I TA L I A N H A B I TAT S
Arid meadows
12
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
I TA L I A N H A B I TAT S
Scientific coordinators
Alessandro Minelli · Sandro Ruffo · Fabio Stoch
Editorial committee
Aldo Cosentino · Alessandro La Posta · Carlo Morandini · Giuseppe Muscio
"Arid meadows · Grassland in near-drought conditions"
edited by Alessandro Minelli
Texts
Lucio Bonato · Alessandra Di Turi · Alessandro Fontana · Alessandro Minelli · Simonetta Peccenini ·
Margherita Solari
In collaboration with
Paolo Giordani · Paolo Maria Guarrera · Paola Sergo · Mirca Zotti
English translation
Elena Calandruccio · Gabriel Walton
Illustrations
Roberto Zanella
Arid meadows
Graphic design
Furio Colman
Grassland in near-drought conditions
Photographs
Nevio Agostini 8, 16, 52 · Archive Museo Friulano di Storia Naturale (Tomasi) 39/2, 45, 46, 47 ·
Claudio Aristarchi 9, 10, 23, 34, 66, 68/1, 69/2, 70/1, 134 · ARPA Regione Veneto 12 ·
Pietro Baccino 27, 28/2, 29, 31, 37, 38, 39/1, 40/1, 41, 43, 54, 55, 63/2, 63/3, 64, 68/2, 69/1, 70/2, 137 ·
Giuseppina Barberis 63/1 · Stefano Bossi 93, 94, 95 · Eugenio Busetto 7, 35, 61, 132 ·
Gianfranco Canderan 103, 138 · Giuseppe Carpaneto 124 · Andrea Dall’Asta 111, 116 ·
Adalberto D'Andrea 140 · Giuseppe Delitala 126 · Vitantonio Dell’Orto 6, 21, 24, 44, 50, 57, 72, 73, 74,
77, 80, 102, 104, 106, 118, 120 · Dario Ersetti 11, 81, 97 · Tiziano Fiorenza 96, 110, 119 ·
Paolo Fontana 82, 85/2 · Luca Gardini 67 · Maria Manuela Giovannelli 78 · Gianluca Governatori 48 ·
Luca Lapini 114 · Giuliano Mainardis 83, 84, 85/1 · Chiara Mazzanti 146 · Ugo Mellone 22, 25, 71, 122,
130, 143 · Michele Mendi 101, 108, 113 · Francesco Orsino 26, 32, 40/2, 42, 53, 59, 60 ·
Roberto Parodi 19, 49, 78, 99, 100, 105, 107, 109, 112, 115, 117, 121, 123, 125 ·
Simonetta Peccenini 36, 58 · Roberto Pizzutti 139 · Giovanni Salamanna 30 · Mauro Sanna 127, 128 ·
Sonja Siljak-Yakovlev 28/1 · Ermido Traverso 51 · Marco Uliana 75/2, 75/3, 76, 86, 88, 91 ·
Augusto Vigna Taglianti 75/1, 79, 85/3, 87 · Roberto Zucchini 18, 148
© 2005 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 23 9
ISSN 1724-6539
Cover photo: Arid meadows near Capo d’Otrano, Apulia (photo by U. Mellone)
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
Italian habitats
Contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Alessandro Minelli · Simonetta Peccenini
Geography, climate and pedology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Alessandro Fontana
1
Caves and
karstic
phenomena
2
Springs and
spring
watercourses
3
Woodlands
of the Po
Plain
4
Sand dunes
and beaches
5
Mountain
streams
Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Simonetta Peccenini · Alessandra Di Turi
Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Alessandro Minelli
Vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Lucio Bonato
6
The
Mediterranean
maquis
7
Sea cliffs and
rocky
coastlines
8
Brackish
coastal lakes
9
Mountain
peat-bogs
10
Realms of
snow and ice
Conservation and management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Lucio Bonato · Alessandra Di Turi · Simonetta Peccenini
Suggestions for teaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Margherita Solari
Select bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
11
Pools,
ponds and
marshes
12
Arid
meadows
13
Rocky slopes
and screes
14
High-altitude
lakes
15
Beech
forests of the
Apennines
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Introduction
ALESSANDRO MINELLI · SIMONETTA PECCENINI
Totally or almost totally grassy covers
are called meadows or pastures.
Differences in their floristic composition
- seldom visible to casual observers identify various types of grasslands,
which may either differ in origin or have
been altered by man’s presence over
the years.
As regards their origin, meadows are
divided into primary and secondary,
the latter being those from which man
has removed all plants with woody
habit.
Permanent meadows are grassy
coenoses which have been artificially
planted and require fertilization or
irrigation to produce forage crops, but
Trasformation of a meadow to a sparse
woodland in the Ligurian Alps
they are not treated in this volume.
Pasturelands, instead, are natural
grasslands that are not exploited by man and are used exclusively by grazing
livestock. There is also a transitional stage of meadow-pasture, called lea,
which is mowed in spring-summer and used as grazing land in the remaining
part of the year. This type of plant cover is not only composed of sown
species, but is also colonised by spontaneous grasses that contribute to the
formation of partially natural environments. Pastures and leas may compose
arid meadows, on which grow a diversity of herbaceous plants sharing
adaptations to the lack of water, or at least lengthy periods without usable
water. There are two main adaptations to survival in these particular
conditions: temporal and morphological.
Extreme temporal adaptations regard short-lived annual grasses, which
prevail when water becomes available even only for short periods during the
year. These annual species, called therophytes, are typically found in
Pastures in Basilicata
7
8
Mediterranean meadows: they are in seed form most of the time, and bud,
grow, bloom and fructify in a few weeks, perhaps a few months, exploiting the
short rainy season. In the Mediterranean area, this season is generally spring,
although some species may, in the course of the same year, have a second
life-cycle associated with autumn rain.
Morphological adaptations aim at limiting transpiration through the leaves,
which may either be very small or covered by thick hairs. Woolly hairs
covering stems and leaves also protect plants from intense sunlight which, in
addition to increasing evaporation, may also damage plant tissues with its
ultraviolet rays.
Meadows should also be considered as transitional environments between
woodlands and deserts. Italy only features temperate woodlands, whereas
tropical zones, where temperatures are high throughout the year and rain is
abundant, have savannahs. Temperate prairies are generally found inland on
continents, far from the influence of damp sea winds blowing landwards, in
areas with cold winters, warm summers, and little rainfall during the year. In
Italy, therefore, they should be limited to inland Alpine and Apennine areas.
But Italian meadows are in fact far more extensive, because they often
developed - as secondary meadows - in areas once covered by forests,
which man has recently destroyed.
Meadow used for grazing (Santa Sofia, Romagna Apennines)
9
Shrubby arid meadow (Liguria)
Geography, climate and pedology
ALESSANDRO FONTANA
■ General features
There are arid meadows in many Italian
areas and, although varying in extent,
they are found almost everywhere,
along the Alps, in the Po Plain, along
the spine of the peninsula, in the
Apennines and on the islands. These
grassy covers are seldom restricted to
particular geological areas or specific
and easily identified geographicclimatic contexts, as is the case with
beaches, grottoes, or high mountains.
Areas with arid meadows may show
extremely different climatic, geological
and morphological characteristics, but
they all share very dry conditions, not
Cracked substrates with poor soil favour
leaching and therefore arid conditions
only due to the lack of rainfall.
The availability of water is also
influenced by various factors associated with the substrate. In addition to the
lithological and chemical composition of soil, also the size, shape, and
distribution of landscape elements are important.
■ Lithological characteristics
Most arid meadows are found in areas with outcropping calcareous or
dolomitic rocks, with no surface water flow, and presenting karstic phenomena.
However, there are many arid meadows on chalk, clay, marl and, in the centralwestern Alps, even on granite and metamorphic rocks like schist and
serpentinite. Chalk is mainly found in the Emilia-Romagna Apennines and in
Sicily, and the clayey and marly-sandy areas in the central-northern
Apennines, along the southern Italian coast and on the large islands of Sicily
Autumn flowering (Urginea) in an Apulian arid meadow
11
12
and Sardinia. Arid meadows sometimes develop in recent alluvial or flank
deposits, such as screes, moraine bars, landslide mounds, riverbanks, and
alluvial and detritic fans.
These are all highly permeable substrates through which water trickles quickly
underground, and in southern Italy and on the main Italian islands, where they
are typical habitats, arid meadows are even found on volcanic lava and
granite.
■ Pedology: arid soils and meadows
Areas with arid meadows generally
have little, degraded, or totally absent
soil. In them, their inaccessible position,
or the lack of nutrients in the soil,
sometimes caused by climatic factors,
hinder the development of woodland,
which eventually disappears or
becomes rarefied.
In these conditions, substrates cannot
hold moisture, especially along
southern flanks.
Areas with outcropping rocks may
contain soil that is produced over
extremely lengthy periods and, if
eroded, is unlikely to form again.
Stratified scree slope composed of very
permeable gravel and sand, with thin soil (Monti
For instance, the chemical composition
Lessini, Veneto)
of limestone hinders pedo-genetic
processes, and its karstic dissolution produces clay residues rich in iron
compounds, which are infertile. This is the so-called red soil, typically found in
Mediterranean karstic environments, particularly all round the southern Italian
coast from Gaeta (Latium, south of Rome) to the Gargano Peninsula (the “spur”
on the “boot” of Italy), on the Adriatic, and the main islands.
Human activities, such as agriculture and the grazing of livestock, have often
accelerated the development of arid meadows by turning woodlands into
sometimes permanently barren land. Deforestation, intensive grazing and
repeated fires have facilitated flank impoverishment and soil erosion in many
areas.
Elsewhere, extensive cultivation of certain plants on unsuitable land has
impoverished the soil.
Arid meadows and glaciations
In central-northern Italy, the present
distribution of arid meadows was
directly or indirectly influenced by the
last glaciation affecting the Alps. This
Ice Age, called the Würm, occurred
between 110,000 and 11,500 years
ago, when the average temperature of
the Earth had fallen by a few degrees.
The resulting enormous glaciers
covered the Alpine valleys and
reached the Po Plain, where the debris
they carried created morainic cirques.
This epoch entered a particularly cold
stage between 25,000 and 18,000
years ago, and ended 11,500 years
ago, when climatic conditions became
similar to present-day ones.
That period marks the end of the
Pleistocene and the beginning of the
Holocene.
The formation of glaciers in the Alps
and in part of the Apennines caused
vegetation to disappear over
extensive areas.
Alessandro Fontana
Several plant species found refuge in
a few mountain chains devoid of
glaciers, especially in the Pre-Alps
and, when the glaciation ended, they
were finally able to migrate to more
suitable areas and colonise locations
the glaciers had abandoned.
Therefore, although present-day arid
meadows are still found in “refuge
areas” of the glacial epoch, others
have developed inland along the Alps,
in ecological “corridors” which have
favoured the penetration of
thermophilous species.
The situation was quite different in
southern Italy and its islands, where
glaciation was restricted or totally
absent. In the Apennines, glaciers
were only present inland at high
altitudes, e.g., along the Emilian ridge,
on the highest massifs of the Italian
peninsula, and in Calabria. The only
glacier still surviving is the Calderone
on the Gran Sasso.
Greatest expansion of extensive Alpine glaciation during the Würmian climax
13
15
14
TN
TN
AO
MI
VE
AO
TS
TO
MI
TS
VE
TO
BO
GE
GE
BO
FI
FI
AN
AN
PG
PG
AQ
AQ
ROMA
ROMA
CB
CB
BA
NA
BA
NA
PZ
CA
PZ
CA
RC
RC
PA
Less than 0°C to 5°C
PA
Less than 500 mm to 600 mm
Between 5°C and 10°C
Between 14°C and 18°C
Between 600 mm and 900 mm
Between 1500 mm and 3000 mm
Between 10°C and 14°C
Exceeding 18°C
Between 900 mm and 1500 mm
Exceeding 3000 mm
Map of average annual temperatures in Italy
Map of average annual rainfall in Italy
16
■ Climatic features: differences between northern and central-southern Italy
As Italy is an extremely long peninsula, its northern areas have a cold,
subcontinental, temperate climate, which changes to warm and subtropical typically Mediterranean - in the south. Changes in latitude are implemented by
the peculiar effects caused by the complex orographic features of Italy - the
mountain chains of the Alps and the Apennines - which give rise to significant
variations. This is why Mediterranean characteristics become less marked as we
proceed inland and gain altitude. The boundary between Mediterranean and
temperate areas runs along the northern Apennines, thus leaving all northern
Italy, except for Liguria, in the subcontinental region. Rainfall in areas south of
the southern boundary of the Po Plain is also called Mediterranean, with
maximum rainfall only for a few months in winter (November-April). In southern
Italy, where drought and intense summer sunlight coincide, giving rise to lengthy
periods of water shortage, arid meadows are common and extensive.
In northern Italy, rainfall is abundant and distributed throughout the year, with
a winter minimum, a main peak in spring, and another in autumn. Only a few
areas with particular geographic conditions have less rainfall than the
surrounding regions - for instance, some Alpine valleys. The distribution of arid
meadows is therefore restricted, and associated with very permeable soil.
17
TN
AO
MI
TS
VE
TO
BO
GE
FI
AN
PG
AQ
ROMA
CB
NA
BA
PZ
CA
RC
PA
Temperate subtropical
Eroded areas in Tuscan-Emilian Apennines calanches (“badlands”)
Temperate warm
Temperate cool
Temperate coastal
Temperate cold
Temperate subcontinental
Cold - freezing
Division of Italy into climatic areas according to temperature
18
■ Main arid meadows in Italy
Po-Alpine area. Arid meadows are not continual, but fragmented.
● Alpine-steppe valleys: in many valleys oriented east-west, parallel to the
Alps, damp sea air hardly ever penetrates, giving rise to a continental climate,
with little rainfall, between 600 and 1100 mm/year. Examples are Valle di
Susa, Valle d’Aosta, Valtellina, Val Venosta, Val Camonica and Val Pusteria.
