which is highly justified, since the oil
has considerable disinfecting, resolvent
and curative properties.
kind of disturbance, is quite rapid, but
the periodic passage of fire causes
continual rejuvenation.
In acid soils in Liguria, aspects with
abundant green heath, honeysuckle
(Lonicera etrusca) and narrow-leaved
phillyrea may be identified.
Elsewhere, the destruction of holm oak
and cork oak groves has favoured
groupings composed of spiny broom,
daphne (Daphne gnidium) and sageleaved rockrose (Calicotomo infestaeEricetum arboreae), or spiny broom,
lentisk and rosemary (Pistacio lentisciCalicotometum villosae) or, in damper
locations, broom, myrtle and lentisk
(Erico arboreae-Myrtetum).
Dwarf palm (Chamaerops humilis)
Maquis with carob and wild olive. This type of maquis (Oleo-Ceratonion) is
the most heat-loving shrubby formation found in Italy. It extends all along the
central-southern Tyrrhenian coast and on the islands, composed of shrubs of
wild olive, carob, tree euphorbia, lentisk and Cneorum tricoccum. It is the permanent vegetation in cliff or beach locations with little soil, or where heatloving holm oak retreats.
In Italy, the distribution of this type of maquis is limited to low altitudes,
except for unusual locations in Calabria, Sicily and Sardinia, where it may
reach 600 metres.
According to which species are present and how they are associated, there
are different types with reciprocal dynamic affinities and relations.
● Maquis with wild olive and tree euphorbia. On rocky limestone coasts, we
find the association with Oleo-Euphorbietum dendroidis: tree euphorbia, wild
olive, prasium (Prasium majus), Jupiter’s beard, buckthorn and lentisk. Tree
euphorbia prevails, mainly along the warmest places of the mainland, definitely making its mark on the landscape.
● Maquis with lentisk and dwarf palm. Low maquis with lentisk, dwarf palm,
broom and rockrose develop along coastal limestone cliffs in Sicily and Sardinia, deriving both from holm oak groves and broom covers and representing
the Pistacio-Chamaeropetum humilis association, characterized by often
abundant and sometimes dominant dwarf palm.
Maquis with large-berried juniper and lentisk. Pistacio-Juniperetum macrocarpae, with large-berried juniper and various other types of broom, lentisk
and buckthorn, grow on both active and fossil dunes on sandy coasts.
Large-berried juniper is a type of juniper which, in Italian, is called ginepro coccolone, since it produces coccole, or large, hard, dark berries which, despite
their name (indicating sweetness and tenderness) are not edible.
● Maquis with shrubby scorpion vetch and diss grass. Coronillo valentinaeAmpelodesmetum mauritanicae, with diss grass (Ampelodesmos mauritanica),
shrubby scorpion vetch (Coronilla valentina), Spanish broom and tree euphorbia or Osyris alba, is a type of low maquis which grows on partially consolidated stony debris.
● Maquis with prickly juniper and lentisk. Pistacio lentisci-Juniperetum
oxycedri, with prickly juniper, lentisk, madderwort, strawberry tree, buckthorn
and diss grass, is a high, shrubby, pre-forest type of vegetation.
● Maquis with scorpion senna and many-flowered heath. In more
mesophilous locations, we find associations with many-flowered heath, scorpion senna (Coronilla emerus ssp. emeroides) and prickly juniper (Coronillo
emeroidis-Ericetum multiflorae).
● Maquis with Phoenician juniper and prasium. Teucrio-Juniperetum
phoeniceae, with Phoenician juniper, prasium, lentisk and buckthorn, forms
Low maquis in northern Sardinia
Phillyrea and Spanish broom
Gaudenzio Paola
Narrow-leaved, broad leaved phillyrea
(Phillyrea angustifolia, P. latifolia)
Spanish broom
(Spartium junceum)
Family: Oleaceae.
Small trees, up to 5 m tall, or more
often shrubs, evergreen, bushy, highly
ramified, with cylindrical, glabrous
Simple opposed hard leaves, with
short stalks, lanceolate or ovate,
rounded and slightly cordate at the
base, margins entire or saw-edged,
from obtuse to sharp at the apex,
shiny dark green above and paler
underneath. The two species are
distinguished according to the aspect
of the leaves, lanceolate and linear in
narrow-leaved phillyrea (Phillyrea
angustifolía) and ovate in the broadleaved variety (P. media).
Small, greenish flowers, arranged in
short racemes, axillary, with short
stalks; rotated corolla with 4 obtuse,
elliptical petals, greenish-white in
colour. Flowers from March to June.
The fruit is a spherical, apiculate
drupe, black when ripe.
The wood, pale brown with indistinct
rings, is hard and gives off an
unpleasant smell. It is excellent as fuel.
Family: Leguminosae.
A shrub growing to 2-3 m, with rushshaped, erect, cylindrical,
compressible branches. Leaves are
oblong and linear, hairy underneath,
and fall early.
Flowers are papilionaceous, large and
perfumed, in racemes; peduncles with
2 bractlets. The calyx is scarious, cleft
from the top to the bottom, 5denticulate. The corolla is golden
yellow, the vexilum being longer than
the other petals, subrounded. The
keel, with touching but free pieces, is
sharply rostrate. Flowers from May to
July. The fruit is a linear compressed
legume (4-8 cm x 5-7 mm), first silky,
and later bare.
Bees are attracted to the flowers, and
the many brown seeds are particularly
sought after by birds.
The leaves and young branchlets may
be used to dye fabrics yellow or
green, and the branches as rough
supports for vegetables in kitchengardens. Coarse twine may also be
dense, sun-loving and heat-loving maquis, and is found on rocky limestone
coasts, e.g., Oleo-Euphorbietum dendroidis.
● Maquis with large-berried juniper and lentisk. This type (Pistacio-Juniperetum macrocarpae) or, locally, ephedra (Ephedro-Juniperetum macrocarpae),
colonizes coastal dunes.
Sarsaparilla (Smilax aspera) with fruit
Juniper groves. Several species of
juniper form part of that type of
maquis called phytosociologically
Juniperion turbinatae: Phoenician,
prickly and, in cooler locations, common juniper (Juniperus communis),
but only the first two are typical of particular kinds of maquis.
● Maquis with Phoenician juniper. This
type of maquis is found along centralsouthern coasts and on the islands,
from sea level to 500 metres. On mainEverlasting flowers (Helichrysum stoechas)
land Italy, it is rather fragmentary and
discontinuous because, being a
coastal formation, it has been severely affected by man’s activities. Instead, in
Sardinia, it still composes a coastal belt with more or less extensive fragments.
Phytosociologically, it is differentiated: with tree heath on acid soil, wild olive on
limestone, and dwarf palm in the most arid spots. These juniper groves are
enriched by tree euphorbia on cliffs and helichryse, or everlasting flowers
(Helichrysum stoechas) on low rocky coasts. Where sand accumulates, we also
find large-berried juniper. More developed locations from the vegetational viewpoint may have some specimens of holm oak, cork oak or thorny oak.
● Maquis with large-berried juniper. This type colonizes sand dunes and is
therefore found only along beaches. It is a form of transitional vegetation
between the pioneering phytocenoses of sandy areas and the forest ones of
holm oak. As well as large-berried juniper, on dunes we also find lentisk,
clematis, Mediterranean honeysuckle, Phoenician juniper, buckthorn, narrowleaved phillyrea, sarsaparilla and many-flowered heath, with myrtle and pine
slightly further inland behind the dunes.
The many combinations of ecological factors which create the dune habitat
give rise to many different vegetal associations - Asparago-Juniperetum
macrocarpae, Spartio-Juniperetum macrocarpae, Phillyreo angustifoliaeJuniperetum turbinatae, and Phillyreo angustifoliae-Ericetum multiflorae.
Gaudenzio Paola
(Cistus albidus, Cistus creticus, Cistus
monspeliensis, Cistus salvifolius)
Family: Cistaceae.
Several species of the genus Cistus
grow in the Mediterranean maquis or
on the edges of evergreen woodland.
In spring and early summer, rockroses
enrich the landscape with their colours
and scents, thanks to their superb
flowers and the intense, characteristic
perfume released during the hottest
hours of the day by the leaves (the
flowers themselves are almost without
Members of the genus Cistus are
evergreen shrubs growing to 1-1.5 m
in height, more or less perfumed, with
opposed leaves, calyx composed of 35 persistent sepals, corolla with five,
easily detachable petals, and many
Fire often helps to spread the seeds of
the two species with white flowers,
Cistus monspeliensis and C. salvifolius.
The Montpellier rockrose (Cistus
monspeliensis) has particularly sticky,
sessile, lanceolate or linear leaves,
with sharp tips, downy on the
underside and with revolute edges.
The flower stalk and calyx are covered
with hairs; the flowers, in groups of 28, are white and highly scented, and
■ Low open maquis
This approaches garrigue, and mainly contains plants common to it.
may reach 3 cm in diameter.
The sage-leaved rockrose (Cistus
salvifolius) is so called because its
leaves recall those of sage. They are
ovate or elliptical, with one rib, and a
wrinkled surface. The flowers, up to 5
cm in diameter, have white petals,
longer than the sepals. Pink rockroses
(Cistus albidus and C. creticus) are
shrubs with greenish-grey downy
leaves, and flowers which are either
solitary or grouped in bunches or 2 or
3, with pink or pale purple wrinkled
The name of this family derives from
the Greek kistho, meaning “little box”,
which describes the shape of the
mature fruit: a capsule which,
suddenly opening, shoots out the
seeds. Traditionally, the plant is
attributed the significance of lightheartedness and inconstancy,
perhaps because the ephemeral petals
only last one day.
Maquis with rosemary. Sparse maquis, with garrigue shrubs like rosemary,
bush-like scorpion vetch (Coronilla juncea), shrubby thyme (Thymus capitatus), common thyme (Thymus vulgaris), Thymelaea hirsuta, many-flowered
heath, and various species of rockrose are the aspects of dynamically less
evolved maquis, frequent throughout mainland Italy and the islands. It is generally sparse and low, abundant in regions where fire is customarily used to
create grazing areas, and is one of the most beautiful of all Mediterranean
types of vegetation: in spring, in full bloom, it becomes a sea of large white
rockroses of various kinds. It grows on poor, only slightly evolved or even
eroded soils. Phytosociologically, these plants are part of the Rosmarinetea, of
the order Rosmarinetalia.
In coastal slopes in Liguria, we find communities with pink rockrose (C.
albidus), rosemary, Aleppo pine, Staehelina dubia and Coris monspeliensis,
forming part of Rosmarinion. In central and southern Italy and on the islands,
there are thermo- and meso-Mediterranean coastal formations with Cretan
rockrose (Cistus creticus ssp. eriocephalus) shrubby thyme, many-flowered
heath, and Greek savory (Micromeria graeca), belonging to the Cisto eriocephali-Ericion multiflorae alliance. This includes maquis on limestone with
Jupiter’s beard and globularia (Globularia alypum), dune formations with yellow rockrose (Halimium halimifolium) and many-flowered heath, populations
preferring lava cliffs, coastal sandstone with sage-leaved rockrose, and many
Maquis with Julian savory and Cretan rockrose. Again, this is a very varied
type of maquis, because it is only slightly evolved and thus depends greatly on
soil conditions. Here grow Cretan rockrose, Julian savory (Micromeria juliana),
thorny burnet (Sarcopoterium spinosum), and whorled heath (Erica manipuliflora). This type of maquis often contains plants which grow in the form of
semi-spherical cushions, in order to cope better with the harsh ecological conditions in which they manage to grow.
In Italy, it only occurs along the southern coasts of the Adriatic and Ionian and
on the reliefs of the southern Appennines, even quite high (on the promontory
of Gargano, it grows on windy limestone slopes between 500 and 680
metres). Phytosociologically, it is part of the Cisto cretici-Micromerietea
julianae class.
Maquis with wild lavender and rockrose. The more acid-, sun- and heatloving maquis which develops on only slightly evolved soils subject to erosion, in a dry Mediterranean climate, degraded by repeated fires, is mainly
formed of Montpellier rockrose, sage-leaved rockrose and wild lavender, all
plants which prefer siliceous soils. In western Liguria at altitudes between 200
and 1000 m, we find specimens of this formation, with green heath. In eastern
Liguria, the degradation of maquis with heath and strawberry tree produces
formations with thyme, spiny broom and hairy broom (Genista pilosa), as well
as rockroses. Phytosociologically, these vegetation types are part of the Cistoladaniferi-Lavanduletea stroechadis class.
Maquis with broom. This type characteristically presents several kinds of
broom endemic to the coasts and small islands of the southern Tyrrhenian and
northern Sardinia. Phytosociologically, it belongs to the Calicotomo villosaeGeniston tyrrhenae alliance.
■ Other shrubby formations
Maquis with oleander. Oleander (Nerium oleander) may reach tree size, but in
Italy, in the spontaneous state, it mainly grows as a bush. It prefers the
warmest parts of the Mediterranean region, especially Calabria, Sicilia and
Sardinia, where it is found as a shrub along watercourses and the almost
always dry beds of small rivers. Elsewhere, it is much rarer and probably
grows wild after having once been cultivated. Its northern limit on the mainland
is in the province of Salerno, but it is said that at Camporosso (“red field”), a
village in Liguria at the mouth of the seasonal river Nervia, there used to be
spectacular flowerings of oleander in the riverbed, before the intense
anthropization which now dominates the area. On the island of Capraia (Tuscan Archipelago), there is a type of riverbed maquis with oleander, buckthorn,
sarsaparilla, honeysuckle and ivy (Festuco corsicae-Nerietum oleandri), confined to areas along short watercourses.
Thus, special environmental conditions allow the spontaneous growth of oleander at the extreme south of the Italian mainland and in the large islands. Maquis
with oleander is almost completely independent of true Mediterranean maquis.
It is a stable formation of edaphic type, i.e., associated with a particular soil.
The areas of maquis with oleander which are found along the rivers of southern Italy and the islands are probably the remains of vegetation which in
ancient times was far richer. Their distribution in Italy is limited to low altitudes.
In river-bank and riverbed vegetation, oleander is accompanied by alder
Myrtle and laurel
Gaudenzio Paola
Myrtle (Myrtus communis)
Laurel (Laurus nobilis)
Family: Myrtaceae.
An evergreen shrub, sometimes a small
tree, up to 5 m high, thick, highly
ramified, with opposite branches.
Simple leaves, opposite or in whorls of
3, sessile, ovate-lanceolate, sharp, with
entire margins, dark green in colour,
shiny on top and paler and more
opaque underneath, very sweetsmelling.
Flowers are large, hermaphroditic,
solitary or in pairs, axillary, pedunculate;
the calyx has 5 free, pointed sepals; the
corolla has 5 obovate, white petals,
hairy and glandulose at the edges, and
very numerous stamens, longer than the
petals, with yellow anthers. It flowers
from summer to autunm. The fruit is a
spherical or ovate berry, blue-black in
colour, sweet in taste and sweetscented, and containing many white
The wood, hard, pinkish-brown, and
with indistinct rings, is used to make
small lathe-turned or carved articles,
including objects like handles and
walking-sticks. As a fuel, the wood
burns wells and produces excellent
charcoal. The myrtle was sacred to
Venus, and myrtle crowns were used as
nuptial garlands. In cooking, the fruit are
used to enhance flavour.
Family: Lauraceae.
An evergreen species, generally a thick
shrub, 2-5 m tall. In favourable
ecological conditions, it may even be
found as a tree exceeding 10 m in
height, with very dense, spherical
foliage, and smooth, dark brown bark.
Leaves are simple, alternate, 5-10 x 2-4
cm, elliptical, pointed, hard, shiny, with
undulate edges, stalk 6-10 mm long,
sweet-smelling if rubbed or crushed.
Small, yellowish-green, unisexual
flowers, in inflorescences at the axils of
the leaves. The fruit is a black, ovoid
berry, slightly more than 1 cm long.
A species of uncertain origin, probably
from western Asia, cultivated since
ancient times. The fruit is considered
delicious by birds, which are
responsible for the considerable
expansion of this species in some
The laurel was dedicated to the god
Apollo, and still enjoys fame as a
glorious and beneficial plant. Laurus is
the ancient Latin name of this species.
According to some, it derives from the
Celtic word blaur (green) and, to others,
from the Latin laus (praise), referring to
the crown of laurel leaves used by the
Romans as a sign of recognition for
noble actions.
Spiny broom and sarsaparilla
Spiny broom (Calicotome spinosa)
Family: Leguminosae.
A very thorny shrub which may exceed
2 m in height, with few branches.
The leaves, silky underneath, petiolate
and subdivided into three leaflets,
characteristally fall in summer, leaving
only shiny, black legumes and thorny
branches. Small, yellow-gold flowers,
solitary or in bunches of 2 or 3, appear
from February to May.
Spectacular and abundant spring
flowering adorns the hottest and most
arid slopes.
However, the shrub is untouchable,
due to the thorns which it has
developed as a form of adaptation to
attack by herbivores, and which
involve the transformation of the tips
of the branches into very sharp,
strong thorns. Precisely for this
particular form of defence, it was used
to create impenetrable hedges to
protect cultivated areas.
The whole plant defends itself even
further by containing an alkaloid,
which is poisonous to animals.
Gaudenzio Paola
Sarsaparilla (Smilax aspera)
Family: Liliaceae.
A liana-like evergreen plant, with a
woody stem, highly ramified and
twisted, glabrous and tenacious, with
backward-facing thorns, which are also
found along the edges of leaves and
on ribs, whence its popular Italian
name of “tears-your-trousers”.
Leaves are simple, alternate, hard,
shiny, heart-shaped, 8-10 cm long,
pointed round the edges and along the
ribbing of the underside. The stalk is 23 cm long with 2 tendrils at the base.
Flowers are small, yellow or green in
colour, in inflorescences at the axils of
the leaves or at the tip of the stem.
Fruit are globous berries with 1-3
seeds, first green, then yellow, and
turning to red at full ripening. Although
not toxic, they are insipid to the taste,
but form an excellent source of food
for birds, since they persist on
branches even in winter. The medicinal
varieties of sarsaparilla originating from
central and south America were once
believed to cure syphilis. Nowadays,
true sarsaparilla (Smilax officinalis) is
only recognized for its diaphoretic and
purifying properties, which are due to
the saponines it contains.