Arid meadows include windbeaten pastures and flanks up to 1500-2800 m.
Soil is carboniferous in the eastern and central Alps, and metamorphic in the
western Alps.
● Pre-Alpine zone: arid meadows cover many flanks overlooking the Po Plain
and the Veneto-Friuli region. Rainfall is abundant (1400-2000 mm/year) and
the xerophilous habitat is associated with calcareous and dolomitic substrates
affected by karstic phenomena, e.g., the Vallese and Lombard Pre-Alps, and
areas surrounding the large Lombard lakes, such as Garda, Maggiore, Como,
Iseo and Orta. These lakes make the climate milder, and thermophilous
species can live on the hills of the large morainic cirques in the plains between
Piedmont and the Veneto. In the Veneto Pre-Alps, there are arid meadows on
the Grappa chain, the hills surrounding Asolo, along the southern Altopiano
del Cansiglio and the Sette Comuni. Particular cases are the volcanic
Euganean and Berici Hills, which rise, isolated, to a maximum of 600 m in the
plain but are less than 30 km from the coast.
● Italian karst (NE): this is an area showing typical karstic phenomena, with an
average altitude of 300 m, stretching as far as the Adriatic coast, where the
Mediterranean climate is cold. Eastwards, the continental influence affects
both climate and living species.
● Po and Veneto-Friuli Plain: at the foot of the Alps, the gravelly and coarse
sandy sediments forming the high plain are particularly permeable, and natural
hydrography is almost absent. In the few uncultivated areas that have been
deforested, there are arid meadows, e.g., along the river Serio, and in dry
moorland on terraces along the river Ticino. Recent cultivation has eliminated
meadows from Pleistocene river dunes (called sabbioni) in the Lomellina
(Lombardy). In Friuli, the watercourses Cellina and Meduna have formed two
wide alluvial cones in which, despite 1700 mm rainfall a year, large particlesized gravel and its totally carbonatic composition, impoverish the soil greatly.
The word magredi (thin) indicates the poor, untilled land of this area,
Pastures in Julian Pre-Alps (Friuli Venezia Giulia)
Magredi in Cordenons, in alluvial fan of Cellina-Meduna (Friuli Venezia Giulia)
The distribution and extent of arid meadows along the Italian peninsula fall
into three main areas: Po-Alpine, the Apennines, and the south including the
islands.
19
20
particularly along the beds of these two watercourses, which are 2-5 km wide
and lack surface water as far as the karstic springs. A similar, although less
extensive condition, is typical along the banks of many large rivers, such as
the Brenta, Piave, Tagliamento and Isonzo in the Veneto-Friuli Plain, and the
Scrivia and the major Lombard rivers in the Po Plain.
● Langhe and Monferrato: these hills have an average altitude of 600 m and
are the northernmost spurs of the Ligurian Apennines. They are composed of
limestone alternating with eroded clay, sandstone and sand. In some areas,
rainfall does not exceed 800 mm/year and arid meadows may be found near
passes, usually called “Bric delle forche”.
Apennine area. This volume treats areas that typically contain arid
meadows, bearing in mind that they are widely distributed along the Italian
peninsula and that, proceeding southwards, the climate gradually becomes
more arid.
● Emilia-Romagna and Umbria-Tuscan Apennines: these mountains have
clayey and sandy soils, sometimes containing chalk. Such substrates often
give rise to landslides and erosion furrows, which cause the removal of soil, as
in Oltrepo’ Pavese (Pavia), near the Calanchi dell’Abbadessa (Bologna), in Val
Marecchia, and several areas of the Emilian Apennines. There are arid
meadows near the Gessi Bolognesi, which also show karstic phenomena and
where the climate is dry. On the Romagna flank and in the Apuan Alps, rainfall
reaches 2000 mm, due to the vicinity of the sea, but is much less in centralsouthern Tuscany. Here are the typical meadows around Siena and Volterra,
with eroded soil, gravelly riverbanks, and the so-called crete and biancane,
i.e., clay reliefs moulded by run-off, the surfaces of which are covered with a
whitish salt efflorescence.
● Mountains of the central-southern Apennines: from Umbria to Calabria, the
largest mountain chains (Vettore, Terminillo, Gran Sasso, Maiella, Velino,
Sibillini, Meta, Vulture, Volturno, Sirino and Pollino) have arid meadows at
medium-high altitudes, where the climate is temperate and sometimes Alpine.
Except for the Vulture, a now inactive volcanic chain, the rocky substrate
always contains limestone and dolomitic karstic rocks. Southwards, rain falls
heavily on the mountains, preventing damp sea air from reaching the valleys
inland, which are therefore very dry: examples are the Piana dell’Aquila and
Piana del Fucino. South of the carbonatic chain of the Pollino mountains, less
permeable magmatic and metamorphic rocks outcrop, enabling the flow of
surface water. There are arid meadows in several areas of the Sila and
Aspromonte mountains, even on schist and gneiss.
Southern Italy and islands. Here there are many, sometimes extensive arid
meadows. Although they are associated with the Mediterranean climate, they
are often the result of man’s degradation of original woodland. Rainfall is
concentrated on the Tyrrhenian flank of the mountains, whereas the Adriatic
side is notably more arid.
● Southern coast: all round the coast from Gaeta (Latium, Tyrrhenian) to the
Gargano (Apulia, Adriatic) and on the main islands, summers are prolonged
and dry, and arid meadows are found wherever rocks outcrop, especially if
flanks are calcareous and steep. This association is also typical of clayey
areas, generally composed of red soil, and sometimes of Pleistocene alluvial
or marine deposits, as in the Ionian Plain and in a few areas in Calabria and
Apulia.
● Lucanian eroded soil and dolomites: extensive areas in Basilicata are
composed of clay, marl and sand with eroded soil, and arid meadows are
frequent. The most typical areas are those of Tursi, between the rivers Sinni
and Agri. Inland, there are arid meadows near the so-called Dolomiti Lucane
(Dolomites of Pietrapertosa and Castelmezzano), which are made of
sandstone.
● Areas near the Gargano and the Salento Peninsula and Murge (the
“heel”): these constitute most of Apulia and are well-known for their karstic
Eroded areas with arid meadows in Basilicata; flat surfaces are farmed calanches (“badlands”)
21
landscape. There is very little rainfall - southwest of the Gargano it is less
than 500 mm/year. Extreme aridity favours a steppe climate, even in
uncultivated areas of the Tavoliere (table mountains) and the Bradanica
(between Apulia and Basilicata), made up of Pleistocene alluvial and marine
deposits.
● Sicily: arid meadows are everywhere, especially on the smaller Aeolian
islands and along the southern coast, where the climate is subtropicaltemperate, with winds blowing from the Sahara. Restricted rainfall
concentrates in a few areas, and there may even be seasons of total drought.
Except for the summit areas of Mount Etna, the average annual temperature
never drops below 11°C along the coast and 9°C inland. On the volcano,
conditions are similar to those of high-altitude deserts, due to the permeability
of lava and lapilli.
● Sardinia: aridity is exceptionally high, with limited rainfall subject to
variations over the seasons. Near the Campidano plain, rainfall never exceeds
500 mm/year. Along the southern coast, the climate is subtropical
Mediterranean, and temperatures never drop below 10°C anywhere. The soil
is peculiar, being composed of granite and lava. A few small areas have a
calcareous substrate and clay sediments that sometimes host thermophilous
species, as in the Piana di Ozieri, in the north.
22
Flowers in a meadow in Salento (Apulia)
Animals grazing on limestone on Monte Albo (Sardinia)
23
Vegetation
SIMONETTA PECCENINI · ALESSANDRA DI TURI
■ Primary and secondary meadows
Large areas above the tree-line, the
extent of which is determined by
sunlight and soil, are covered by
Alpine prairie, i.e., hosting types of
spontaneous grasses, sometimes
impoverished and discontinuous,
stretching as far as the upper
vegetation line (3000-3100 m).
These are the so-called high-altitude
primary grasslands. They are scarcely
Karstic area used for grazing (Apulia)
affected by man and are therefore
more constant in extent and richer in
scented plants and colourful flowers. In the Alps, Apennines and the larger
Italian islands (Sicily and Sardinia), they are found on the highest summits,
interrupted by ravines and rocky slopes, or at their highest possible limit,
where they are replaced by high-Alpine and discontinuous, pioneer snow
vegetation.
Sometimes there are meadows even where there should be woodland: these
are called secondary because they are not natural, but the result of
deforestation at various altitudes. They would certainly be re-colonised by
woodland if continual grazing and recurrent mowing did not prevent the
development of new plantules. In areas where grazing has ceased or has been
forbidden for reasons of environmental protection, woods spontaneously
close in glades, thus re-conquering spaces that had previously been taken
from them.
Man’s presence on leas and pastures has been and continues to be a sort of
ecological factor, selecting the species living in these environments. Flora in
these herbaceous formations is very similar, even in areas far apart. Arid
meadows are mostly composed of various grasses, e.g., Sesleria, Festuca
and Nardus.
Orchid Ophrys fuciflora
25
26
■ History of the populations of arid mountain and submountain Alpine
meadows
At the end of the last Ice Age, about 10,000 years ago, large numbers of plants
and animals migrated towards south-western Europe. In that dry, cold period,
European flora and fauna mixed thoroughly. Alpine valleys were covered by
thick ice from which rocks emerged, hosting impoverished and discontinuous
vegetation similar to that of the present high-Alpine and snowline plants in the
Alps. In the plains, the vegetation was typically that of cold steppes, and
developed as soon as the glaciers began to retreat.
This north-eastern vegetation, mainly composed of grasses, included joint
pine (Ephedra distachya), sea buckthorn (Hippophae rhamnoides), an
excellent pioneer plant of barren land, and wormwood.
The westward post-glacial migration of these plants was associated with
alluvial soil and morainic cirques exposed to the sun, below the line of the
retreating glaciers. Vegetation could therefore colonise these areas during the
final stages of glaciation.
The advance of the steppe-like component was arrested by a radical change
in the climate, which became more and more oceanic, and led to the formation
of deciduous and silver fir woodland.
All around the Alps, heliophilous steppe species were totally eliminated by
woodland, which prevented them
from migrating inland, towards the
Alps, where they are now relicts.
The milder climate, impossible for
many steppe species, favoured the
northward migration of Mediterranean
elements.
It was probably in this period that subMediterranean and Mediterranean
mountain species crossed the Alpine
passes to colonise inland areas north
of the Alps. Later, climate variations
decreased in intensity, but man’s
presence increased. Man has totally
changed the look of the landscape by
deforestation, actively contributing to
the development of steppe-like
Joint pine (Ephedra distachya)
vegetation.
■ Meadows in the Alps and Pre-Alps
In the Alps, from the vegetational and ecological viewpoints, there are four
types of arid meadows, which correspond to four different phyto-sociological
types:
● Sub-Alpine xeric and acidophilous Alpine grassland (Caricetalia)
● Sub-Alpine xeric and calciphilous Alpine grassland (Seslerietalia)
● Continental grassland (Festucetalia vallesiacae)
● Mediterranean sub-mountain and mountain xero-thermophilous grassland
(Brometalia)
Obviously, the boundaries between the various types are not clearly marked,
but the first two generally prevail at high altitudes.
Acidophilous Alpine grassland (Caricetalia). The flanks of siliceous
mountains, up to 2800 m, are covered by more or less dense herbaceous
vegetation, which constitutes Alpine acidophilous grassland. Pioneer grasses
often colonise areas near landslides and invade siliceous rocky slopes,
beginning from open turf and developing into continuous grassy cover.
Mature grassland stages are composed of more complex vegetal groups,
which give rise to proper grasslands: typical examples are fescue covers with
Haller’s fescue (Festuca halleri), and a plant association (Curvuletum)
Snowline vegetation in the Cottian Alps, with hummocks of Silene
27
29
28
Alpine chrysanthemum (Leucanthemopsis alpina)
Moss campion (Silene acaulis)
dominated by the sedge Carex curvula. Associations with Haller’s fescue are
the early colonising stages on detritic soil derived from the erosion of siliceous
rock. These are extremely stable grasslands, not particularly xeric, with
fragmented groups along the Italian side of the Alps, and extensive ones on
the high mountains of Valtellina and Val Chiavenna.
In favourable conditions, Haller’s fescue, a slender grass with thin, violetbrownish spikes, associates with rock bent (Agrostis rupestris), rampion
(Phyteuma hemisphaericum), large-flowered cinquefoil (Potentilla grandiflora),
Alpine cat’s ear (Hypochoeris uniflora) and various types of hawkweed
(Hieracium sp. pl.). These associations are typical of siliceous mountains, but
they may also be found on calcareous ones, if the soil is neutral or slightly
acid. In addition to Haller’s fescue, these grasslands contain shrubby forage
grasses, as well as others with no food value.
Grasslands with Haller’s fescue are very stable associations, the best for the
high-altitude valleys of the Braulio (South Tyrol). They develop from natural
pioneer associations on siliceous rocky slopes, or from areas which, in the
past, were deprived of scrub. In this case, they give rise to secondary
meadows.
The other closed grasslands found higher up on siliceous mountains are
sedge fields or associations with a typical Alpine sedge (Carex curvula). More
extensive and xerophilous than Haller’s fescue grasslands, they carpet with
ochre-coloured curved sedge the highest mountain tops dried by the winds,
contrasting with the bright green of fescue fields below. Curved sedge
grasslands are associations found from the Alps to the snowline, between
2200 and 2800 m, but may reach the snowline at about 3000 m.
Eyebright (Euphrasia minima)
White groundsel (Senecio incanus)
These meadows, which develop on very acid soil in the central Alps, are
climax associations, i.e., they are, in the locally prevailing conditions, the most
stable vegetation, towards which all the other plant associations of the upper
Alps evolve.