(Alnus glutinosa), narrow-leaved ash (Fraxinus angustifolia), lesser maple (Acer
monspessulanus), various types of willow, and rarely with true maquis components (buckthorn, lentisk, phillyrea, wild olive, wild vine, ivy, honey-suckle, sarsaparilla). The shrubby vegetation along riverbeds is phytosociologically part
of the Nerio-Tamaricetea association, typical species being oleander, tamerisk
(Tamarix africana, T. gallica) and chaste tree (Vitex agnus-castus). The plants of
this association thus compose warmth-loving, hygrophilous, riverbank undergrowth, physiognomically characterized by oleander and tamerisk, associated
with a warm Mediterranean bioclimate and suited to periodic drying up of
watercourses. They also grow along the alluvial terraces of riverbeds, are only
exceptionally influenced by seasonal flooding, and are widespead in the
hottest parts of the Mediterranean region. There are three different types in
Calabria and Sicily: Tamarici africanae-Viticetum agni-casti, Spartio-Nerietum
oleandri and Rubo-Nerietum oleandri. The first, Tamarici africanae-viticetum
agni-casti, composed of river-bank undergrowth, typically contains African
tamerisk and chaste tree, with oleander, willow and brambles. It is found at the
mouths of seasonal rivers where the water-table is not too deep.
Spartio-Nerietum oleandri is made up of riverbank undergrowth with oleander,
Spanish broom, spiny broom, tamerisk and bramble (Rubus ulmifolius). It is
found almost exclusively on pebbly soil in the narrowest stretches of riverbeds and has been described for Sicily. Instead, Rubo-Nerietum oleandri
forms riverbank undergrowth with oleander, common bramble and African
tamerisk, preferring quite steep riverbeds with rocky outcrops, and favouring
stretches affected by seasonal flooding.
Maquis with laurel. Laurel may be found spontaneously or subspontaneously
in small, warm valleys, protected from too much sunlight, along the shores of
the large North Italian lakes, whereas in mainland Italy it is found on the inner
and upper margins of cooler valleys in coastal hills, in depressions, and along
damper slopes. Laurel is accompanied by other plants called lauriform, due to
the shape, anatomy and physiology of their leaves which are typically broad,
hard and evergreen.
Maquis with laurel is the extreme expression of Mediterranean maquis in the
“hot-humid” direction. Currently, only a very tiny, marginal part of Mediterranean maquis reaches this aspect, but in the past, during warmer, damper climatic periods - like the Tertiary, or more recent interglacial periods of oscillations in climate towards oceanic types - this type of formation was more widespread. Today, therefore, it is reduced to a few fragments. In any case, in Italy,
laurel is definitely not spontaneous everywhere. It appears to have been intro-
duced in many places, and later
became naturalized. But there are
some aspects of vegetation with prevailing laurel, in shrub and tree form,
which unquestionably indicate its
spontaneous origin. Its compact
foliage creates deep shadow inside
the undergrowth, so that only butcher’s broom and ivy can occasionally
Eastern hornbeam (Carpinus orientalis)
grow. But it does mix with other plants:
further north are many broad-leaved
trees: pubescent oak, chestnut (Castanea sativa), terebinth, eastern hornbeam
(Carpinus orientalis), hazel (Corylus avellana), fig, and manna or flowering ash,
with very few evergreens, such as butcher’s broom, laurustine, and some
ferns. Further south, evergreen maquis species increase (holm oak, wild olive,
phillyrea, sarsaparilla, rose, lentisk, strawberry tree, tree heath), but pubescent
oak, terebinth and flowering ash remain here and there.
Fragments of maquis with laurel occur all along the Italian coastline, but only in
the larger islands, in the heart of the Mediterranean region, are there true
examples of this type. They confirm their characteristic as relicts from another
climate, plants which took refuge in favourable microclimatic areas, at the
boundary between the area of Mediterranean evergreen forest and submountain deciduous forest. As regards altitude, due to orographic factors, maquis
with laurel may descend to the coast, but generally grows on south-facing
slopes between 500 and 600 metres.
Pseudo-maquis. This type of maquis grows outside its proper ecological
boundaries as regards altitude and latitude, and thus appears in less typical
forms, removed from its true Mediterranean character. It is also called
mesophilous maquis.
● Pseudo-maquis with box. Here, we find box associated with holm oak, terebinth, prickly juniper, spiny broom, winter savory (Satureja montana) and rockrose. In the Ligurian Appennines, it may grow between 200 and 800 m and in
the Central Appennines between 500 and 800 m.
● Pseudo-maquis with gorse. Gorse (Ulex europaeus) is a very thorny shrub,
which prefers siliceous soil and has a sub-Atlantic distribution.
Due to its climatic requirements, it may be considered a moorland plant. In
Italy, it is limited to the Tyrrhenian flank of the mainland, and some fine examples exist in southernmost Liguria (Levante) and above all in Tuscany.
Gorse (Ulex europaeus)
Fungi of the maquis
Mirca Zotti
There are many species of fungi which
grow in the Mediterranean maquis.
Conditions favouring abundant fungal
flora in maquis environments (closely
associated with the degree of humidity)
only occur from late autumn to the end
of winter and in spring.
Among the Amanita, the best-known and
certainly most appreciated, due to its
delicious flavour, is royal agaric, or
Caesar’s mushroom (Amanita caesarea).
This mushroom prefers high maquis with
prevailing holm oak. It has a fine reddishorange cap, dark ochre or yellow
lamellae and stalk, and white volva. In its
early stages of development, it looks like
a whitish “egg”, and may be confused at
this stage with several other species, all
belonging to the genus Amanita and
occurring in the same environment, like
the lethally poisonous Amanita
phalloides and A. verna. Other amanitas
of the high maquis with holm oak are
A. echinocephala, white or pale grey in
colour, clearly identifiable by the thick,
pointed warts decorating the whole
surface of the cap; A. aspera, with a
greyish-brown cap covered with minute
pale yellow warts; A. ovoidea and
A. proxima. Typical of the low maquis in
sunny spots and on sandy soils are
A. curtipes and A. ponderosa, both with
a relatively short, stubby stalk, well sunk
into the earth. They are typically
Mediterranean species; in Italy, they are
found in the south and the islands.
Many showy fungi belong to the boletus
family, and are mainly characterized by
having their fertile portions
(hymenophore) under the cap, not
composed of lamellae but of a set of
tightly bound tubules arranged so as to
form a compact mass, easily separated
from the flesh of the cap.
Among the mushrooms of the boletus
family associated with holm oak, the
most common and best-known for its
delicious flavour is Leccinum lepidum.
The colour of the cap is very variable
and ranges from yellowish or yellowishbrown to dark brown. The hymenophore
is first yellow, but then becomes ochre
yellow with olive-green tones.
Another species belonging to the
genus Leccinum, more typically found in
the low maquis and believed to be
closely associated with Montpellier
rockrose, is L. corsicum, an edible
mushroom widespread in the south of
Italy and the islands.
Also very good to eat and found growing
near holm oak or cork oak is Boletus
impolitus, with a velvety, irregularly fawncoloured or brown cap, and flesh which
does not change colour when cut. It is
frequent in the south and very common
on the islands, but is quite rare in the
north. Another warmth-loving species,
with a brilliant carmine red stalk and
pores is Boletus rhodoxanthus.
Again with a tubular hymenophore but
smaller than the above species are
Xerocomus persicolor, X. rubellus and
X. dryophilus. The last two have brilliant
caps, ranging from red to bright pink.
The genus Russula, with a generally
stubby stalk, never with a ring, a brightly
coloured cap, and tender sweet- or soursmelling flesh, occurs in the maquis, and
has both edible and inedible types.
Other cosmopolitan and very common
species found in these environments are
Russula cyanoxantha, R. vesca,
R. heterophylla, R. delica and
R. nigricans. More closely associated
with holm oak are R. ilicis, R. prinophila,
R. nuragica and R. atramentosa.
R. monspeliensis and R. cistoadelpha
may often be noted under rockrose.
Milkcap (genus Lactarius), similar to the
above, has flesh which is structurally
similar, but weeps more or less
copiously when cut and may change
colour upon contact with the air. Species
occurring in the maquis include Lactarius
tesquorum, L. mairei and its variety ilicis.
One of the most frequent types, clearly
visible because of its bright tawnyorange cap, is L. atlanticus.
Members of the Hygrophorus family,
identified by the widely spaced lamellae
decurrent down the stalk, have
numerous species. H. unicolor is beigeochre in colour, with a reddish-brown
centre to the cap. H. pseudodiscoideus
var. cistophilus may be identified
because it grows under rockrose and
has tender flesh. Neither is considered to
be edible. Instead, H. nemoreus, and
H. russula are considered good to eat.
The maquis also contains many species
belonging to the genus Cortinarius. None
of these beautifully coloured species is
considered edible. Many of them prefer
both high and low maquis, like
Cortinarius glaucopus, with an ochrecoloured cap; C. splendens, with
golden-yellow cap, lamellae and stalk, is
definitely poisonous.
Of the very many other species of fungi
which may be found in the various types
of Mediterranean maquis are Clitocybe
font-queri, Hebeloma cistophilum and
Ripartites strigiceps, commonly growing
with rockrose, and some very frequent
species growing on wood, like Polyporus
meridionalis, found on twigs of rockrose
and rosemary embedded in the soil;
Phellinus torulosus, growing at the base
of holm oak trunks; and Ganoderma
lucidum, sought after thanks to its
therapeutic properties.
Amanita ovoidea
Leccinum lepidum
Hygrophorus russula
Cortinarius ionochlorus
■ From one kind of maquis to another all round Italy
A more or less wide belt of Mediterranean vegetation runs the whole way round
Italy, from the Ligurian Riviera to Trieste, only interrupted by the mouth of the Po
as far as Friuli. Sicily and Sardinia, except for their highest reliefs, are almost
entirely covered.
Leaves of dwarf palm emerge from thick maquis (Circeo promontory, Latium)
The Tyrrhenian coast. In Liguria, the polymorphism of the maquis is further
complicated by the climatic, geomorphological, lithological and phytogeographic factors so typical of the region. Coastal Liguria enjoys a mild climate,
with relatively high annual rainfall. This particular climate, which is anomalous
with respect to latitude, allows plant species and associations from temperatehot climates to grow, especially near sea level.
The western Ligurian riviera (Ponente) contains maquis with rockrose, ascribable to Rosmarino-Ericion, and shrubby formations (Quercetea ilicis) of two different types: heath plus strawberry tree, and buckthorn plus lentisk.
Maquis with tree heath and/or green heath plus strawberry tree colonize moderately steep slopes, generally facing south or west, between 50 and 400 m
above sea level, mainly in central and eastern Liguria. The most “closed” types,
exclusive to western Liguria (Levante), with a tall tree layer reaching about 5 m
and a lower shrubby layer to about 1.5 m, may be considered as variants, and
are differentiated by broad-leaved phillyrea, greater spleenwort (Asplenium
onopteris), friar’s cowl, or Italian Jack-in-the-pulpit (Arisarum vulgare) and black
bryony (Tamus communis). This aspect locally represents the most highly
evolved stage, preceding the formation of holm oak woods. In much of the
region, less highly evolved stages are much more widespread. These formations are normally less “closed” and in any case have a single shrubby layer
between 1.5 and 2 m in height, mainly differentiated due to the presence of
spiny broom, yellow rockrose and daphne. The central and western areas contain a variant exclusive to acid soils, with narrow-leaved phillyrea, green heath
and Mediterranean honeysuckle.
Warmth-loving maquis with buckthorn and lentisk, attributable to the Pistacio
lentisci-Rhamnetum alaterni association, is exclusive to the Riviera di Ponente,
where it is located at altitudes between 10 and 350 m above sea level, mainly
on limestone or limestone-marly soils. It represents the degradation of holm
oak on limestone soils or a later, garrigue stage of Rosmarino-Ericion. Transitional stages are frequent, where species of the Pistacio-Rhamnetum association mingle with typical garrigue species such as rosemary, Staehelina and
helichryse. These formations almost always contain Aleppo pine, a relict of
previously planted woodland or spontaneously growing after the passage of fire.
Then there are fragmentary, small formations, representing particular coastal situations, mainly on cliffs, in the warmest areas of the two Rivieras, western and
eastern, with constant, often abundant tree euphorbia. Asparagus, madderwort,
greater spleenwort and sarsaparilla indicate that these formations are stages of
holm oak reconstruction rather than the Oleo-Euphorbietum dendroidis association, definitely more heat-loving and widespread in areas further south. Landslides and the abandon of agriculture favour slope instability and the expansion
of coarse, well-aerated soils, factors which favour the growth of tree euphorbia.
Most of these aspects may be found along the coast between the frontier with
France and the town of Savona, and on the southern flanks of the promontories of eastern Liguria (Portofino, Punta Manara, Punta Baffe, Punta Mesco,
the five villages composing the Cinque Terre, and Portovenere).
Another particular case is that of the shrubs in the central part of the region,
between the villages of Varazze and Cogoleto, where ophiolithic formations
with metagabbro and serpentinite outcrop. Here we find shrubby associations with tree heath and green heath, structurally very similar to EricoArbutetum but with floristic composition slightly impoverished in species of
the Pistacio-Rhamnetalia alaterni order and without strawberry tree, probably
because of the soil.
Locally, the pseudo-maquis is dominated by box (Buxus sempervirens) with
Salzmann’s broom (Genista salzmannii), junipers (Juniperus communis, J.
oxycedrus) and tree heath. Further east, with higher humidity, are examples of
pseudo-maquis with gorse in the undergrowth of pine woods.
The Tuscan coast also has a considerable variety of plants, with occasional formations of hard-leaved evergreen maquis on coastal cliffs, consolidated dunes
and back-dunes, and hills further inland. Where the coast is low, there is only a
narrow belt of spontaneous vegetation, bounded on one side by the beach and
by fields and buildings on the other.
On the plain of the river Arno, between Viareggio and Livorno, this belt is formed
of woods of maritime pine with maquis underneath. In the coastal stretch north
of the Arno, this maquis contains large-berried juniper and Spanish broom
(Spartio juncei-Juniperetum macrocarpae), characterized by Spanish broom,
wall germander (Teucrium flavum) and wild pear (Pyrus pyraster), and a great
reduction in holm oak species, following the slightly damper climate of the area.
In the Migliarino-San Rossore area, again characterized by a local microclimate
tending towards more continental conditions, consolidated dunes are colonized by shrubs with dominant Spanish broom and narrow-leaved phillyrea
(Spartio juncei-Phillyretum angustifoliae). These formations are often in contact
with maquis with large-berried juniper and Spanish broom, and represent either
a pioneering stage or degradation after fires.
Between Ardenza and Rosignano, hills approach the coast, allowing the expan-
Maquis with Phoenician juniper, buckthorn, rosemary, Montpellier rockrose and holm oak (Tuscany)
Maquis along rocky coastline on island of Elba (Tuscany)
sion of natural vegetation. But the true triumph of maquis may be admired on
the promontories which extend towards the islands of the Tuscan Archipelago Piombino, Punta Ala, the mountains of Uccellina and Monte Argentario - and on
the islands themselves.
Punta Ala hosts a low, coastal or internal maquis, on limestone soil, with rosemary, Montpellier rockrose, many-flowered heath, tree heath, lentisk, narrowleaved phillyrea and holm oak (Erico-Rosmarinetum). On the hills further
inland, with siliceous soil, is a type of low maquis with acid-loving species like
ling, green heath, wild lavender and yellow rockrose (Ampelodesmo-Ericetum
scopariae). On the cliffs grows a type of maquis with Phoenician juniper and
wild olive (Teucrio-Juniperetum phoeniceae), together with prasium, lentisk,
buckthorn and sarsaparilla, or heat-loving types with tree euphorbia (OleoEuphorbietum dendroidis) with Jupiter’s beard, shrubby scorpion vetch,
lentisk, Phoenician juniper and dwarf palm. Dunes hold maquis with largeberried juniper and lentisk (Pistacio-Juniperetum macrocarpae) with clematis,
Mediterranean honeysuckle, Phoenician juniper, buckthorn, narrow-leaved
phillyrea, sarsaparilla and many-flowered heath, while further inland behind
the dunes are myrtle, many-flowered heath, narrow-leaved phillyrea, largeberried juniper and pines (Phillyreo-Ericetum multiflorae).
On the mountains of Uccellina, we find holm oak woods, according to exposure
and the trends of slopes and valleys, tending towards mixed groups of hardand broad-leaved deciduous trees, groups of hard-leaved evergreens and various types of maquis: slightly evolved, deriving from holm oak on siliceous soils
with myrtle, lentisk, narrow-leaved phillyrea and spiny broom; coastal maquis
with Phoenician juniper (Teucrio-Juniperetum phoeniceae) with germander,
prasium, lentisk and wild olive; heat-loving maquis with tree euphorbia (OleoEuphorbietum dendroidis) located on limestone coasts, with Jupiter’s beard,
shrubby scorpion vetch, lentisk, Phoenician juniper and dwarf palm; and, lastly,
low maquis with rosemary, many-flowered heath, Montpellier rockrose and
globularia, defined phytosociologically as Erico-Rosmarinetum.
On the Maremma coast (Punta Ala, Lago di Burano), the more inland dunes, now
consolidated, are colonized by formations of pioneer maquis with large-berried
juniper and asparagus, representing transition vegetation between pioneering
phytocenoses and forest types typical of holm oak woodland. These are dense
formations, the floristic cortège of which shows good cover of shrubby species
such as buckthorn, lentisk, and both narrow- and broad-leaved phillyrea, of the
Asparago acutifolii-Juniperetum macrocarpae association and its sub-association juniperetosum turbinatae, if with Phoenician juniper.
On the island of Elba, in Maremma, along the Cecina littoral north of Rome, and
at Tombolo di Feniglia (Orbetello), maquis with large-berried juniper and
asparagus is often replaced by shrubs dominated by Phoenician juniper, lentisk
and narrow-leaved phillyrea (Phillyreo angustifoliae-Juniperetum turbinatae) on
the most consolidated inland dunes, often in contact with marshy depressions.
The oldest consolidated dunes along the stretch of coast between Punta Ala
and Orbetello have thick maquis, between 1 and 3 metres tall, with prevailing
narrow-leaved phillyrea, many-flowered heath and myrtle, often growing under
artificially planted woods of maritime pine and umbrella pine (Pinus pinea). This
is part of the Phillyreo angustifoliae-Ericetum multiflorae association, representing degradation or a pioneering stage of the associations quoted above, with
which it is dynamically and spatially in contact. Garrigue formations, typical of
post-fire situations, are linked with these kinds of maquis, with Cretan rockrose,
canary clover (Dorycnium hirsutum) and everlasting flowers, called phytosociologically Helichryso stoechadis-Cistetum eriocephali.
Succession of vegetation along sandy coast of Tuscany, from the sea inland. Psammophilous
formations ( 1 ) colonize the first dune front, as far as its summit; immediately behind the dune, maquis
with large-berried juniper begins to appear ( 2 ), more or less wide according to area available; further
inland, this type of maquis is replaced by narrow-leaved phillyrea and many-flowered heath ( 3 ), in turn
giving way to hard-leaved Mediterranean formations (holm oak, 4 )
On Elba and along the Tuscan littoral near Orbetello, there are also formations
of garrigue with rosemary and rockrose (Cisto eriocephali-Rosmarinetum officinalis), linked dynamically with maquis containing many-flowered heath and
narrow-leaved phillyrea.