On the Rhaetian Alps, curved sedge, with its typical leaves which yellow
precociously, associates with a grass species (Oreochloa disticha) with small
blue-green spikes, Avenula versicolor, a beautiful oat-grass with glossy spikes
shimmering from violet to golden yellow, rock bent, and dwarf blue speedwell
(Veronica bellidioides). Sedge covers brighten up with the flowers of rampion,
primrose, Alpine chrysanthemum (Leucanthemopsis alpina var. alpina), moss
campion (Silene acaulis), gentian (Gentiana alpina, G. punctata), Alpine
dandelion (Taraxacum alpinum) and eyebright.
The competitive capacity of curved sedge is great and only partially hindered
by some grasses or other plants particularly adapted to the Alpine climate.
Pastures may efficiently thwart the expansion of curved sedge due to the
effects of grazing, which interrupts the grassy cover and favours colonisation
by species which are less competitive but which have high food value.
However, excessive grazing favours the development of mat grass (Nardus
stricta), which has no food value.
Sedge fields are extensive on the Spluga, between 2300 and 2600 m, and in
Valle d’Aosta, where they are found on both flanks from 2400 m. In Valle di
Susa, they are less frequent, although present between 2500 and 2800 m,
along gently sloping flanks. They contain large numbers of species, such as
curved sandwort (Minuartia recurva), curved sedge, glandulous hawkweed
(Hieracium glanduliferum) and white groundsel (Senecio incanus).
30
Lichens in arid meadows
Paolo Giordani
Lichens are important elements of
biodiversity in arid meadows. In these
habitats, terrestrial lichens and
mosses have important ecological
functions, such as regulating water
flow, and they sometimes also reveal
interesting adaptive solutions.
For instance, some lichens grow on
mosses or wood and decaying leaves:
in dry habitats, these niches remain
damp for lengthier periods, enabling
the lichen to supply itself with the
water necessary for photosynthesis.
Many lichen species are symbionts
with cyanobacteria, tiny organisms
which can capture nitrogen directly
from the air. One of the most
widespread is Leptogium lichenoides,
a leafy lichen with grey thallus and
papery texture. It lives between
terrestrial mosses in meadows along
the Tyrrhenian hills.
The same environment also hosts
species of the genus Collema, which
jellify with the rain or dew.
Mediterranean arid meadows, both
along the coast and in the hills,
usually host several species of
Squamarina, like S. cartilaginea, a
type of lichen that grows in limestone
cracks. It is made up of tiny pale
green-brown scales covered with a
prickly whitish patina, used for
protection against the sun’s rays.
Cladonia rangiformis and C. convoluta
live on limestone in Mediterranean
arid meadows. They are fruticose
lichens resembling sturdy shrubs, but
less than a few centimetres tall. The
former is more branched than the
latter, which has larger and flatter
entwining lobes.
Arid meadows above the tree-line in
the Alps and Apennines host many
other Cladonia species.
Some of them spread due to
fragmentation of their thalli caused by
grazing animals.
New specimens may therefore form
from tiny pieces of the original thallus
already containing the two symbiontic
partners, lichen and alga.
Cladonia furcata
Cladonia digitata
Sedge fields are extensive in Val
Masino, near Livigno, and as far as the
high-altitude valleys of the Braulio.
On early and mature stages of
crystalline rocky soil, there are other
covers, such as fescue grasslands
with various fescue (Festuca varia),
which give rise to very solid turf and
constitutes the Alpine grasslands of
the warmest, driest and sunniest
flanks, between 1800 and 2800 m, but
there are also other grasses. Various
species of fescue, with their large tufts
of long, thin leaves alternating with
short, sharp ones, often dominate, but
other species are visible, if not for their
numbers, for their gaudy flowers which
Alpine yarrow (Achillea moschata)
last until early winter, due to the
favourable microclimate. Among these are ramous gentian (Gentianella
ramosa), rock speedwell (Veronica fruticans), Alpine yarrow (Achillea
moschata), star hare’s-ear (Bupleurum stellatum), Haller’s sermountain
(Laserpitium halleri) and bearded bellflower (Campanula barbata).
Sedge fields with various fescue cover areas from the Spluga pass to Stelvio,
on the ridges of south-facing rocks or detritic slopes below. Various fescue is
a pioneer species on inland valleys of the Rhaetian flank, but it is more
extensive on the southern chain overlooking the Adda valley. It is the most
frequent type of grassland on the Eastern Alps on silica soil and is extremely
important in stabilising slopes.
Sedge fields with various fescue up stretch as far as the Carex curvula stands
and sedge fields with Haller’s fescue, thus occupying the most arid sites.
Grasslands with various fescue have no grazing value, especially when
sloping and southward-facing. If animals do graze on them, they soon turn
into secondary grasslands with mat grass. This association is important in
protecting the soil in deforested areas cleared of scrub.
A type of very dense grassland is found in Valle di Susa, between the Alpine
and upper mountain lines, containing Alpine meadow foxtail (Alopecurus
gerardi), Alpine avens (Geum montanum), and Swiss violet (Viola calcarata), all
of which develop in grassland containing mat grass. These species may be
transitional in more degraded areas, due to intensive grazing and frequent
31
32
fires, evolving towards another association which colonises sunny, dry and
warm flanks of the Southern Alps.
It contains singleflower knapweed (Centaurea uniflora), panicle fescue
(Festuca paniculata), spotted catsear (Hypochoeris maculata) and mountain
clover (Trifolium montanum ssp. rupestre), which associate with large
numbers of asphodel (Asphodelus albus), St. Bruno’s lily (Paradisea
liliastrum) and Nottingham catchfly (Silene nutans). Mat grass is also very
frequent here.
These species have food value for animals, but are not used for grazing due to
the presence of panicle fescue, which has sharp, hard leaves and whose tufts
grow deep into the soil, more protected from fire than other grasses.
Alpine grassland on limestone (Seslerietalia). On dolomitic and calcareous
surfaces, it is more difficult for grasslands to find soil, especially on large,
mobile detritic cones.
On these mountains, grasslands are dry and calciphilous, and develop on
flanks receiving little snow. The most typical and complete example of
grassland on limestone, especially in the central and eastern Alps, is the
association with various moor grass (Sesleria varia) and evergreen sedge
(Carex sempervirens): the Seslerietum-Semperviretum.
Various moor grass (Sesleria varia)
These are mature associations, and have great colonising and covering
capacity. They are typical environments for edelweiss, which may flower
extensively if undisturbed. In their early stages of development, SeslerietumSemperviretum forms continuous green terraces which, seen from below, look
like uninterrupted covers. If slopes are steep and debris keeps bringing down
tufts, these terraces may become permanent.
Grazing pressure increases interruptions in turf. If disturbance by animals
decreases or ceases completely, the association closes up and becomes
continuous, whereas excessive grazing gives rise to receding associations
with patches of barren land and colonisation by Firmetum, the plant
association dominated by glaucous sedge (Carex firma). Further treading
favours colonisation by mat grass.
Although the two typical species of this association have different
environmental requirements, they join other species to form one of the best
defined Alpine grasslands. Various moor grass, with its pale blue, silvery
spikes, is a typical pioneer species, which does not require much humus,
humidity or light. Evergreen sedge, with its tufts of hard, shiny leaves, is
saprophytic and heliophilous, indifferent to calcareous or siliceous substrates.
Many dolomitic grasslands, such as those on the top of Monte Baldo, the
bright green of which contrasts with grey rocky slopes and bare cliffs, are
typical examples of this association. Accompanying the two main species are
hawkweed (Hieracium villosum, H. bifidum), edelweiss (Leontopodium
alpinum), chamois ragwort (Senecio doronicum), houseleek (Sempervivum
tectorum), Alpine pasque flower (Pulsatilla alpina ssp. alpina) with large white
flowers, buckler mustard (Biscutella laevigata), cinquefoil (Potentilla aurea, P.
crantzii) and Alpine kidney vetch (Anthyllis vulneraria ssp. alpestris), all with
golden flowers, Swiss violet, with large pale blue flowers, and Alpine aster
(Aster alpinus), with yellow and violet heads.
On limestone, between 2000 and 2900 m, Alpine grasslands are composed of
associations of a few species, such as mountain avens (Dryas octopetala).
This plant of the rose family grows in dense patches which then join together
to form extensive covers.
These early stages are followed by others with increasing numbers of species,
like glaucous sedge, blue saxifrage (Saxifraga caesia), Clusius’s gentian
(Gentiana clusii), Kerner’s hawksbeard (Crepis kerneri) and various moor
grass. These give rise to typical Firmetum, pioneer Alpine grassland
associations resistant to cold and wind. Although Firmetum grasslands have
little food value, they are extremely important because they colonise mobile
slopes, hindering erosion and protecting soil.
33
34
Mosses in arid meadows
Although mosses are forced to live in
damp, shady areas because they lack
roots to absorb water and vascular
tissues to carry it, they have
developed adaptations which enable
them to survive even in difficult
conditions - to the extent that some of
them have not only colonised arid
environments like meadows, but have
even become characteristic species,
used to describe species of grasses.
One such example is Syntrichia (=
Tortula) ruraliformis, a typical species
of Artemisio campestris maritimaeKoelerietalia albescentis, with tufts 3-5
cm high, which lives on neutral or
alkaline sandy soil on shore dunes.
However, most species are not
associated with one particular habitat,
substrate or altitude, but with the
simultaneous occurrence of certain
ecological conditions. Some of them
are found in the Emilian PreApennines, which are particularly rich
in thermophilous steppe species. The
most typical are the rare Tortula
revolvens and its variety obtusata,
which lives in drier environments. They
are small, densely-tufted olive green
and brown plants which grow in chalk
cracks and hollows.
Siliceous substrates host Crossidium
squamiferum, a small plant with small,
dense, greyish-white tufts, often found
in warm, open areas in Val Venosta
(Upper Adige).
Another, typical mountain species,
also found on acid soil and sand in
sunny meadows, is Racomitrium
canescens, whose small sprigs of 3-6
cm form loose greyish covers.
Among the mosses living on limestone
there is Pleurochaete squarrosa, which
forms lax, yellow-green shrubs in sandy
Alessandra Di Turi
Tortula sp.
substrates. It is found on the upland
plains of the area near Finale, an
extensive limestone area in western
Liguria, which also hosts
Pseudocrossidium (=Barbula)
hornschuchianum, with loose pale or
olive green tufts and delicate sprigs, the
leaves of which are arranged in star
shapes; Homalotecium sericeum, with
silky dark green or yellowish-green tufts
1-3 cm thick, stolons, and erect, slightly
bent twigs; and Homalotecium
(=Camptothecium) lutescens with
adjacent stamens 1.5 cm apart, forming
loose, bright yellow hummocks.
Another typical moss on calcareous,
and even siliceous substrates is
Tortella tortuosa. Its name is due to
the behaviour of its leaves, which
twist when dry. It is a mountain
species with branched, entwined
stamens 2-5 cm long, and forms
dense tufts on rocks and soil.
As far up as 3000 m, windy slopes that are easily dried and undergo
temperature variations are colonised by an association which can withstand
the lack of snow (the snow cover lasts a maximum of 2 months). This is
Elynetum, a small, fragmented community that forms reddish-brown covers,
considered to be one of the most impressive vegetal associations of highaltitude mountains. Mousetail-like elyna (Elyna myosuroides) associates with
black Alpine sedge (Carex atrata) and various gentians (Gentianella tenella,
Gentiana nivalis, etc.).
Elynetum covers windbeaten ridges and dolomitic rocks and, although it is not
a pioneer species, it follows Firmetum; if soils are sufficiently acid, it may
evolve towards Curvuletum.
Elynetum grasslands are stabilised by extreme conditions, especially strong
winds that prevent snow from covering crests and ridges, exposing vegetation
to their action and to great temperature variations.
Along the Alpine line of Valle di Susa (Piedimont), between 2500 and 2750
m, Elynetum prevails on unstable soils, and on steep slopes facing south
and south-east.
Typical species, in addition to elyna, are short-leaved gentian (Gentiana
brachyphylla), Swiss milk-vetch (Oxytropis helvetica), oneflower fleabane
(Erigeron uniflorus) and often various moor grass.
Abandoned terraces turned into pasture, Ligurian Alps
35
36
Primary grasslands on the Ligurian Alps
Continental steppe grasslands
(Festucetalia vallesiacae). Alpine
valleys have a continental climate.
These sub-steppe valleys in the Alps,
surrounded by huge mountain chains
which hinder the passage of damp
winds, develop like islands, with
varying altitude and extent, in Valle di
Susa, the high Val Chisone, Valle
d’Aosta, Valtellina, Val Venosta and
Valle dell’Isarco.
The climatic conditions in these
valleys leads to low annual rainfall and
its distribution over the seasons so
that, in summer, vegetation suffers
from lack of water. The continental
climate in inland valleys gives rise to a
Central Alpine milk-vetch (Astragalus
centralpinus)
higher isothermal line, which raises all
altitudinal limits. As aridity increases,
species, plant and animal associations, crops and even towns all move
higher up in the mountains. The valleys and lower slopes have lost their
original woodlands, which were exploited and destroyed from the Stone Age
onwards, to the extent that erosion has even removed part of the soil. Here
are secondary arid grasslands with xeric grasses and heaths with wormwood
and dwarf shrubs (Lavandula angustifolia, Thymus vulgaris, Hyssopus
officinalis).
Arid valleys host large numbers of steppe species, isolated in Europe, and
witnesses of the migration that occurred during the final stages of the last
glaciation. A few have remained in only a couple of sites, others are scattered
in isolated xeric sites: forage kochia (Kochia prostrata) and the extremely rare
central Alpine milk-vetch (Astragalus centralpinus), which only lives in Valle
d’Aosta; yellow pheasant’s-eye (Adonis vernalis) is only found in Friuli;
Austrian milk-vetch (Astragalus austriacus) only in Valle di Susa, and Austrian
dragon’s head (Dracocephalum austriacum) only in Val Venosta and
Moncenisio. Many of these steppe immigrants, which are receding or have
totally disappeared in some areas, have geographic distributions and
similarities showing their Sarmatic, Iranian-Caspian, and even central Asian
origin. They are more numerous in the west, because the continental, subMediterranean components and steppe flora decrease eastwards.
37
38
Pre-Alpine and central Alpine valleys
are colonised by a peculiar association
that has adapted to extremely arid
conditions - sedge fields with Wallis
fescue (Festuca valesiaca).
These grasslands are highly xeric and
closely associated with south-eastern
European steppes. They are actually
the westernmost element of the
eastern European steppe which has
penetrated into the Alps. These
grasslands are found on the warmest
and driest southern slopes in Valle
d’Aosta, Valtellina and Val Venosta
where, in fragmented populations,
they may reach 2000 m on wellexposed slopes. They are marked by
Globe thistle (Echinops ritro)
the dense grey shrubs of Wallis fescue,
hair-like feather grass (Stipa capillata) and feather grass (S. pennata), a wavy,
feathery species, dwarf milk-vetch (Astragalus excapus), yarrow, cinquefoil
and sedges.
In Valle di Susa, xeric grassland grows along the mountain and sub-mountain
line, especially on the left flank of the valley, and contains xerophilous species
such as squinancywort (Asperula aristata), Somerset hair-grass (Koeleria
vallesiana), smooth thyme (Thymus glabrescens), tunic flower (Petrorhagia
saxifraga), and Spanish catchfly (Silene otites).
At mountain levels, arid meadows with Somerset hair-grass and rock-rose live
on sunny flanks, where the calcareous soil is locally eroded, as in the high
valley between Exilles and Oulx and in the deep Bardonecchia valley. Typical
species are saintfoin milk-vetch (Astragalus onobrychis), hairy milk-vetch
(Oxytropis pilosa), toadflax (Thesium linophyllon), bulbous meadow-grass (Poa
bulbosa) and white hawksbeard (Crepis albida). Among these, saintfoin milkvetch and toadflax are ecologically important and are also found in different
associations.
The main species are Somerset hair-grass, globe thistle (Echinops ritro), silky
scabious (Scabiosa holosericea), white rock-rose (Helianthemum apenninum)
and field wormwood (Artemisia campestris). In the Susa area, from Novalesa
to Meana, where species colonise lower areas of mountain levels, are
honewort (Trinia glauca) and hair-like feather grass meadows, with central
Wormwoods
Simonetta Peccenini
Wormwoods are typical steppe
species. Common wormwood
(Artemisia absinthium), native to the
Near East, was introduced and
naturalised in pre-Roman times. It is
traditionally used for medicinal
purposes and for making liqueurs. It is
now frequent in dry, uncultivated
Alpine land, and is found on the
Apennines as far as Basilicata.
Calcareous arid meadows along the
Italian peninsula host the spontaneous
male wormwood (Artemisia alba).
Dwarf wormwood (Artemisia borealis)
and Wallis wormwood (Artemisia
vallesiaca) are found in Alpine valleys
with a continental climate.
A species of wormwood, Artemisia
cretacea, features the most peculiar
distribution, as it only lives in eroded
clay soil and valleys of Pliocene age,
called calanches, and is endemic to
areas near Siena and Bologna.
One of the rarest species is
chamomile-leaved wormwood
(Artemisia chamaemelifolia), found in
the Alps along arid, rocky flanks with a
continental climate, between 500 and
2400 m. Its distribution area stretches
from the Pyrenees to Iran. In Europe, it
is found in the south-western Alps,
Pyrenees, Cantabrian Mountains, and
in north-western Bulgaria. In Italy, it
only lives near Cogne, in Valle d’Aosta,
and at Fenestrelle (Val Chisone) in
Piedmont, in Val Varaita and near
Garessio, in a location called “Garbo
della Luna”, from the name of the cave
opening in calcareous soil in the Costa
di Monte Merlino.
Wormwood (Artemisia absinthium)
Male wormwood (Artemisia alba)
39
40
European steppe species, such as
Haller’s pasque flower (Pulsatilla
halleri). These associations contain
branched bastard-toadflax (Thesium
divaricatum), Haller’s pasque flower,
small hare’s-ear (Bupleurum baldense),
mountain pasque flower, upright
brome (Bromus erectus), white rockrose, field wormwood, and meadow
globe daisy (Globularia punctata).
These species live on steep, westwardfacing slopes, partially leached and
Mountain pasque flower (Pulsatilla montana)
swept by the wind - one of the factors
responsible for the xeric conditions in
the area.
The association with Cleistogenes
serotina and tanglehead (Heteropogon
contortus) is more widespread and
lives on the left-hand flank of the
valley, along the sub-mountain line,
sometimes stretching as far as the
mountain line.
It has xerophilous populations, typical
species of which are broom grass
(Chrysopogon gryllus), Cleistogenes
serotina,
Mediterranean
fumana
White rock-rose (Helianthemum apenninum), a
Western European species
(Fumana ericoides), miniature restharrow (Ononis minutissima) and
common crupina (Crupina vulgaris). Less spontaneous and more localised are
tanglehead, small rest-harrow (Ononis reclinata), and Ligurian viper’s bugloss
(Onosma fastigiatum). Numerous and locally dominating are field wormwood,
white rock-rose, feather grass and field eryngo (Eryngium campestre).
The Valle d’Aosta also contains steppe-like environments, i.e., grassy and
shrubby plants distributed inland in the valleys, at Villeneuve, Sarre, Saint
Pierre, Aymavillesa and Gressan, between 500 and 1100 m. Here, meadows
with Somerset hair-grass, hair-like feather grass, Wallis wormwood and Swiss
viper’s bugloss grow on both flanks.
In the same areas, the continental climate has also affected soil genesis, giving
rise to a true crust, impermeable to roots and similar to that found in arid North
African countries. Here are meadows
with ciliate melick (Melica ciliata) and
forage kochia, a xeric chenopod native
to central Asian steppes, together with
many other annuals, unlike the usual
situation in the Alps.
These meadows always host tuft
false brome (Brachypodium rupestre
ssp. caespitosum), crested hair-grass
(Koeleria macrantha), pyramidal hairgrass (K. pyramidata), and Wallis
fescue, all species of little grazing
Field eryngo (Eryngium campestre)
value. Higher up in the mountains,
where farmland has been abandoned,
vegetal evolution is already visible
after just a few years, its transitional
stages marked by large numbers of
wormwood.
In the Valtellina (Lombardy), at
medium and low altitudes, fragmented
xeric grassland stretches on the right
bank of the river Adda as far as 600 m,
from the beginning of the valley to the
town of Tirano.
These populations are similar to those
with Cleistogenes serotina and Wallis
Carthusian pink (Dianthus carthusianorum ssp.
vaginatus)
fescue. Typical are the greenish-grey
tufts of Wallis fescue, a continental
species, and other gramineae with small grey, waxy leaves, revealing the lack
of water, such as Bothriochloa ischaemon, Cleistogenes serotina and crested
hair-grass, the most common. These associations grow along flanks in the few
spaces not occupied by vineyards, and invade tilled land when cultivation is
suspended, thus showing their climatic potential. The distributions of
meadows containing Cleistogenes serotina and Wallis fescue and vineyards
usually coincide.
Some xeric grasslands growing fragmented in the low Poschiavo valley and
above Tirano contain tuft false brome and the knapweed Centaurea bracteata.
These meadows also grow along the Rhaetian flank between Sondrio and
Tirano, up to 1400 m. Among their typical species, the most common are
41
42
Centaurea bracteata and Carthusian
pink (Dianthus carthusianorum ssp.
vaginatus). They colonise impoverished
soil resulting from the accumulation of
crystalline schist debris, the moistened
layer of which is never more than 5 cm
thick, with average acidity and no
limestone.
Brachypodietum associations, i.e.,
false brome meadows, are found
between 580 and 1400 m, far from
vineyards.
Although the soil is as impoverished
as that of Cleistogenes serotina and
Wallis fescue meadows, it seems
more stable, unaffected by the
movements of the various plants. It
Meadow clary (Salvia pratensis)
may evolve into Scotch pine
woodland and often derives from the burning of hay meadows.
At higher altitudes in Valtellina, as far as the Bormio valley and in the lower part
of Valdidentro, xeric grassland is composed of tunic flower and field
wormwood, feather grass, Wallis fescue and catchfly. It may be considered as
the continuation of sedge fields with Wallis fescue and, unlike the xeric
grassland in the lower valley, here grassy turf is continuous. It may be
degraded, as it is used for grazing, thus showing its inadequacy for this
purpose. Xeric grasslands include wormwood associations.
Xero-thermophilous sub-mountain and mountain Mediterranean
meadows (Brometalia). Grasslands on the southern and lower portions of
the Pre-Alps are composed of Brometum grasslands - impoverished, arid
meadows typical of sunny, low altitudes that stretch along slopes. These
meadows used to be regularly mown, but are now abandoned or only
occasionally mown.
Brometum grasslands are made up of various associations, according to the
varying arid conditions. They usually contain brome, a type of grass with
bearded panicles, found all over Italy and Sardinia with various similar types. It
grows in dry, sunny calcareous soil. In Europe, brome is one of the species
which has migrated furthest west during immigrations of steppe elements, and
usually associates with similar species. Primary Brometum grasslands are
discontinuous, found on outcropping
rocks, with rarefied carpets of brome
or Bothriochloa ischaemon. Meadows
are interrupted by turf covered with
debris, moss, and lichen. However,
most Brometum grasslands are
secondary and are probably connected
with man’s past abandonment of
farmland.
In late spring, both primary and
secondary Brometum grasslands
brighten up with colourful flowers,
from the pale blue of meadow clary
(Salvia pratensis) and meadow globe
daisy, the violet of big-flowered selfheal (Prunella grandiflora), the yellow
of common rock-rose (Helianthemum
Cone knapweed (Leuzea conifera)
nummularium) and kidney vetch
(Anthyllis vulneraria) and the pink of Carthusian pink. The most xerophilous
Brometum grasslands are physiognomically discontinuous and grow along the
driest and warmest slopes. They usually host large numbers of Mediterranean
species and are called Xero-Brometum. They stretch along the margins of
plains, the driest and barest fluvial-glacial terraces, as far as the sunny,
calcareous lower slopes of the Pre-Alps.
Typical of the lower belt are brome and, further up, pyramidal hair-grass
(Koeleria pyramidata). Brome associates with crested hair-grass (Koeleria
macrantha), fescue (Festuca gr. ovina), purple-stem catstail (Phleum
phleoides), Bothriochloa ischaemon and wall germander (Teucrium
chamaedrys), which form a kind of Mediterranean garrigue. The warmest and
driest areas host broom grass, Bothriochloa ischaemon and Cleistogenes
serotina, which also live in Alpine xero-thermal protected spots.
Lack of fertilisation favours the development of tuft false brome, of little
grazing value, to the detriment of important forage grasses. This is the case of
the eastern and western flanks of Monte Baldo, where the slopes are steep
and the soil is arid and thin, so that turf degrades. These meadows also
contain crested hair-grass, pyramidal hair-grass and Wallis fescue, all of little
value as food for livestock.
The main species is brome, which associates with other regional species. At
the southern foot of the Alps and along the Pre-Alps, from Piedmont to Carnia
43
is mountain pasque flower and honewort. Other interesting thermophilous
species are small rest-harrow in Val Grana and Valle di Susa, along Lake Garda
and the Veneto Pre-Alps, upright micropus (Micropus erectus) in the southern
Alps and along the Veneto hills, strawflower (Helichrysum italicum) in Val
Grana, the Veneto Pre-Alps and Euganean Hills, and Narbonne flax (Linum
narbonense) in Val Grana, the Veneto and Julian Pre-Alps and the Italian karst.
These species also grow in Apennine meadows: one of them, cone knapweed
(Leuzea conifera), is worthy of mention, as it is a xerophyte of steppe origin
with fragmented distribution. In Italy, this western Mediterranean species only
grows in one dry Alpine valley, at Falcemagna, near Bussoleno (Valle di Susa),
and in the Langhe, along arid, Mediterranean calcareous slopes of western
Liguria, on Monte Argentario (Tuscany) and in Sicily.
Semi-arid meadows, called Meso-Brometum, contain false brome, brome,
broom grass, Bothriochloa ischaemon, narrow-leaved flax (Linum tenuifolium),
wall germander, various scorpion-vetch (Coronilla varia), lady’s bedstraw
(Galium verum) and its relative Galium lucidum. These meadows develop on
the steepest slopes along the highest hills or even near the summits. Semiarid meadows often develop on terraces that were once vineyards. Their soil is
shallow, dry and poor in nutrients, with organic matter that retains humidity
and acts as a fertiliser. Their associations are composed of many colourful
species adapted to arid conditions, high temperatures, and lack of nitrogen.
Among these are many orchids - Orchis ustulata, O. morio, O. provincialis, O.
maculata, Ophrys bertolonii, O. apifera, O. sphegodes, Gymnadenia
conopsea, G. odoratissima, Anacamptis pyramidalis, Serapias vomeracea,
Limodorum abortivum, Cephalantera longifolia - all species loving open, dry
areas. There are also some steppe species, such as Siberian bellflower
(Campanula sibirica), kidney vetch, dropwort (Filipendula vulgaris) and
scabious.
44
Ophrys sphegodes
Arid grassland in the Karst area. The
so-called Chrysopogonetum with
Triestine knapweed (Centaurea cristata)
is a typical formation found at sea level
up to 400 m, mainly in the warmest and
sunniest areas along the Isonzo karst,
near Monfalcone and Gorizia, and even
further east and south. This is one of
the most thermophilous associations in
the area, with large numbers of
Winter savory (Satureja montana ssp. variegata)
45
Mediterranean species such as silvery
broom (Argyrolobium zanonii) and
Balkan species such as Carniola
broom (Genista sylvestris ssp.
sylvestris) and Triest pink (Dianthus
sylvestris ssp. tergestinus). It is an
evocative association, especially in late
summer and early autumn, when
Bothriochloa ischaemon, one of the
The Karst knapweed Jurinea mollis ssp. mollis
most numerous grasses, is in full
bloom. Beauty is enhanced by the
flowers of winter savory (Satureja montana ssp. variegata), Cleistogenes
serotina male wormwood, etc.