The small islands of the Tuscan Archipelago, like areas along the mainland
coast, have low maquis dominated by rosemary, heath and Jupiter’s beard
(Cisto eriocephali-Rosmarinetum officinalis). These formations are low, thick,
and characterized by a prevailing shrubby layer covering between 90 and
100% of the land, on average 80 cm high, with only a few sun-loving suffruti-
cous plants of the genus Cistus and many tendentially limestone- and heatloving shrubs like rosemary, many-flowered heath, germander and lentisk. The
most highly developed vegetational stage on the islands of the Tuscan Arcipelago are aspects of maquis with lentisk and Jupiter’s beard. Also found on the
small islands of Giannutri and Capraia and on the flanks of Uccellina, these phytocenoses belong to the Myrto communis-Pistacietum lentisci association.
Low maquis with rosemary and many-flowered heath cover many parts of the
coastline and islands of Tuscany: Punta Ala, Maremma, M. Argentario, the
islands of Giannutri and Elba, and Orbetello. The littoral north and south of
Rome (Latium) is similar to that of Tuscany: a very narrow, fragmented strip of
maquis near the low coast (a few specimens only remain at Palo Laziale,
Ladispoli, Castelporziano and Sabaudia), with broad-leaved phillyrea, tree
heath, strawberry tree, lentisk, buckthorn and myrtle, or holm oak and tree
heath. Instead, at Furbara, Macchiatonda and Pratica di Mare, we find remaining fragments of coastal woodland with elm (Ulmus minor) and laurel. Fregene,
Torre Astura and Sabaudia lie on the remains of dunes colonized with largeberried juniper, Phoenician juniper, broad-leaved phillyrea, daphne, rockrose,
sarsaparilla, asparagus and madderwort. But more frequently along these
consolidated dunes of the Roman littoral, we find garrigue with Cretan rockrose
and everlasting flowers (Helichryso stoechadis-Cistetum eriocephali).
To an even greater extent than in Tuscany, the maquis penetrates inland on
south-facing slopes along the hills of Latium. Maquis with holm oak, buckthorn
and laurel may be found on the cliffs above the narrow gorges of the river Fiora
(Montalto di Castro, Vulsini mountains). On the Rocca della Tolfa are cork oak
with maquis composed of tree heath, strawberry tree, germander, broad-leaved
phillyrea, flowering ash, cerris or turkey oak, cyclamen and butcher’s broom.
On the Tolfa mountains, we find the first examples of the southern Mediterranean seasonal rivers, with African tamerisk and chaste tree in the riverbeds
and maquis with trees from the Eastern Mediterranean such as Judas tree (Cercis siliquastrum) and Christ’s thorn (Paliurus spina-christi). The Cimini mountains contain broom preferring acid soils, with European broom and, on the
sunniest slopes, Cretan rockrose, sage-leaved rockrose, and Spanish broom.
The valleys of the rivers Treia and Tevere (Tiber) (e.g., on Monte Soratte or at
Civitella San Paolo) contain pockets of Mediterranean climate and vegetation,
with holm oak, lesser maple, black hornbeam (Ostrya carpinifolia), flowering
ash, laurustine, lentisk, buckthorn, sarsaparilla, asparagus and madderwort
and, at Nazzano, on south-facing sandstone cliffs, even rosemary and wild
olive. On Monte Gennaro, maquis with holm oak, myrtle, broad-leaved
phillyrea and wild olive reaches altitudes of more than 1000 metres (a phenomenon also found on the Ernici and Lepini mountains), often with interesting
plants of eastern origin such as Judas tree, eastern hornbeam, storax (Styrax
officinalis) and Christ’s thorn.
The Alban Hills near Rome contain some maquis with holm oak, strawberry tree,
laurustine, buckthorn, tree heath, box, butcher’s broom, and occasionally laurel.
The maquis expands in particular on promontories, mainly the Circeo headland,
which is a protected National Park. Monte Circeo contains extensive tree heath,
Vegetation on southern flank of Circeo promontory. The highest belt has best preserved vegetation,
mainly high, hard-leaved Mediterranean maquis, with prevailing holm oak ( 1 ); lower down, shrubby
formations with myrtle and lentisk ( 2 ) appear, giving way in a degraded fashion to plant cover with diss
grass, rockrose and other species associated with the repeated passage of fire ( 3 )
Christ’s thorn (Paliurus spina-christi)
The maquis slowly recolonizes slopes degraded by fire (Circeo promontory, Latium)
Tyrrhenian coast, rustic and heat-loving, colonizes abandoned terraces together
with tree heath, sage-leaved rockrose, wild lavender and Montpellier rockrose.
On the island of Zannone is a coppiced area of holm oak with strawberry tree
and tree heath on the northern slopes, and low maquis with myrtle, lentisk and
wild lavender on the southern ones. The littoral of the Gulfs of Gaeta and
Naples, and the plains of Volturno and Garigliano have Mediterranean maquis
with holm oak, laurustine, terebinth, strawberry tree, and plants of eastern origin such as oriental hornbeam, black hornbeam, Judas tree and flowering ash.
The abundance of deciduous trees, even along the coastline and on the
islands, denotes a climate with high rainfall. Thus, a true heat-loving Mediterranean belt with wild olive, lentisk and Phoenician juniper is missing, except
for some cliff sites on promontories and adjacent reliefs (Phlegraean Fields,
Vesuvius, Lattari mountains).
On the southern slopes of the Alburni mountains and the Appennines in
Lucania, maquis with tree euphorbia and wild olive reaches the highest altitudes in mainland Italy (600 m) and the greatest distance from the sea, exploi-
myrtle, green heath, flowering ash, asparagus, and a few specimens of a strange
oak (Quercus crenata), probably originating from hybridization of cork oak with
turkey oak. The northern flanks are covered with tall, thick maquis or evergreens,
mainly holm oak, whereas the southern flanks have tall maquis, mainly with holm
oak, Phoenician juniper and strawberry tree, and degraded situations with diss
grass, myrtle, lentisk and rockrose. However, the most beautiful formations are
those fragments of maquis which manage to sink their roots in limestone cliffs,
with heat-loving species such as dwarf palm, tree euphorbia, many-flowered
heath, Jupiter’s beard, prasium and Phoenician juniper, or mesophilous varieties
like holm oak, broad-leaved phillyrea and flowering ash.
The Ausoni and Aurunci mountains, along the coast, are composed of carbonatic rocks, and thus host abundant maquis. The type with the most thermophilous trees is located on the cliffs of Sperlonga and Gaeta. These cliffs
also host a type of maquis with tree euphorbia, dwarf palm, Phoenician juniper,
Jupiter’s beard, wild olive and lentisk, while further inland are other types with
holm oak, cork oak and laurel, or with strawberry tree and tree heath.
On the island of Ponza and its accompanying tiny islands, are fragments of tall
maquis tending towards holm oak, with abundant lentisk, and low maquis with
species of the Oleo-Ceratonion association mixed with holm oak. Occasionally,
according to location, predominant plants are tree heath, tree euphorbia,
Artemisia arborescens and, especially on abandoned terraces, Spanish broom.
Another type of broom (Genista tyrrhena), endemic to the central-southern
Cross-section of vegetal cover along high coasts of island of Ponza. A: original vegetation, composed of
low maquis ( 1 ) on high, more exposed areas and holm oak on cooler slopes ( 2 ); maquis and holm oak
are respectively replaced by cultivated land and vineyards, the latter on artificial terraces. B: abandon of
cultivation allows spontaneous species to recolonize these areas. Previously cultivated flat land is
covered with arid Mediterranean meadows ( 3 ), whereas terraces are first invaded mainly by broom with
high colonizing power (Genista tyrrhena, 4 ) which, later, is accompanied by maquis species ( 5 ).
The development of both situations moves towards the reconstitution of a maquis cover, as was
originally the case
Carob (Ceratonia siliqua)
Winter savory (Satureja montana)
Terebinth (Pistacia terebinthus)
ting the fact that vegetation can penetrate along the valley of the river Sele.
The heavy anthropic burden borne by
the coastline of Campania (Bay of
Naples, Sorrento peninsula and nearby
islands) has meant that space available
for natural vegetation has been greatly
reduced. On Capri and Ischia and along
the coast, flourishing remnants of
Mediterranean maquis only remain in
the least accessible small valleys. The
Cilento area, instead, contains some
splendid examples of maquis, sometimes heat-loving, now protected by the
creation of a National Park. However,
over long stretches of the territory, overgrazing and fires in the past have
degraded the vegetation to garrigue.
On the mountains behind the Lucanian
Tyrrhenian coastline, from sea level to
about 300 metres, maquis with wild
olive and carob is particularly rich in
myrtle, a species preferring conditions
of relatively damp soil and atmospheric
humidity. The maquis here is mainly
composed of myrtle and many-flowered heath (Myrto-Ericetum multiflorae)
with lentisk, wild olive, and broadleaved phillyrea. At higher altitudes, it
becomes garrigue, in which many-flowered heath and myrtle are accompanied
by spiny cytisus (Chamaecytisus
spinescens), rosemary and winter
savory (Chamaecytiso subspinescentiEricetum multiflorae). Nearer the sea,
facing south, maquis with tree euphorbia and wild olive prevails. The northern
slope of the island of Dino (off the Calabrian coast) is occupied by a strip of
holm oak with terebinth, flowering ash and black bryony, while the windier
slopes have prevailing high maquis with broad-leaved phillyrea; the warmer
ones host maquis with wild olive and carob, with myrtle, lentisk and tree euphorbia. Instead, on the nearby island of Cirella, we only find maquis with myrtle,
lentisk, wild olive and tree euphorbia.