Karst cliffs also host another grass that is deeply affected by the bora (the
typical strong wind which so often blows in Trieste), the Seslerietum with silky
broom, which mainly contains Sesleria juncifolia, but also sedges. It is a
primary meadow, a true cliff steppe, found along high-gradient cliffs
(sometimes over 70°) and unaffected by man.
Other extensive associations stretching from Trieste inland, as far as the upper
karst, with constant ecological and floristic characteristics, are sedge fields
with cliff knapweed, composed of sedge and brome. Typical are Karst
knapweed (Jurinea mollis ssp. mollis) and silvery plantain (Plantago argentea).
46
Autumn in the Karst (Friuli Venezia Giulia)
Po Plain meadow fragments: low ridges, sand dunes and magredi (poor,
untilled land). In the Po Plain, grey hair grass (Corynephorus canescens) was
once certainly a common species on the sand dunes of Lomellina, the
moorland of Gallarate and riverbanks in Piedmont and Lombardy. In 1855, in
Piedmont, prairie rocket (Erysimum rhaeticum) was still found on the dunes of
Cambiano near Chieri, and in sand of the Po riverbed. Today, these species
are declining, due to the destruction of their natural environment for
agricultural, industrial or reafforestation purposes, and their Po Plain sites
have generally disappeared.
In Friuli, extensive stretches of arid meadows are called magredi. They
constitute the typical vegetation of the Friuli plain, with very arid grasslands.
They are very valuable, due to their peculiar flora. However, they are also very
vulnerable, and any disturbance may remove the grassy turf, causing
irreversible degradation. The aridity of the magredi is due to highly permeable
substrates, composed of calcareous and dolomitic-calcareous pebbles, on
top of which is a thin layer of clay, called ferretto, which is never more than 50
47
48
Magredi in the high Friuli plain (Friuli Venezia Giulia)
cm in depth, lacking in calcium and
organic matter, but rich in ferric and
aluminium nitrates. The colonising
vegetation is very uniform, giving rise
to a monotonous landscape which is,
in fact, very rich in plant species of
Eastern, Alpine and local origin.
Species are distributed in the various
phytocoenoses according to the
Friuli kale (Brassica glabrescens)
maturity of the vegetation which, in
turn, depends on soil evolution.
As modifications only occur gradually,
there are several transitional areas, the
boundaries of which are less clear-cut
than in the two main types: primitive
magredi, probably of primary origin,
on only slightly evolved soil, is
covered by various moor grass,
Tatarian sea kale (Crambe tataria)
Kerner’s spurge (Euphorbia triflora
ssp. kerneri), mucronate sedge (Carex
mucronata), common fumana (Fumana
procumbens), Friuli kale (Brassica
glabrescens) and the rosy-yellow
knapweed (Centaurea dichroantha).
More mature magredi is found on
ferretto, with more available water,
probably deriving from the destruction
Alpine stock (Matthiola fruticulosa ssp. valesiaca)
of mixed woodland, with common
bent (Agrostis tenuis), broom grass,
Yorkshire fog (Holcus lanatus), cliff fescue (Festuca rupicola), a species of
garlic (Allium cirrhosum), Pampanini’s rattle (Rhinanthus pampaninii) and
Ressmann’s scabious (Knautia ressmannii).
Among Oriental species is steppe kale (Crambe tataria), a majestic crucifer
living in the magredi near Pordenone, totally separated from its main
Pannonian-central Asian range.
The floristic variety of these pre-steppe associations is also due to the
phenomenon of dealpinisation, i.e., floodwaters bring species down to valleys,
where they find their ideal, impoverished soil. Examples are mountain avens
(Dryas octopetala) and Alpine stock (Matthiola fruticosa ssp. valesiaca).
49
50
Mushrooms and arid meadows
Mushrooms can develop on many
types of substrates and environments.
No wonder, then, that large numbers of
mushroom species may even be found
in particular biotopes like arid
meadows. Some are rare and
interesting from the mycological
viewpoint; others are common, but far
more in demand because they are of
good quality and edible. Interesting
species belong to the genus
Hygrocybe and, among the edible
ones, are Agaricus, Macrolepiota,
Pleurotus, Calocybe, Marasmius,
Lycoperdon, Langermannia, Calvatia
and Vascellum. Alkaline soil often hosts
several species of Hygrocybe, the
bright, lively colours of which are
clearly visible despite their tiny size (the
diameter of their caps is only 0.5-7 cm;
the length of their stalks is 1.5-7 cm).
Hygrocybe coccinea, commonly found
among grasses, grows in mountain and
plain meadows, either in clusters or
solitary. The surface of its cap is a
waxy, bright crimson; the upper part of
the stalk is also crimson; the base is
yellowish; the gills are pale yellowishorange.
Hygrocybe spadicea grows in low-lying
plain meadows up to the mountains,
even on sand and gravel; its cap is a
characteristic brown-grey, brown-sepia,
which contrasts with the citrine yellow
and orange-yellow of gills and stalk.
H. calyptriformis is rare, generally
found only in meadows at 1000 m, and
is easily recognised because it is the
only Hygrocybe with lilac, pink and
lilac-violet colours.
Another infrequent but interesting
species is Amanita nivalis, which grows
solitary in summer on Alpine meadows
and pastures. It is tiny, with a whitish-
Mirca Zotti
grey cap, no ring, delicate flesh, and
sheathing volva at the base of the
stalk. The genus Amanita includes
typical ecto-mycorhizal symbiontic
woodland fungi, i.e., their development
depends on their symbiosis with the
roots of a wide variety of trees, both
conifers and broadleafs. In Alpine
meadows, the presence of a species
like A. nivalis may appear anomalous,
as meadow fungi are generally
saprophytic (feeding on dead organic
matter). The reason is that A. nivalis
may be symbiontic on dwarf willows
(Salix herbacea, S. retusa, S. reticulata)
and on viviparous knotgrass
(Polygonum viviparum).
Edible species of the genus Agaricus
(e.g., A. campestris, A. macrosporus,
A. spissicaulis), commonly called
meadow mushrooms, are perhaps the
best-known fungi. They grow in fields
and meadows in plains, hills, and
mountains and in groups - often in
circles known as “fairy rings”. These
rings are both due to the distribution of
the mushrooms and to the circular
Parasol mushroom (Macrolepiota procera)
growth of the surrounding grasses.
Circular areas where vegetation is lush
and bright green alternate with sterile,
almost bare areas, due to substances
produced by the underground fungous
mycelium, which can either hinder or
favour plant growth.
Agaricus species are recognisable by
their brownish-white colours, typical
rings around their stalks, and gills
ranging from white, pink, or brownishgrey to brownish-purple when mature,
and flesh ranging from yellow to
pinkish-red. However, some Agaricus
species are toxic (e.g., A.
xanthoderma). These toadstools may
be identified by the colour of their
flesh, which is lemon yellow at the stalk
base (when cut or scraped), the
unpleasant smell, which recalls ink or
phenol, and intense yellowing when
their surface is scraped.
In Sicily and Sardinia, some typically
meadow species of the genus
Pleurotus are well-known and popular
due to their organoleptic properties.
These are a series of species similar to
Pleurotus eryngii, which is closely
associated with its substrate.
Pleurotus grows on the decaying roots
and stipes of some umbellifers like
giant fennel (Ferula communis), eryngo
and sermountain (Laserpitium
latifolium). It ranges in colour from
creamy-white to brownish-grey, with
cap margins initially folding
downwards, then stretching, and gills
along the stalks, which are inserted in
the middle of caps, with elongated,
root-like bases.
Other species, among the many found
in arid meadows, are Macrolepiota
procera and M. excoriata. The former
is commonly called parasol mushroom
and is very large. The diameter of its
cap may measure 30-35 cm, and is
slightly umbonate and covered with
gills.
The stalk is covered with conspicuous
brown scales and is bulbous at the
base, with a mobile, multi-layered ring.
M. excoriata is also edible and differs
from M. procera in that it is smaller,
with a smooth stalk and a singlelayered ring. It has a felty cap surface
that soon cracks open towards the
star-shaped margin.
Mushrooms commonly called puffballs
belong to various genera, such as
Lycoperdon, Langermannia, Vascellum
and Calvatia. They grow in meadows
and pastures, even in the mountains,
do not generally have caps or stalks,
but a globose shape, with double
membranes, the outer part of which
frays into variably-sized gills.
They are generally edible while
immature, and their flesh is firm and
white. Langermannia gigantea can be
extremely large, up to 50 cm in
diameter.
Scarlet hood (Hygrocybe coccinea)
51
52
■ Apennine meadows
Inland Apennine slopes are covered by secondary grasslands growing at
altitudes between 300 and 1200 m and, in the driest areas, up to 1700-1900
m. They usually develop along the sunniest and warmest flanks and derive
from the degradation or destruction of original broad-leaved woodland (oak
and beech woods) that were felled centuries ago to create space for farming
and grazing land.
These are discontinuous xeric meadows which often resemble Mediterranean
garrigue rather than true grassland, especially when environmental aridity and
shallow, even rocky soil increase the numbers of suffruticous and fruticose
chamaephytes.
However, these grassy associations look like Asian steppes, and mainly
contain grasses. They are impoverished meadows with large quantities of
brome, which grows from maquis glades to Apennine cliffs. Brome is a
widespread grass along the hilly, sub-mountain belt and is the dominating
species where grasslands become grazing pastures, giving rise to stable
meadows which remain such as long as the pastures last. When grazing
stops, the Brometum grasslands become scrub. Brome meadows are
widespread in Europe and in the Apennines on limestone and sandstone
Meadows on cultivated eroded land (northern Apennines)
mountains, but are absent in volcanic areas, in Sicily and Sardinia. The
Apennines are their southern European limit, due to the oceanic climate that
hinders their growth.
Apennine Brometum grasslands may roughly be divided into northern and
central-northern Apennines, and southern and central-southern Apennines,
although this distinction is not always very clearcut and the various coenoses
often mix.
It is therefore more sensible to divide mountain xeric grasslands according to
ecological and floristic factors, thus identifying two large groups:
● Markedly thermo-xerophilous formations, distributed at the foot of
mountains (Brachypodietum/false brome meadows)
● Less xerophilous formations distributed along the hills and mountains of the
Apennines (Brometum/ brome meadows).
Brometum grasslands. According to the varying environmental conditions
(microclimate, sunlight, soil, etc.), Apennine Brometum grasslands may
change their floristic composition, although the typical species of arid
meadows are always present. Among the most frequent, in addition to brome,
is false brome, salad burnet (Sanguisorba minor), yellow bedstraw, wall
germander, mouse-ear hawkweed (Hieracium pilosella), small scabious
Large yellow rest-harrow (Ononis natrix)
53
54
(Scabiosa columbaria), rock-rose, and
purple woodruff (Asperula purpurea).
The grassy associations found on the
calcareous-marly and marly-sandy soil
of the central-northern Apennines,
sometimes as far as the centralsouthern (Abruzzi and Molise) and
southern Apennines (Lucania) belong
to a homogeneous group called
Xerobromion.
They are generally arid grasslands
growing in glades of mixed woodland,
but are even found at higher altitudes,
along cliffs at 1200-1300 m, which are
composed of the ubiquitous brome,
Bothriochloa ischaemon, large yellow
rest-harrow (Ononis natrix) and
Narrow-leaved flax (Linum tenuifolium), found
in meadows with a limestone substrate
bulbous plants with attractive flowers
such as tassel grape hyacinth
(Leopoldia comosa) and St. Bernard’s lily (Anthericum liliago).
The most interesting species from the floristic viewpoint are some orchids of
the genus Ophrys, which bloom in spring. Xerobromion grasslands bloom
until autumn, with the pale blue flowers of autumn squill (Scilla autumnalis)
and amethystine eryngo (Eryngium amethystinum) and the yellow of yellow
odontite (Odontites lutea). Spanish broom (Spartium junceum) is also
frequent. Some of these typical associations grow in the low Val Borbera
(Piedmont), where large areas alternate cultivated or tilled land with
abandoned fields.
Where agriculture has ceased, mown fields evolve towards arid meadows.
Together with grasses of high grazing value, there are typical xeric species,
such as Centaurea bracteata, white bedstraw (Galium album) and greater
knapweed (Centaurea scabiosa), which enhance biodiversity and floristic
abundance. These grasslands cover the Adriatic flank of the Apennines
where grazing land once existed, growing in marly-sandy and sandy-silty
soil.
Other Brometum grasslands typical of Romagna and the Tuscan-Marches
Apennines (540-1000 m) grow in the eroded marl and marly ravines of Val
Curone, and contain species such as the tiny scorpion-vetch (Coronilla
minima), mountain flax, Montpellier milk-vetch (Astragalus monspessulanus),
Villars’s hawkbit (Leontodon villarsii),
mountain
germander
(Teucrium
montanum) and male wormwood. They
usually grow in surface soil with a xeric
microclimate and are considered as
pioneer species, due to the steepness
of the slopes. Arid meadows of the Po
Plain flank of the Ligurian Apennines
and the hills of the Oltrepo’ Pavese, as
far as the Langhe (Lombard plain), are
again different.
Val Bormida has a series of eroded
marly surfaces which, in the south, are
covered with steno-Mediterranean
species.
Their penetration into the Po Plain is
favoured by the low altitude (below
Montpellier aphyllantes (Aphyllantes
monspeliensis)
1000 m) of the Apennine divide.
Vegetal populations contain thyme, a
pioneer species widespread in Piedmont (Val Curone, Val Borbera and Val
Bormida). Less well distributed are other steno-Mediterranean species, like
felty germander (Teucrium polium) and Montpelier aphyllantes (Aphyllanthes
monspeliensis), a typical plant of various types of garrigue. Other
characteristic plants are rough feather-grass (Achnaterum calamagrostis),
Tyrrhenian knapweed (Centaurea aplolepa), brome and Bothriochloa
ischaemon.
One type of pioneer vegetation found on reliefs between Liguria and
Piedmont, which are composed of conglomerate with coarse pebbles and
rocky soil, is very similar to Mediterranean garrigue. Its associations grow on
very unstable flanks, and contain large numbers of chamaephytes, such as
thyme, Italian rock-rose (Helianthemum oelandicum ssp. italicum), a small
chamaephyte of mountain meadows, the fescue (Festuca robustifolia), an
endemic hemicryptophyte found along the Apennines, and two stenoMediterranean species, Mediterranean bedstraw (Galium corrudifolium) and
globe thistle.
The hills surrounding Bologna (300-600 m), along gentle slopes where erosion
reveals the underlying sandstone, contain other xerophytic grasslands with
steppe and arid meadow species like hoary rock-rose (Helianthemum canum),
Bothriochloa ischaemon, catchfly, and mountain masterwort (Peucedanum
55
56
oreoselinum). Arid locations with little soil are colonised by feather grass and
broom grass.
All xerophytic or semi-mesophytic associations along the mountains and
hills of the limestone Apennines may be included in the Brometum
grasslands. Examples are found on the calcareous-sandy soil near Forlì (low
Marecchia valley, Emilia-Romagna), at low altitudes (300-450 m) and along
the warmest flanks. They contain cock’s head sainfoin (Onobrychis caputgalli), wall germander, bindweed (Convolvulus cantabrica), and Cleistogenes
serotina.
Southwards, between Romagna and the Marches (400-550 m) the northern
flanks have semi-xerophilous meadows, once cultivated, with large numbers
of hemicryptophytes such as Southern daisy (Bellis sylvestris), Siberian
bellflower, houseleek, the blueweed-relative (Onosma echioides), star
anemone (Anemone hortensis), Southern timothy (Phleum ambiguum) and a
hawksbeard species (Crepis lacera).
Xeric grasslands of the karstic area stretching to the foot of the south-western
Alps are very different, and contain peculiar plants which do not grow in the
Apennines. Here are species typical of Mediterranean arid meadows which
develop inland, in the warmest karstic areas, and decrease with altitude and
falling temperatures, as mountain species increase.
Plants usually found in shallow, calcium-free soil are dwarf sedge (Carex
humilis), globe daisy (Globularia cordifolia), feather grass, mountain
germander, silky broom (Genista sericea), narrow-leaved fleabane (Inula
ensifolia), and Austrian viper’s grass (Scorzonera austriaca var. stenophylla).
Southwards, on calcareous soil between southern Umbria and Latium (but
also in Basilicata, with slight differences) grows a Brometum with winter
savory (Santureja montana), and other species, some of which are endemic,
like knapweed (Centaurea deusta ssp. deusta) and Apennine wallflower
(Erysimum pseudorhaeticum), and many other frequent plants, such as
meadow globe daisy, woodruff (Asperula aristata ssp. scabra), fleshy
penny-cress (Aethionema saxatile) and strawflower. These are also
secondary grasslands, which came into being as a result of deforestation
and grazing.
Along the main limestone reliefs, from the Sibillini mountains to Calabria, there
is a pioneer grassy formation, absent in northern Italy, which grows between
1600 and 2100 m, where the substrate outcrops and alternates with fine soil.
The most frequent species found there are Majella thrift (Armeria majellensis),
Tenore’s yarrow (Achillea tenorii), potentilla, mountain clover (Trifolium
montanum) and Genua false brome (Brachypodium genuense).
57
Ophrys apifera
Ophrys insectifera
On the southern flanks of the Umbria-Marches Apennines, along the
mountains and particularly the hills, are arid meadows with purple woodruff
and brome. These herbaceous plant communities grow on calcareous
substrates with very little soil and plentiful rocky surface debris. Although
these areas are extremely arid, they are brightened by the varied flowers of
round-headed leek (Allium sphaerocephalon), hawksbeard (Crepis lacera),
amethystine eryngo, pink (Dianthus ciliatus), Atlantic hyacinth (Muscari
atlanticum), male wormwood, and several orchids of the genus Ophrys.
Mountain Brometum grasslands also stretch along the highest reliefs of the
Latium Apennines. At the foot of the Monti Ernici are xeric grasslands eroded
by excessive grazing, with large areas containing outcropping calcareous
rocks, and poorly evolved soil, conditions which give rise to arid environments
and restricted tree species. Since ancient times, grazing, particularly of sheep,
has been the most important factor in balancing and maintaining these
coenoses which, in the Mediterranean area, are considered the forefront of
eastern European steppe vegetation. On the Apennines, in addition to steppe
species, there are local, endemic and Mediterranean-orophilous plants, such
as dusty miller (Cerastium tomentosum), hawksbeard, Apennine stock and
purple toadflax (Linaria purpurea), which contribute to the biodiversity of these
associations. Brome is often replaced by Genua false brome, which is
58
endemic to the Apennines and typical of acid substrates, together with mat
grass. If the latter species dominates, it gives rise to Nardetum grasslands,
with marked floristic impoverishment.
A very special type of grassland, more similar to garrigue than true meadow,
develops on outcropping calcareous rocks on the arid slopes of the LepiniAusoni-Aurunci mountains, between 700 and 1200 m, but also in the central
Apennines, in the Marsican valley. The main species in these associations is
sage (Salvia officinalis), which usually grows in Dalmatia and is only found
locally in Italy. The plants of these discontinuous meadows are thermoMediterranean species, due to the low altitudes. The most common are yellow
woundwort (Stachys recta ssp. labiosa) and a kind of parsley (Elaeoselinum
asclepium). There are also small shrubs, such as strawflower, Genua broom
(Genista januensis), spiny broom (Chamaecytisus spinescens) and spiny
spurge (Euphorbia spinosa), which make this vegetation look like garrigue
covering other Apennine areas. There are other rare species, like prostrate
broom (Cytisus decumbens) and Gasparrini’s pearlwort (Buglossoides
gasparrinii). Mediterranean plants include Salerno’s violet (Viola pseudogracilis
ssp. pseudogracilis), endemic to the central-southern Apennines, which finds
its northernmost distribution area along the Aurunci, and cliff knapweed
(Centaurea rupestris), which here grows in its southernmost limit.
Summer flowering on limestone in Sirente-Velino National Park (Abruzzi)
Brachypodietum grasslands. Sunny,
siliceous, leached soil all along the
Apennines hosts meadows the main
grass of which is false brome.
These formations are therefore called
Brachypodietum (false brome meadows)
and often contain species from
Xerobrometum grasslands. This is
another type of secondary grassland,
as it derives from the degradation of
oak woodland.
Marly-sandy substrates between
Umbria and inland Latium, at low
altitudes (300-1200 m) are covered
with Brachypodietum, grasslands
dominated by cliff false brome
(Brachypodium rupestre), the rhizomes
Rayed broom (Genista radiata)
of which have great capacities for
vegetative development, enabling it to colonise clayey soil. It is very common
in Latium, from the valleys upwards, as far as 1500-1600 m, where it
competes with brome and other grasses, especially on calcareous-marly or
clayey soil. It grows in wood clearings, and also along the margins of
deforested areas.
Brachypodietum grasslands, with the endemic yellow milkwort (Polygala
flavescens), spiny rest-harrow (Ononis spinosa ssp. spinosa), rockcress (Arabis
collina), birdsfoot trefoil (Lotus corniculatus), etc., typically grow on plains, on
clayey soil containing little organic matter.
Among Brachypodietum grasslands is a type of shrubby-grassy vegetation
typical of the central and northern Emilian-Tuscan Apennines. In addition to
cliff false brome, it contains rayed broom (Genista radiata), a small shrub
growing in very dry areas.
These plants are not affected by grazing, and constitute natural vegetation
colonising dry, rocky slopes and detritic valleys. False brome has sturdy tufts
which tangle around rocky debris, consolidating it; they also produce a great
quantity of organic matter, which contributes to the creation of soil.
These often inaccessible populations growing on rocky crests also contain
horned rampion (Phyteuma scorzonerifolium) and stout umbellifers such as
moon carrot (Seseli libanotis) and Austrian milk-parsley (Peucedanum
austriacum).
59
60
■ Coastal meadows of southern
Italy, Sicily and Sardinia
Along the coasts and in the regions of
southern Italy, in addition to perennial
grasslands mainly composed of
tufted hemicryptophytic grasses,
similar to those of the Apennines,
there are meadows composed of
annuals (therophytes), which make up
the so-called therophytic meadows.
This vegetation often creates a mosaic
of therophytic and hemicryptophytic
elements, although these communities,
despite their similarities, have differing
structures, ecology and floristic
components.
Ampelodesmos mauritanicus
Perennial steppe meadows (Lygeo-Stipetea). These steppe grasslands are
composed of perennial thermo-xerophilous species with tufted gramineae.
Most of them are grasses, such as Ampelodesmos mauritanicus, Thatching
grass (Hyparrhenia hirta), esparto grass (Lygeum spartum) and rice millet
(Oryzopsis miliacea). This vegetation grows in the most xeric Mediterranean
regions, often covering extensive areas, and is also found on hills and
submountain locations in southern Italy. It makes up primary meadows on
eroded soil and outcropping rocks, and is the most mature vegetation growing
in these areas. It often constitutes extensive secondary meadows, due to
man’s disturbance (deforestation, grazing, fire) over time.
Esparto grass meadows (Lygeo-Stipetalia). On eroded clay, perennial
grasses are composed of esparto grass. This gramineae colonises very steep
eroded surfaces, consolidating clay substrates with its sturdy stolons and
hindering surface erosion. Esparto meadows are brightened by the yellow
heads of viper’s grass (Podospermum laciniatum).
This type of vegetation has specific ecological requirements, as it lives
exclusively in clay soil with a warm Mediterranean climate and long arid
summers. It grows in the driest southern Mediterranean areas; in Italy, it is
found in southern Latium, Campania, Basilicata, Apulia, Calabria, Sicily and
Sardinia. Other thermophilous esparto meadows contain south Mediterranean
species, such as Sicilian caper (Capparis sicula) and moricandia (Moricandia
arvensis), which grow in eroded clay along the Ionian coast and in Sicily.
In areas with a cooler climate, on the Ionian flanks of Basilicata and Calabria,
meadows contain Tenore’s knotgrass (Polygonum tenoreanum) and other
viper’s grass species endemic to eroded soil in southern Italy.
In the western Mediterranean, there is branched false brome (Brachipodium
retusum = B. ramosum). This localised perennial grass grows everywhere in
central-southern Italy, Sicily, Sardinia and, occasionally, in Liguria. It generally
lives on gentle slopes, in slightly evolved, calcareous rocky soil. It requires an
oceanic climate, with high relative humidity in the air and cryptoprecipitations throughout the year. If these conditions fail, large tufted
gramineae and chamaephytes take over, being better adapted to a
continental or xeric climate. These are discontinuous grassy covers, the
average height of which never exceeds 40 cm, and are composed of dense
populations of branched false brome and many chamaeophytes,
hemicryptophytes and geophytes.
These meadows cover restricted surfaces, and sometimes create a mosaic
with garrigue and basophilous, therophytic formations. The latter prevail in
conditions of intense disturbance, e.g., frequent fires, excessive passage of
livestock and grazing. Their evolution enables the colonisation of small shrubs
Spring flowering in Sardinia
61
62
such as rock-rose (Cistus salvifolius, C. monspelliensis, C. creticus ssp.
eriocephalus, C. albidus), thyme-leaved fumana (Fumana thymifolia),
thymelaea (Thymelaea tartonraira), cat thyme (Teucrium marum) and rosemary
(Rosmarinus officinalis), which are favoured by the presence of grazing
animals, because they have no food value.
A particular Brachypodietum, the main species of which is rush-like feathergrass (Stipa offneri), together with branched false brome and shrubby clover
(Dorycnium pentaphyllum ssp. suffruticosum), colonise rocky, south-facing
flanks, between 700 and 1200 m, along the calcareous plateau that makes up
the typical blocks of central-southern Sardinia, especially the Tacco di Osini
and Montarbu di Seui.
Again in Sardinia, along the coastal hills of Arbus, on ancient alluvial soil, there
are Brachypodietum grasslands containing warty milk-vetch (Astragalus
verrucosus), a paleo-endemic species restricted to a very small coastal stretch
in south-western Sardinia, and lesser feather-grass (Stipa bromoides).
In north-eastern Sardinia, at higher altitudes (200-900 m) along the western
and northern slopes of Monte Albo, in Supramonte di Urzulei and on the
island of Tavolara, an island off north-east Sardinia, Brachypodietum
grasslands host an oat-grass species (Trisetum splendens) which is endemic
to Sardinia and Sicily.
Brachypodietum grasslands sometimes contain large numbers of geophytes,
such as Narbonne star of Bethlehem (Ornithogalum narbonense), and two
squill species (Urginea fugax and U. undulata), all rare, south-Mediterranean
species.
The calcareous mountain tops of the Tacco di Montarbu, with deep crevices
and eroded substrates due to surface karstic phenomena, contain calcicole
orophytes, like grass-leaved buttercup (Ranunculus gramineus), a southwestern European species seldom found in Sardinia, Barbagia moor grass
(Sesleria insularis ssp. barbaricina), a Sardinian endemite, and Illyrian sea
daffodil (Pancratium illyiricum), a Sardinian-Corsican geophyte.
Thatching grass meadows (Hyparrhenietalia). Different, although not clayey
substrates (limestone, schist, gneiss, marl, calcarenite, conglomerate, sand)
host thermo-xerophilous perennial grasses, mainly graminaceae, such as
thatching grass, Ampelodesmos mauritanicus and rice millet.