Compatibly with problems caused by over-grazing and fire, maquis is also found
along the Calabrian coastal chain and on the Serre chain. At sea level is maquis
with strawberry tree and tree heath, and Pulicaria odora, which at Capo Bonifati
is enriched by spiny cytisus and deciduous oak, lentisk and myrtle near Paola,
and cork oak around Pizzo Calabro. At higher altitudes, the maquis typically
contains deciduous vegetation (black hornbeam and flowering ash), together
with holm oak, myrtle, honeysuckle, sarsaparilla, asparagus and butcher’s
broom. The Aspromonte mountains host formations of high maquis developing
on limestone, with wild olive, lentisk, the oak Quercus virgiliana, a heat-loving
deciduous species, prickly juniper and asparagus (Oleo-Quercetum virgilianae),
widespread from sea level to about 500 m in the most southerly portion, which is
also the driest. The limestone rocks of the hilly belt on the Ionian side contain the
Aceri-monspessulani-Quercetum virgilianae association, with lesser maple,
holm oak, terebinth, madderwort, asparagus and butcher’s broom. On schist,
gneiss and granite, the maquis is composed of holm oak with tree heath, strawberry tree, wild olive, lentisk and spiny cytisus (Erico-Quercetum ilicis) or, more
often, from 200 to 1000 m along the Tyrrhenian side or more sporadically on the
Ionian one, of the same species with Sicilian germander (Teucrium siculum) (Teucrio siculi-Quercetum ilicis). Instead, high maquis with Quercus virgiliana, tree
heath and spiny cytisus are found on siliceous rocks from sea level to 900 m,
from 0 metres on the Tyrrhenian side and only above 400 m on the Ionian side,
which is hotter and drier (Erico-Quercetum virgilianae).
Up to 800 m, the Tyrrhenian side has maquis preferring siliceous soils with cork
oak, tree heath, Sicilian germander and spiny cytisus (Helleboro-Quercetum
suberis), whereas the Ionian side has types with Quercus frainetto, holm oak,
tree heath and broad-leaved phillyrea (Quercetum frainetto-ilicis). Heat-loving
maquis is represented by types with tree euphorbia, occurring everywhere on
rocky spurs, both along the coast and inland; by a kind of maquis with myrtle
and lentisk, only along the coast, in the driest parts between Brancaleone and
Capo dell’Armi; and by a few fragments with Phoenician juniper and wild olive,
and secondary silicaceous-loving maquis with predominant tree heath, prickly
juniper, on the Tyrrhenian side from 0 to 1000 m, or with myrtle lower down. The
Calabrian and Lucanian seasonal rivers have wide, pebbly beds with fine clumps
of maquis with oleander, tamarisk and chaste tree.
Maquis covers part of the cliffs along the Trieste coastline (Friuli Venezia Giulia)
The Adriatic coast. In Salento (the “heel” of Italy), in the few stretches of spontaneous vegetation, we find primary littoral maquis with juniper along the high
portions of the dune system, parallel to the coast, with prickly juniper, Phoenician juniper, Aleppo pine, prasium, lentisk and myrtle. This maquis is often
degraded to a type of garrigue with shrubby thyme and thorny burnet with
sage-leaved rockrose, or replaced by high or low secondary maquis with myrtle and prickly juniper. Further inland are holm oak with phillyrea, butcher’s
broom, cyclamen and honeysuckle.
The most degraded portions of the Apulian limestone coast, with rosemary and
Cretan rockrose (Cisto eriocephali-Rosmarinetum officinalis) are linked dynamically with maquis with many-flowered heath and narrow-leaved phillyrea
(Phillyreo angustifoliae-Ericetum multiflorae). Very widespread on the limestone
cliffs of the Gargano promontory (the “spur” of Italy) is a low type of maquis,
with wild olive and lentisk (Oleo-Lentiscetum var. adriaticum), Phoenician
juniper, prasium, Aleppo pine, myrtle, broad-leaved phillyrea, madderwort and
asparagus, modelled by the action of the wind. The Gargano promontory also
has a type of maquis with tree euphorbia and wild olive (Oleo-Euphorbietum
dendroidis), together with prasium, lentisk, buckthorn, madderwort and, of
course, wild olive in particularly hot locations. The sandy dunes host juniper
(Juniperetum macrocarpae-phoeniceae) in which, towards the sea, exposed
to salt-laden winds, large-berried juniper prevails, while Phoenician juniper
prevails on the opposite side. The most widespread formation at greater altitudes is holm oak and ash undergrowth (Fraxino orni-Queretum ilicis), with
lentisk, broad-leaved phillyrea, madderwort, asparagus and black bryony;
cooler slopes host holm oak, black hornbeam, flowering ash, butcher’s broom,
madderwort, asparagus and cyclamen (Ostryo-Quercetum ilicis).
The limestone arcipelago of the Tremiti Islands has the same geological and
vegetational features as the Gargano. The largest island, San Domino, has fine
woods of Aleppo pine, with maquis composed of wild olive, myrtle, rosemary
and rockrose. The island of San Nicola only has low maquis, with lentisk, tree
euphorbia, rosemary and rockrose. Northwards along the Adriatic coast from
the Gargano towards the Conero, the maquis is progressively impoverished in
heat-loving species. The hills along the Adriatic littoral host undergrowth of holm
oak and ash (Fraxino orni-Quercetum ilicis) with lentisk, buckthorn, prickly
juniper, sarsaparilla, asparagus and madderwort on the sunniest slopes. More
mesophilous environments have holm oak, and also prevailing black hornbeam,
laurustine, scorpion senna and ivy (Cephalantero-Quercetum ilicis). Naturally,
the boundary between evergreen and deciduous vegetation is gradual.
Between the Gargano and the Conero, maquis with holm oak and ash often has
a cover of Aleppo pine, degraded into
garrigue of the Cisto-Ericetalia type. In
some cases, e.g., along the Pescara littoral, strawberry tree, myrtle and wall
germander are missing, whereas there
are many mesophilous species.
There are also fragments of maquis in
the inner portions of the Appennines.
On south-facing slopes of the eastern
Maiella are small formations with a type
of daphne (Daphne sericea), shrubby
scorpion vetch, strawberry tree, box,
honeysuckle and scorpion senna.
On the hilly slopes of the Marches,
mainly north-facing, on deep soil, we
still find maquis, coppiced in the past,
with holm oak, flowering ash, scorpion
Cross-section of vegetal cover of the coastal
senna, Mediterranean honeysuckle,
part of the Conero promontory (Ancona).
sarsaparilla, asparagus, myrtle, strawOnly at a certain height above sea level does
slope configuration allow a small amount of
berry tree and pubescent oak (Fraxino
soil to develop, where low maquis formations
with shrubby scorpion vetch and diss grass
orni-Quercetum ilicis). On partially concan grow ( 1 ), often accompanied by Aleppo
solidated rocky debris, with no or very
pine ( 2 ); high holm oak maquis ( 3 ) can only
appear on the top part of the cliff, where soil is
little soil, a type of low maquis grows,
more abundant
with shrubby scorpion vetch and diss
grass (Coronillo valentinae-Ampelodesmetum mauritanicae), Spanish broom,
Aleppo pine, asparagus and madderwort. The best examples are found on the
Conero promontory, with tree euphorbia on rocky spurs near the sea and Osyris
alba in more wooded locations. The seaward-facing slopes of the Conero also
have thick maquis with prickly juniper and lentisk (Pistacio lentisci-Juniperetum
oxycedri), strawberry tree, buckthorn, holm oak, sarsaparilla, asparagus and
The same formations with holm oak and ash, but with fewer heat-loving plants,
may be seen inland (Gole di Frasassi, Gole di Pioraco, Valle Scappuccia), both
north as far as Pesaro and to Val Marecchia. Holm oak and ash undergrowth
also occurs on the Appennines along the Umbria-Marches boundary (M. Catria,
M. Subasio, Monteluco, Valnerina, Valle del Serra), but mainly on limestone and
in particular microclimatic situations. There is a certain amount of mixing
between evergreen and deciduous vegetation, and more mesophilous varieties
with woodland species (Cephalantero-Quercetum ilicis).
On inland Appennine reliefs (Valle del
Serra, Ferentillo, Pietracamela), on sunny slopes at 200-300 metres, are types
of maquis with scorpion senna and
emeroidis-Ericetum multiflorae), with
prickly juniper, Spanish broom, Aleppo
pine, broad-leaved phillyrea, holm oak,
terebinth, Cretan rockrose and madderwort.
At the Gole di Pioraco and on Monte
Nerone, at an altitude of slightly more
than 400 metres, is a type of pseudomaquis with smoke tree (Cotinus coggygria) and prickly juniper (Junipero
oxycedri-Cotinetum coggygriae) with
terebinth, box, flowering ash, holm oak,
scorpion senna and asparagus. Even
Mediterranean honeysuckle (Lonicera implexa)
more widespread, slightly lower down
(250-450 m) at the Gole di Pioraco, between Genga and Sassoferrato, Gola del
Furlo and Valle del Serra, is another type of pseudo-maquis with Osyris and
asparagus (Asparago acutifolii-Osyridetum albae), with prickly juniper, Cretan
rockrose, pubescent oak, ash and Spanish broom.
Moving now further north, the climatic situation and geographic morphology of
the Po Plain both mean that there are suddenly many changes before true
maquis is again found near Trieste.