These typically Mediterranean and Macaronesian associations grow in hilly,
submountain soil with a warm, arid climate, and are secondary meadows, as
they derive from the degradation of shrubby and woody maquis and ilex
vegetation.
Areas with deep, mature soil, rich in silt
and clay, contain steppe-meadows
with Ampelodesmos mauritanicus, a
large tufted grass growing in the
south-western Mediterranean area,
the primary locations of which are in
cliff habitats. They have fewer
bioclimatic requirements than thatching
grass meadows. These grasslands
have few flowers and are favoured by
repeated fires, because Ampelodesmos
mauritanicus regenerates its subaerial
parts immediately after being burnt
down. In addition to Ampelodesmos
mauritanicus, these grasslands contain
gypsophila (Gypsophila arrostii) and
Sicilian oat-grass (Helictotrichon
convolutum), eastern Mediterranean
species found in Basilicata, Calabria
and Sicily.
Thatching grass, a Mediterraneantropical plant frequent in very arid,
sunny environments of coastal and
hilly areas, grows together with twospike
beardgrass
(Andropogon
distachyus), Cachrys libanotis and
Greek micromeria (Micromeria graeca
ssp. graeca) in perennial grassy
associations in surface soil with
outcropping rocks. These grasslands
grow in the most arid parts of the
central and western Mediterranean
and are usually secondary, associated
with degradation and colonisation by
abandoned crops.
In extremely arid soil, these formations
are replaced by meadows with
thatching grass and tufted steppe
gramineae, such as a species of
63
Rush-like feather-grass (Stipa offneri)
Grassy-leafed buttercup (Ranunculus gramineus)
Illyrian sea daffodil (Pancratium illyricum)
Annual thermo-xerophilous meadows. One characteristic of Mediterranean
vegetation, as may be noted along the southern Italian coasts and the islands,
is the occurrence of usually small areas completely covered by annual,
precocious dwarf plants.
Glades in the maquis and small, natural patches of soil between rocks are the
favourite locations of various small species, as well as others which would
normally grow taller, but instead are dwarves that flower, bear fruit and die
without ever reaching their normal height. Later, the same locations are
colonised by generations of larger plants, which erase all traces of the
previous vegetation.
This annual, precocious, short-lived dwarf vegetation is called “precocious
Mediterranean microflora”. It usually grows in the maquis, but is not
associated with particular substrates, as it is found in various types of soil,
from calcareous, on the island of Giannutri, to granitic on the island of Giglio
(Tuscany).
“Nanism” or dwarfism is a biological adaptation to the environment for various
reasons. Areas occupied by microflora are not colonised by perennials and
therefore, with the early autumn rains, the soil is suitable for the germination of
many seeds. In this soil free of competitors, the young plants develop well
initially, but soon start competing with each other. They lack space and
nourishment, and therefore their growth is arrested. In the meantime, other
normally small plants have fully developed.
However, even microflora may flower and bear fruit, because its neighbours,
which are also small, do not deprive it of light and air. When the life-cycle of
precocious microflora ends, later plants flower and continue growing, so that
young plants are commonly seen amid flowering and fructifying dwarf
plants, giving rise to a richer second vegetation. This is also annual and its
life-cycle ends before the summer heat, leaving the soil free until a new
germination begins.
M E D I T E R R A N E A N V E G E TAT I O N
SUB-MEDITERRANEAN
V E G E TAT I O N
CLIMAX
OF OLIVE AND CAROB
CLIMAX
OF ILEX
CLIMAX
OF PUBESCENT OAK
(Oleon Ceratonion)
(Quercion ilicis)
(Quercion pubescentis)
littoral garrigue
garrigue
pseudo-garrigue
littoral steppe
steppe with Brachypodium
xero-brometum
direction of decreasing heat needs
evolutionary series
regressive series
transition and mixes
Diagram illustrating evolution of Mediterranean and sub-Mediterranean vegetation
evolutionary series
threeawn, Aristida caerulescens, and a
species of foxtail grass, Tricholaena
teneriffae. They are MediterraneanSaharan-Arabian species and show the
typically xeric characteristics of these
meadows. In Europe, Tricholaena
teneriffae is only found in southern
Calabria, and near Messina in Sicily. It
is a Saharan-Arabian species that
colonises cliff environments and steep
sandy deposits.
Man’s activities, which favour soil
erosion and degradation by destroying
the woody components of the maquis,
have accelerated the growth of these
grasslands,
which
now
cover
extensive
areas.
They
are
relict
plants
False yellowhead (Dittrichia viscosa)
of the Tertiary epoch, and today grow
in a few isolated areas with a very arid climate, e.g., along the coast between
Reggio Calabria and Capo Spartivento. Some perennial steppe grasses are
associated with synanthropic, sub-nitrophilous environments, such as road
edges and abandoned fields. This type of vegetation contains rice millet and
false yellowhead (Dittrichia viscosa), together with steppe hemicryptophytes
and geophytes, and grows in hilly, submountain Mediterranean areas.
Disturbed stretches are colonised by wall barley (Hordeum leporinum) or fourleaved allseed (Polycarpon tetraphyllum).
regressive seies
64
65
66
Therophytic meadows on salty, eroded cliffs, where normal evolution is
impossible, may be primary, although they are generally secondary and
constitute the last degradation of shrubby Mediterranean maquis.
According to the ecological conditions in which they grow, they may be
divided into:
· Basophilous thermo-xerophilous meadows (Stipo-Trachynietea distachyae)
· Acidophilous thermo-xerophilous meadows (Tuberarietea guttatae)
· Mediterranean halophilous meadows (Saginetea maritimae)
Basophilous meadows. Ephemeral
xerophilous,
neutrophilous
or
basophilous vegetation grows on
calcareous, dolomitic, marly, clayey,
sandstone,
sandy
or
volcanic
substrates, in the driest Mediterranean
and Macaronesian areas. These
meadows have winter-spring cycles and
contain large numbers of therophytes
and sometimes small geophytes.
This type of pioneer vegetation occurs
on rocky substrates, with surface or
poorly evolved soil. It lives in
Mediterranean coastal areas, in
southern Italy and on the islands, even
in the mountains. These are usually
secondary ephemeral meadows, in that
Iris (Gynandriris sisyrinchium)
they are the final stages of degraded
scrub and woodland. They are primary only along rocky crests and cliffs.
They contain many annual xerophytes typical of the south Mediterranean or
Mediterranean-Iranian-Turanian areas, such as Atractylis cancellata, fascicled
brome (Bromus fasciculatus), the halophilous hare’s-ear (Bupleurum
semicompositum), the iris (Gynandriris sisyrinchium), purple brome (Bromus
rubens), French ground pine (Ajuga iva), chicory (Hyoseris scabra), annual
catsear (Hypochoeris achyrophorus), curved hard-grass (Parapholis incurva),
annual feather-grass (Stipa capensis), the grass Trachynia distachya, and
many others.
Ptilostemon stellatus, narrow-leaf crucianella (Crucianella angustifolia),
Southern cudweed (Filago eriocephala), sphaeric vetchling (Lathyrus
sphaericus), Neapolitan melilot (Melilotus neapolitana), and cock’s-head
Useful plants in arid meadows
Paolo Maria Guarrera
In Sardinia, several species of asphodel thorny parts, are cooked like artichokes
and the leaves, deprived of the thorns,
are used to make the typical woven
are added to soups. The fruit (achenes)
panniers (canisteddu) and baskets
contain silimarine, are useful in liver
(corbule or colbe), which are decorated
disturbances, and are employed in
with geometrical patterns obtained by
popular medicine.
alternating paler fibres (from the inner
In Tuscany, strawflower (Helichrysum
parts of the plant) with darker ones
italicum), an aromatic plant with anti(from the outer parts).
inflammatory properties, is used as a
The most common patterns are stars,
digestive in veterinary and
flowers, roses, birds, leaves, etc. In
homeopathic medicine.
order to prepare them, stalks are
Ampelodesmos mauritanicus is used to
uprooted and collected in bundles in
make ropes, fish baskets, and cereal
early spring, dried in the sun for about
and vegetable containers, as well as to
three weeks, cut into strips, and then
dried again in the open air. Before being cover demijohns, large bottles, chair
seats, etc. South-east of Rome, the
woven, they must be soaked for a few
long inflorescences of Ampelodesmos
hours. Asphodels are particularly used
were used to make torches and thatch
in Ollolai where they are called iscraria,
for roofs. In ancient times, in southern
Olzai, Montresta, Tinnura, and Flussio
Latium, Ampelodesmos was used as
(in the province of Nuoro) (and in the
twine to tie up vines, as its name
province of Sassari, where they are
suggests (from the Greek words
called almuttu and iscareu in the local
ampelos = vine, and desmos = twine). It
dialect, Logudurese). Asphodels are
actually means “vine-twine”, and the
typically found in long-abandoned
attribute tenax - used in the past as a
pastures impoverished by excessive
synonym of the specific epithet - refers
grazing. In the past, near Anzio
to its robust fibre. In Sicily the
(Latium), boiled asphodel tubers
stalks of giant fennel (Ferula
were applied to relieve
c. ssp. communis), called
reddened skin. Milk
fella, were cut at the
thistle leaves (Sylibum
base and dried to
marianum) can be
make stools called
eaten despite
firlizze. The stalks
their thorns. In
were collected in
Latium,
summer, and cut
Basilicata,
at the nodes into
Calabria, and
several segments
other Italian
of equal length.
regions, even the
These stools are still
young stalks are
used in country
peeled of their cuticle,
houses in Mazara del
chopped, and boiled or
Vallo and Ristretta,
fried. In Basilicata, milk
and also in parts of
thistle heads, deprived
Sardinia.
of their fibrous and
Basket made with asphodel
67
69
68
Rough dogstail (Cynosurus echinatus)
Buckshorn plantain (Plantago coronopus)
sainfoin give rise to ephemeral, highly xerophilous meadows, associated with
extremely arid conditions. They grow on various types of substrates: marl,
clay, calcarenite, conglomerate and occasionally sand. Eroded clay and
esparto meadows also host small, ephemeral, pioneer vegetation with
Calabria plantain (Plantago amplexicaulis), a south-Mediterranean species
which only grows in eroded areas of the Aspromonte.
Ephemeral subhalophilous, thermo-xerophilous meadows with precocious,
short-lived microflora grow next to cliff and meadow halophilous thatching
grass in Sicily, Sardinia and southern Italy. These pioneer species are found in
coastal environments and may be identified by the nearby presence of sea
ryegrass (Catapodium marinum) and by the flattened rosettes of buckshorn
plantain (Plantago coronopus).
Acidophilous meadows. Ephemeral, annual, acidophilous, and nitrofuge
meadows with winter-spring life-cycles colonise areas in scrub and perennial
meadows. They grow in compact or sandy siliceous soil, with surface or
poorly evolved soil.
Along the plains, gentle cliffs and in garrigue glades are ephemeral
acidophilous, therophytic phytocoenoses living in oligotrophic (lacking
nutrients) and slightly evolved soil with a sandy-silty texture. They are
carpeted by grasses such as fescue (Vulpia bromoides, V. geniculata, V.
Small-flowered catchfly (Silene gallica)
Hair-grass (Aira cupaniana)
ligustica, V. myuros), bristle tail grass (Psilurus incurvus) and rough dogstail
(Cynosurus echinatus), and dotted with the yellow flowers of sea centaury
(Centaurium maritimum), orange birdsfoot (Ornithopus pinnatus) and umbrella
milkwort (Tolpis umbellata), and the white ones of thale cress (Arabidopsis
thaliana) and asterolinon (Asterolinon linum-stellatum).
Degraded rock-rose meadows contain silver hair-grass (Aira caryophyllea),
Andryala integrifolia, great quaking grass (Briza maxima), annual lupin (Lupinus
angustifolius), narrow-leaf cottonrose (Oglifa gallica), small-flowered catchfly
(Silene gallica), haresfoot clover (Trifolium arvense), and spotted rock-rose
(Tuberaria guttata).
More pioneer, ephemeral, and strictly acidophilous meadows growing in
immature, arid surface soil contain the delicate flowers of hair-grass (Aira
cupaniana, A. elegans) and yellow daisy (Coleostephus myconis). The
ephemeral pale yellow corollas with central dark spots of spotted rock-rose
add enhanced colours.
The southern sandy hills near Reggio Calabria host highly specialised
ephemeral meadows, with very rare xerophilous therophytes, such as
wahlenbergia bellflower (Wahlenbergia nutabunda) and Chian chamomile
(Anthemis chia). In mountainous parts of Calabria, Sicily and Sardinia, small
rocky clearings with surface soil host ephemeral Mediterranean-mountain
species such as forget-me-not (Myosotis incrassata), annual knawel
70
Harestail grass (Lagurus ovatus)
Dyer’s alkanet (Alkanna tinctoria)
(Scleranthus annuus), and the smallish violet Viola parvula. In Aspromonte, at
altitudes over 1400 m, well-exposed, windy locations on weathered schist
and gneiss substrates are covered by ephemeral orophilous meadows with
heath pearlwort (Sagina subulata) and annual knawel.
Coastal sand dunes and sandy deposits inland are colonised by annual
xerophytic meadows with psammophilous species, such as harestail grass
(Lagurus ovatus), small, velvety plants of strand medick (Medicago litoralis),
hair-grass (Corynephorus fasciculatus) and ryegrass (Cutandia maritima).
Ephemeral xerophilous meadows associated with coastal dunes, sometimes
even fossil dunes inland, are colonised by gaudy psammophytes like dyer’s
alkanet (Alkanna tinctoria) and branched malcolmia (Malcolmia
ramosissima). Annual psammophytes with Silene nicaeensis and Cutandia
maritima live in retrodunal areas, on flat sandy surfaces.