The Bosco della Mesola, like the Bosco Nordio, contains holm oak, but the typical species of the holm oak association are not found in sufficient numbers to
define it as such, and so the corresponding maquis is also lacking. On Lake
Garda, in the Euganean Hills and in the Valle del Sarca, special micro-environments contain relicts of thermophilous vegetation, composed of holm oak, laurel, Judas tree, buckthorn, terebinth, nettle tree (Celtis australis), broad-leaved
phillyrea and tree heath. These formations were probably more extensive in the
past and, due to climatic changes, have remained near cliff environments.
Lastly, in the Carso (karst) area of the Gulf of Trieste, we again find true maquis.
The type found in the North Adriatic is special because, due to high rainfall, the
ratio between hard-leaved and malacophyllous plants (ones with thin, expanded leaves) increases in favour of the latter, but the permeability of karst soils
offsets the high rainfall. The main reason for the existence of maquis with holm
oak and black hornbeam (Ostryo-Quercetum ilicis) along the Trieste coastline is
its warmer and drier microclimate.
The high maquis found here is composed almost exclusively of holm oak,
broad-leaved phillyrea, terebinth, flowering ash, lesser maple, scorpion senna
and St. Lucie’s cherry (Prunus mahaleb), with abundant liana species like sarsaparilla, madderwort, clematis and asparagus. There are not many non-woody
species, because deep soil is lacking and there is little light. In certain cases,
only cyclamen (Cyclamen purpurascens) grows on a carpet of ivy.
The large islands. In Sicily, the various types of Mediterranean maquis play an
important role because this is the potential vegetation of almost the whole
island (excluding Etna, the mountain chains of Peloritani, Nebrodi, Madonie,
and other higher inland reliefs). It is greatly limited everywhere by land exploited
for agriculture and grazing, but there are some remarkable examples here and
there. At lower altitudes, between sea level and 300-500 metres, in the OleoCeratonion domain which occupies most of Sicily, wild olive, carob, broadleaved phillyrea, narrow-leaved phillyrea, buckthorn, Pyrus amygdaliformis,
terebinth, wild almond (Prunus amygdaliformis), dwarf palm, germander, tree
euphorbia, lentisk and prickly broom all prevail. Near Ragusa, many formations
contain thorny oak, accompanied by wild olive, carob, dwarf palm, broadleaved phillyrea, lentisk, prickly juniper, shrubby thyme and germander.
Between 300-500 and 800 metres, in the cooler belt of Quercion ilicis maquis, we
find prevailing tree heath with holm oak, prickly juniper, daphne, lentisk, myrtle,
sarsaparilla, Mediterranean honeysuckle, madderwort, asparagus and rockrose.
In the holm oak maquis, characterized by the occurrence of Sicilian germander,
we often find cork oak, exploited by man at the expense of holm oak for its bark,
butcher’s broom, cytisus, asparagus, sarsaparilla, madderwort and, at lower altitudes, heat-loving species like dwarf palm, broad-leaved phillyrea and lentisk.
The widespread exploitation undergone by Sicily since Roman times has
severely influenced its vegetal cover. Natural vegetation was replaced by cultivated cereals, fruit and vegetables, according to historical epoch. After the
abandon of agriculture, the maquis has repossessed ecologically suitable
areas, often with floristic and structural compositions similar to the original
ones but, more frequently, these recolonizing types of maquis are far from the
original vegetation with wild olive and carob or holm oak: species of the Rosmarino-Ericion domain now prevail, with rosemary, shrubby thyme, rockrose,
Thymelaea hirsuta and many-flowered heath, accompanied by germander and
dwarf palm. Completely different types, dominated by species associated with
at least temporarily well-watered soils, with oleander, chaste tree and tamerisk,
are found along the bed of the river Alcantara, beween Catania and Messina,
and other seasonal rivers of the western and northern coastline between
Taormina, Messina and Capo d’Orlando.
Garrigue with asphodel in Giara di Gesturi (Sardinia) in spring, with the famous wild horses
The same area of Giara di Gesturi in summer
In Sardinia, Phoenician juniper is the main component of belts of coastal undergrowth on rock. Along sandy littorals, this formation also contains large-berried
juniper. On the most consolidated dunes in the Nurra area is undergrowth with
large-berried juniper, asparagus, lentisk and madderwort. Schist and limestone
outcrops both have fine examples of dense maquis, sometimes 3-4 metres
high, mainly composed of Phoenician juniper accompanied by tree euphorbia.
The best pioneering formations are composed of dwarf palm, with gradual
additions of lentisk and, later, Phoenician juniper. After fires on coastal limestone, we find a type of maquis with lentisk and dwarf palm, like that found in
similar situations in Sicily. But destruction of forests on the plains has given rise
to a type of maquis with lentisk and whitethorn, with deciduous species like
Pyrus amygdaliformis. Deforestation by fire of the most heat-leaving formations
of holm oak gives rise to garrigue with rockrose, which may evolve into maquis
with lentisk and spiny broom and, in time, tree heath and strawberry tree. Some
very fine examples of all these formations may be admired at Capo Caccia.
In the Sinis area, we find maquis with lentisk, rosemary and many-flowered
heath, accompanied by the yellow cushions of Corsican broom (Genista corsica).
The dunes of the Buggerru-Portixeddu complex contain maquis with large-
Cross-section of coastal vegetation along north-western coastline of Sicily.
Formations with Echinophora spinosa ( 1 ) and Thymelaea hirsuta ( 2 ) occupy respectively areas nearer
the sea and those immediately inland; then maquis with dwarf palm becomes dominant ( 3 ) and, still
further inland, maquis with carob ( 4 ) (Oleo-Ceratonion). On high cliffs inland, vegetation is dominated
by diss grass ( 5 ), revealing the considerable influence of man on the area
berried juniper and lentisk, and, in areas protected from sea winds, peculiar
formations of Phoenician juniper and thorny oak. The toponym “Landiri marru”
refers to the local name for this variety of oak, which flourishes in the area.
Another type of maquis, deriving from the degradation of cork oak groves, is
found on rocky outcrops further inland.
In the Giara di Gesturi, a basaltic plateau located in the central-southern part of
Sardinia, high maquis with broad-leaved phillyrea and strawberry tree is accompanied by whitethorn in cooler areas, whereas lentisk and wild olive favour sunnier parts, and myrtle is found near the ponds locally known as “pauli”. In areas
subjected to fire in the past, rockrose and wild lavender prevail.
On the slopes of Monte Albo, near Siniscola, lower altitudes contain thermoxerophilous maquis with Phoenician juniper, bean trefoil (Anagyris foetida),
Thymelaea hirsuta, sea asparagus (Asparagus aphyllus), prasium, and, to 500
metres, maquis with buckthorn, narrow-leaved phillyrea, lentisk, wild olive and
spiny broom. Higher up, between 500 and 800 metres, are holm oak with broadleaved phillyrea, laurustine, butcher’s broom, madderwort and sarsaparilla.
The flat-topped island of Tavolara, dominating the Gulf of Olbia, and the island
of Cavoli, near Cagliari, have a little maquis rich in wild olive, lentisk, junipers
of various types, spiny broom and rosemary.
On the island of Sant’Antioco, off the south-west coast of Sardinia, the
juniper maquis is xerothermic, with Phoenician juniper, lentisk and dwarf
palm (Pistacio-Juniperetum macrocarpae), like that of Nurra where, at Cala
Gonone, on the eastern coast, there is more tree heath and the situation is
less xerothermic.
Low maquis also occurs with lentisk, wild olive, dwarf palm, narrow-leaved
phillyrea and spiny broom, without myrtle; maquis with holm oak, strawberry
tree, tree heath, lentisk, myrtle, Montpellier rockrose and phillyrea; and low
maquis with Genista ephedroides, rosemary, Montpellier rockrose, wild
lavender, tree heath and strawberry tree. On the island of San Pietro, near
Sant’Antioco, the original extensive cover of juniper is reduced to a type of low
maquis with lentisk, Phoenician juniper, narrow-leaved phillyrea, rosemary
and, after recent fires, Montpellier rockrose and low maquis with Genista
ephedroides, lentisk, myrtle and Phoenician juniper. With respect to the
maquis found in northern Sardinia, dwarf palm is not abundant, but much
more myrtle is found. Other types of maquis are composed of rosemary, green
heath, narrow-leaved phillyrea, strawberry tree and tree heath.
In the most degraded areas, a kind of mosaic of low maquis develops, with
yellow rockrose, rosemary and heaths of various kinds, or Corsican broom,
Montpellier rockrose, lentisk and rosemary.
■ Origin and history of Mediterranean fauna
The origin of Mediterranean fauna is lost in the intricate history of animal populations and of their co-evolution with ancient biomes, starting from the Mesozoic era, more than 200 million years ago. At the beginning of the Jurassic, the
ancient continent of Pangea split into two parts, separated by the Sea of
Tethys. The coastal areas of this sea underwent profound transformations in
their vegetation, and thus fauna, after the climatic change - from continental to
oceanic - which occurred as a result of the mitigating influence of this sea. In
the millennia which followed, along the shores of the Tethys, innumerable
events of speciation and colonization led to continual shifts in the compositions of the biotic communities. Finally, in relatively recent times, two largescale paleogeographic and paleo-ecological events influenced what were to
become the present-day populations: the salinity crisis of the Messinian, and
the glaciations of the Pleistocene.
At the end of the Miocene (Messinian), between 5.6 and 5 million years ago,
the Strait of Gibraltar closed, turning the Mediterranean into an inland sea. The
hot, arid climate of that period caused widespread evaporation and a considerable lowering in sea level, to the extent that the Mediterranean basin was
transformed into a series of enormous lakes surrounded by deserts or steppelike terrains. The latter were mainly colonized by animal and plant species
originally from the African continent and what we now call the Middle East.
During the period of maximum crisis, Corsica, Sardinia and Sicily were all
joined together with Italy and North Africa, making up a large land bridge
which allowed animals to wander from one part of the territory to another. Still
today, the Tyrrhenian maquis and the large islands of the Mediterranean contain faunistic components of North African origin, which were cut off when the
sea level rose after the re-opening of the "Pillars of Hercules".
During the Quaternary, a new series of climatic events influenced the structure
and dynamics of the various ecosystems over a period of almost two million
years. Cold (glacial) periods alternated with hot (interglacial) ones, producing
constant displacements of biomes from north to south and vice versa. During
Serpent eagle (Circaetus gallicus)
this alternation of paleoclimatic phases, the animals which had adapted to
cold or hot climates became extinct or migrated with their elected biome, giving rise to continual changes in both landscape and life forms. For example,
during the coldest periods, the tundra and its reindeer reached the shores of
the Mediterranean, whereas in the hottest ones, the Mediterranean maquis
and fallow deer were to be found as far north as Denmark. In addition, during
each climatic phase, the melting or formation of glaciers alternately freed or
imprisoned enormous quantities of water, respectively causing rising or lowering in sea level.
This series of marine transgressions or regressions, as they are called, continually modified the paleogeography of the Mediterranean, varying its coastline and thus the extent of continental and island masses. For the same reason, during the glacial periods, the coastal Mediterranean maquis of the Italian peninsula found itself part of that of the islands; vice versa, during the
interglacial periods, the same islands were smaller and farther away from the
mainland. Many animals were able to colonize the islands and expand along
the Italian coastal maquis during periods of marine regression, profiting by
the land bridges which emerged where the sea was shallowest. Also, the
closer the islands were to each other or to the coastline of the peninsula, the
greater the number of organisms which could colonize those areas, spreading actively or passively. Some species exploited the larger size or closeness
of emerging land by swimming across short stretches of sea. Others, transported by currents (for instance, on floating tree-trunks) were able to reach
nearby islands and beaches more easily. In this way, many species spread
throughout the evergreen maquis of the Mediterranean, from Andalusia to
During the glacial periods, the maquis only occupied a thin coastal strip, but
then extended far inland during the interglacial ones. Thus, animal species
linked to the maquis continually changed their ranges, which became smaller
or larger according to climatic phase.
Still today, we can find relict heat-loving species on particularly hot Appennine or Prealpine hilly slopes (generally south-facing) where fragments of
Mediterranean maquis are preserved in the middle of landscapes generally
characterized by more continental vegetation. This relict presence reveals the
once more extensive and continual distribution of these species during the
warmest periods.
In the last tens of thousands of years, the history of the Mediterranean maquis
has advanced together with the biological and cultural evolution of man, from
the Neanderthal hunter-gatherers to the polluters, speculators and animalists
of the New Age. The coastal environments all round Italy have suffered
increasing pressure due to the impact
of humans, not only due to deforestation, fire and grazing, but also to the
introduction of allochthonous species
like wild rabbit, fallow deer, spurthighed tortoise, marginated tortoise,
and Central European populations of
wild boar and hare. In addition, the
environment has been subjected to
the severe impacts of DDT, distributed
in enormous quantities in regions near
coastal marshes to curb malaria, and
of pesticides, widespread in all the
intensive crop-growing areas of the
fertile plains with which the fauna of
the Mediterranean maquis interacts
Wild rabbit (Oryctolagus cuniculus)
closely. Lastly, the most dramatic of all
impacts has arrived: speculation by the construction industry. Closeness to
the sea, the favourable climate and the beauty of the Italian coastal landscape have all given rise to a rush towards privatization of the territory, which
has been girded about and chopped up into small plots, enclosed with fencing and other obstacles to the path of animals; gardens are irrigated all year
round, impoverishing water resources in the subsoil; domestic and feral cats
prey on nesting birds and fledgelings, open-air waste dumps favour the
increase in numbers of anthropophile and opportunistic species like rats,
crows and seagulls; artificial bodies of water become infested with mosquitoes and have to be disinfected regularly, impoverishing other animal communities; artificial lighting of inhabited areas at night sows death among nocturnal insects, and so on.
To compensate all these negative aspects, it must be said that a greater naturalistic awareness and environmental vision of life is slowly but steadily
advancing - a vision which leads to initiatives favouring the conservation of
ecosystems. But the institution of protected areas is not enough to guarantee
the survival of vital, self-supporting populations of animals and plants. In order
to protect the Mediterranean maquis - and, indeed, all ecosystems in general
- two essential aims to be achieved are intelligent developmental planning and
sustainable exploitation of resources.
■ Animal communities
The fauna of the Mediterranean maquis is poor in animal species which are
exclusive to it. This applies to holm oak woodland, as well as to shrubby
maquis and garrigue.
Unlike the flora, which contains a considerable number of the hard-leaved
evergreen species so characteristic of coastal and subcoastal environments,
the fauna is mainly composed of organisms which are widespread from sea
level to the mountains. But the number of animal species closely associated
with hard-leaved evergreen formations is low in all taxonomic groups. This is
probably because, during the last glaciation, the Italian coastal vegetation
was reduced to small isolated fragments in the warmest areas of the peninsula and the islands.
In all probability, these fragments of maquis were too small to conserve vital
populations of animals, mainly terrestrial vertebrates, but also many insects
which require abundant trophic resources distributed over large areas.
The Würmian glaciation may thus have been responsible for one mass extinction, in which many species exclusive to this environment disappeared.
Maquis in northern Sardinia
The animals which colonized the environmental mosaic of the Mediterranean
vegetation mainly belong to the following categories:
● Woodland and shade-loving species living in deciduous oak woodland.
Several invertebrate species may be found in the litter of mature holm oak
groves, or are generally associated with oak (and thus also with the holm oak
and cork oak of the Mediterranean forest), or live on the trunks of trees of any
species. The latter include several vertebrates living on trees or in forests.
● Ecotonal species of shrubland and open spaces. Many species, all widespread, both vertebrates and invertebrates, without special needs obliging
them to live in particular environments, find favourable living conditions in all
parts of the evergreen maquis.
● Species of steppe origin. Many animal species living in the open maquis, garrigue, and spaces rendered arid by the destruction of the vegetation, colonized
Italy in relatively recent times from the Eurasian steppes. They found favourable
living conditions in these environments, which have been altered by man.
● Species of subdesertic origin. Several species found in the garrigue and
arid meadows of southern Italy, resulting from the extreme degradation of the
Mediterranean maquis, are very widespread in subdesertic portions of North
Africa and the Near East. They too colonized Italy in historical times, gradually
spreading as man modified the environment.
Good competitive skills are one of the characteristics necessary for success in
the Mediterranean maquis. This is because the conditions of this type of environment are quite favourable to animal life: winters are never very cold and
summers never too hot, because evergreen vegetation always supplies shade
and helps animals to regulate their body temperature. Clearly, these ideal conditions lead to intense competition among animals, most of which find it
advantageous to live in this environment.
The only problem is the relative scarcity of edible fruit and appetizing flowers.
However, the autumn flowering and fructification of several species, such as
holm oak, myrtle, juniper and phillyrea, represent an advantage for many animals which, as in the Maremma of Tuscany, first consume the resources of the
deciduous woodland and then move into the evergreen maquis. This is habitual for several species of large mammals, but above all for birds, which spend
the winter here, exploiting the mild climate. Insects too, even in mid-winter,
can find shelter from predators in the evergreen vegetation.
The secret of the relationships between the Mediterranean maquis and deciduous forest lie precisely in these seasonal dynamics which, partly by means
of the mosaic of ecotones and cultivated land, represent a continual ecosystem characterized by great biological diversity.
■ lnvertebrates
Paratriodonta romana mating on rockrose
Molluscs, nematodes and annelids. In spite of the hot-dry climatic conditions so typical of Mediterranean environments during the summer months,
many species of terrestrial gastropod molluscs (snails and slugs) live in the
maquis or garrigue, where they play a role as primary consumers, thanks to
their special adaptations.
Both during summer drought and winter cold, pulmonate gastropods with
shells (snails) withdraw into their "houses" and secrete mucus which, solidifying, creates a sort of "front door" (epiphragm) which thus allows them to close
out the external world. In this way, the animals minimize loss of water from tissues, and simply wait for better times. Some species pass both hot and cold
months in micro-environments with favourable conditions of humidity and temperature, e.g., in cracks between rocks or under stones. Others directly challenge the heat and dryness of the Mediterranean summer by attaching themselves to the stems of plants, to avoid contact with the overheated ground.
Instead, pulmonates without shells (slugs) can only take refuge right inside
damp, sheltered cavities. There are also species of terrestrial prosobranch gastropods which close the entrance to their shells with a special structure - not
provisional, like the epiphragm, but permanent. This operculum, as it is called,
is a sort of tiny calcareous shield or lid, which the animal produces from its covering epithelium and which is attached to the dorsal region of the back part of
the mollusc's foot. Pulmonate gastropods are hermaphroditic, but the terrestrial prosobranchs (related to the more common marine shell-bearing gastropods) have separate sexes. The most widespread is Pomatias elegans,
typically found in shady environments.
Among the pulmonates, the clausiliid
family includes several species endemic to southern Italy, all provided with
very long spindle-shaped shells. Two
members of this family are Papillifera
papillaris and P. solida, which prefer
maquis on limestone rocks and resist
drought well. The helicid family contains many species with more or less
spherical shells, some of which have
always been used as food by rural populations. Typical Mediterranean species
Eobania vermiculata
are Eobania vermiculata, Cantareus

The Mediterranean maquis 6