Quiet areas are colonised by the chamomile Anthemis tomentosa, an
eastern Mediterranean psammophyte, and by Hypecoum imberbe, a very
rare species in Italy, found in a few sites along the Apulian, Calabrian and
Sardinian coasts.
In southern Italy, flat areas between mobile dunes are colonised by annual
psammophilous plants with more pioneer characteristics, such as sainfoin
milk-vetch and branched malcolmia. They manage to live in coarse, unstable
sand.
Halophilous Mediterranean meadows. Small, pioneer meadows in perennial
halophilous vegetation with sea barley (Hordeum maritimum), curved hardgrass, buckshorn plantain and sea pearlwort (Sagina maritima) colonise
retrodunal hollows, rocky coastal crevices and eroded clay. In these locations,
aridity is not due to the climate, but to physiological causes, as any available
water is too salty to be used by these plants.
Ephemeral macrophytes of these salty environments, such as centaury
(Centaurium spicatum, C. tenuiflorum), annual sea-heath (Frankenia
pulverulenta), annual beard-grass (Polypogon monspeliensis) and lesser sea
spurrey (Spergularia marina) colonise temporarily flooded eroded soil. In early
spring, when the soil is still damp, these meadows are in full bloom, but die out
at the end of the season.
Annual halophilous vegetation in eroded clay, which is very humid in winter,
contain viper’s grass (Podospermum canum), which grows in Sicily and on the
Ionian coast in Calabria.
Expecially on the small islands, huge amount of droppings of large colonies of
seagulls enriches the soil with nitrates. This gives rise to halo-nitrophilous
vegetation with sea knotgrass (Polygonum maritimum), prickly saltwort
(Salsola kali), sea rocket (Cakile maritima), lesser sea spurrey, stock (Matthiola
tricuspidata), etc.
An arid meadow near the Salento coast (Apulia)
71
Invertebrates
ALESSANDRO MINELLI
The environmental feature that most
affects insect populations in arid
meadows is a strong seasonal nature.
This characteristic may be modified by
man by mowing which, paradoxically as
it may sound, sometimes stimulates
new flowering in plants. In any case, the
temporal distribution of the main food
sources (spring buds, leaves, flowers
and seeds) enables most insects in
these habitats to produce a single
generation each year. For instance, the
abundant production of pollen from
grasses coincides with the spring
abundance of flower beetles, such as
Oedemera, Cantharis and Rhagonycha,
which feed on that pollen.
The sexual dimorphism of Oedemera,
Coccinella septempunctata on broom
which have very beautiful metallic
teguments and a pair of very narrow
wing covers (elytra), barely covering their rear wings, is very curious. The males
have arched, swollen rear femura. As the beetles of these three genera
frequently visit flowers, one would expect these legs to play an essential role in
pollination, but this is not so. Their bodies are not very hairy and pollen does
not stick to them. The fact is that they are unreliable, i.e., unlike good pollinators
such as bees, which insist on visiting the same flowers for hours on end, these
beetles, after examining one flower covered with pollen, move on to another of
a completely different species, so that the pollen they might have carried along
is immediately wasted. Moreover, great quantities of pollen fall into the mouths
of ladybugs, like the common, euryecious Coccinella septempunctata. It
generally lives in arid meadows and, as an adult, has a mixed diet of pollen and
small prey (aphids), which is its only food at the larval stage.
The owlfly, or ascalaphid, Libelloides
73
74
Among plant families in arid meadows,
the most appeasing to insects are
grasses, leguminous plants and
labiates. The trophic role played by
orchids is far less important, although
they are very gaudy and locally
abundant. The insects that visit orchid
flowers of the genus Ophrys (their
main pollinators) are the males of a few
genera of solitary bees, or the related
sphecid wasps. The latter, however,
are not looking for this pollen as food,
they land on the flowers attracted by
chemical stimuli which they mistake
for their females’ pheromones. As they
touch the flowers, the sticky pollen of
Ophrys attaches to their bodies, and
this phenomenon is repeated so often
Bee on asphodel
that it guarantees that plenty of pollen
will end up on flowers of the same
species, fertilising them, but none is left for the insects.
As regards grasses, there are three primary conditions that make them
valuable as food. The first is the great number of these plants, although their
leaves have little food value, and are so hard and siliceous that only
phytophages (plant eaters) with robust mouthparts can chew them. This is a
typical characteristic of orthopterans (grasshoppers, crickets, cockroaches)
which are, in fact, very numerous in these environments. The second reason
why grasses are important food sources for insects in arid meadows is the
great quantity of pollen they produce, which is used by many species,
especially beetles. The third reason is the great number of caryopses
produced (a caryopse is a small fruit with integuments, attached to the seed
and contained in a single grain, as in wheat).
Man has exploited this characteristic since ancient times, and civilisations
have always relied on extensive cereal crops which derive from environments
similar to arid meadows and which, when abandoned, evolve towards arid
meadows once again.
Gramineae caryopses are food for granivores and insects, especially ants
and ground beetles (Carabidae), which usually have a very different diet. Arid
meadows are generally inhabited by harvester ants of the genus Messor,
which gather and store grass
caryopses in their burrows.
These large ants are very conspicuous,
and were even mentioned in the Bible.
Their workers are very polymorphous,
i.e., the same ant-nest may contain
non-reproducing individuals of varying
sizes, some of which have enormous
heads. However, their duties are not
assigned according to the size of their
bodies or heads, but change during
their adult life.
These ants are extremely selective
when collecting their seeds: before
seizing them with their sturdy
mandibles and heading back to the
nest, they carefully examine the size,
and perhaps the possible future use of
the seeds. They may even discard 50
of them before making up their minds!
Although most ground beetles are
predators, the species living in arid
meadows are partially or totally
granivorous, at both larval and adult
stages. Among these are species of
the genera Ophonus, Carterus and
Ditomus.
Leguminosae are another large family
of plants in arid meadows. Their leaves
are also well protected from
phytophagous insects, although their
defence is not mechanical, as in
grasses, but chemical. Their tissues
often contain cyanogenic glycosides,
which are poisonous to most of their
potential eaters, both insects and
vertebrates.
However, there are groups of insects
which have developed defence
75
Harvester ants (Messor sp.)
Ophonus ardosiacus
Carterus cordatus
76
mechanisms against the harmful
effects of these plants: their digestive
tubes contain enzymes that turn toxic
substances into precious sources of
nitrogen. They can therefore exploit
food sources unavailable to their
competitors, and this explains the
great success of these species (also in
terms of numbers) - mostly butterflies,
Laemosthenus janthinus
moths and beetles.
Among lepidopterans associated with
leguminosae there are burnets (zygaenid moths), some of which are typical of
arid meadows. A day-flying type of insect - although they are relatives of
many twilight- and night-flying moth families, rather than true day-flying
butterflies such as cabbage butterflies, admirla, peacock and swallowtails their bodies have metallic blue shades and thin, tapering blue wings with
large, crimson spots.
They are gaudy not only for their colouring, but also because they spend their
days on shiny red or mauve flowers, such as those of scabious and
leguminosae, including those on which they spend their larval stages. In
addition to this, they are easily approached and touched, as they rely on their
poisonous tissues (due to the plants they feed on as caterpillars) that make
them inedible. Worthy of mention is Zygaena fausta, a western European
species living as far as the Valle di Susa in northern Italy, where its larvae live
on leguminosae of the genus Coronilla.
From the biogeographic viewpoint, arid meadows are often true “islands”
inhabited by individuals that come from far-away places. As regards arid
Alpine grasslands (and, to a much lesser extent Apennine ones), these are
typical steppe species, relatives of those living in the large arid areas in central
Asia and eastern Europe.
Among ground beetles, for instance, there are Laemostenus janthinus,
Carabus cavernosus, and several species of the genera Amara, Harpalus and
Cymindis.
In pre-Alpine valleys, thermophilous arid meadows such as those in Valle di
Susa and Val d’Adige, or xero-thermal hilly areas in the Veneto plain, like the
Euganean Hills, contain southern, sub-Mediterranean species, which find
their northernmost distribution area here. This is the case of some ground
beetles, embiids (web spinners) and the large poisonous centipede
Scolopendra cingulata.
77
Burnet (Zygaena) on Anacamptis piramidalis
■ Gastropod molluscs
78
The diversity and abundance of
terrestrial molluscs in arid meadows
largely depend on the nature of
substrates.
Calcium carbonate is extremely
important. Typical inhabitants are
Rumina decollata, a Mediterranean
species with an unmistakable shell.
In the early stages of its development,
this shell grows quickly from a rather
Rumina decollata
narrow apex, and after a short time,
the initial part including the first whorls,
is lost and replaced by a kind of calcareous plug, a diaphragm perpendicular
to the long axis of the shell: this explains the specific name of the mollusc.
Other large species in open arid areas are Chondrula tridens, typical of
impoverished meadows, Zebrina detrita, a mountain species found in southeastern Europe and on dry, sunny slopes in central and northern Italy
(eastwards as far as the Trieste Karst), and the calcicole species of the genus
Granaria, which generally lives on sunny, rocky meadows from the coasts to
the foot of mountains.
In the plain, often on plant stalks, are Candidula unifasciata, Cernuella
cisalpina and Xerolenta obvia. The last is only found from Friuli to Lombardy. In
the Apennines, arid meadows typically host Cernuella neglecta and Candidula
spadae.
In arid grasslands, molluscs may lead a less conspicuous life, either at the
base of plants or inside the soil. Among these are the tiny species of the genus
Truncatellina, T. cylindrica, and the rarer T. callicratis, T. claustralis and T.
monodon, Vallonia costata and Pupilla muscorum. Among the species of the
genus Vertigo, those better adapted to less humid biotopes are V. pygmaea,
found in the plain, and V. alpestris, which lives in the mountains.
■ Centipedes and scorpions
In arid meadows, the soil is not suitable for centipedes, myriapods which
prefer cooler environments with greater quantities of litter, branches and twigs
on the ground, although some lapidicolous species (living under stones) may
locally abound. These habitats are particularly suitable for scolopendras, the
bite of which is very painful, although
not dangerous for man. In Sardinia,
there is the smaller Scolopendra
oraniensis which, in Sicily and
southern Italy, lives with the larger
Scolopendra cingulata (up to 10 cm
long). The latter may be found as far as
northern Tuscany and, eastwards, in
the Marches.
Isolated populations also live in the
warmer, southern flanks of the
Euganean Hills. In late winter, young
Scolopendra cingulata
scolopendras with greenish or brown
bodies and bright red heads which
fade during the summer, may be found under the stones that dot arid
meadows. Later, the same environment will host females with their young.
Scolopendras usually lay two or three dozen eggs in a single cluster around
which they wrap themselves, protecting it for several weeks, during which
time they fast. They return to their normal lives only after their eggs hatch.
In arid meadows, another genus of large arthropods living under rocks - and
not only there - is known to provide more complex parental care: scorpions.
The females retain the eggs in their bodies until the young are born, and then
the delicate, whitish, very easily dehydrated young climb on their mothers’
backs, where they stay for several days. The main benefit which the young
scorpions derive from this habit seems to be the humidity they obtain from
their mothers’ bodies.
■ Millipedes
Returning to myriapods, arid environments also host a few millipedes, but for
reasons unlike those of centipedes. Centipedes are predators that catch their
prey with their poisonous appendages, whereas most millipedes feed on dead
leaves or decaying wood, materials which are unavailable in arid meadows.
However, the bases of grasses growing attached to rocks locally host large
specimens of the genus Pachyiulus, with cylindrical bodies supported by
several dozen feet and ready, if touched, to emit great quantities of quinones
from their repugnatorial pores. Quinones are a family of reddish-orange
organic substances that stain the skin for lengthy periods and may greatly
irritate mucous membranes.
79
meadows. Among these are very large and gaudy hymenopterans (bees,
wasps), including scoliid wasps, which are attracted by the numerous flowers
in arid grasslands, but which spend their juvenile lives as parasites on the
larvae of rhinoceros beetles which, in turn, had settled inside old decaying tree
trunks in the woodland nearby.
Many hymenopterans therefore nest on the margins of meadows, where the
first shrubs grow. They collect food in arid meadows and then carry it to their
young.
80
A crab spider (Misumena vatia) preying on a butterfly (left) and another crab spider on Aceras: this
thomisid mimics yellow or white, according to the plant on which it sits in ambush
■ Spiders
Blattarians. Cockroaches are an order of insects mostly living in tropical
regions, although small numbers of species, carried by man, reach temperate
regions and settle in towns. A few, locally abundant species live in arid
meadows, such as Lobolampra subaptera, which lives under stones or the
plant debris on which it feeds. Its name derives from its very short wings,
which make adults look like young that still have to develop their flight
appendages. Similar to this species is the congener L. decipiens, which also
lives in sparse, thermophilous woodland, and is typical in xerothermic habitats
in northern Italy.
Another characteristic cockroach is Phyllodromica marginata, which has an
unmistakable white line circling its small black body.
For a few months each year, spiders abound. The structure of arid vegetation,
which is short and almost exclusively grassy, greatly hinders orb weavers, i.e.,
those which spin the typical sunburst cobwebs, but it does enable the tiny
linyphiids (sheet-web weavers and dwarf spiders, which constitute the spider
family with the highest number of species in Italian fauna, living in grassy
environments) to weave their typical horizontal sheet webs with non-sticky
silk. In spring, when grasses and herbs are visited by many insects, especially
those living on flowers, arid meadows host small wandering spiders which do
not weave webs (philodromids and jumping spiders), but which have the habit
of pouncing on their tiny prey.
■ Insects
Before discussing insects, a preliminary clarification must be made. There are
insects which spend most of their lives on the plants in arid meadows, such as
many orthopterans (grasshoppers, crickets), aphids (plant lice), cicadellids
(leafhoppers) and bugs. Others live on vegetation only as adults, because they
live elsewhere as larvae, perhaps in scrub or even woodland near arid
Large numbers of flowers are extremely attractive to many insects
81