Invertebrates: taxonomy

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Invertebrates: taxonomy
■ Cliff fauna
Parnassius apollo
Molluscs. Although most Italian species
live in humid, shady areas protected by
vegetation, some terrestrial molluscs are
adapted to hostile environments like
rocky cliffs. Among these are numerous
species of the prosobranchs, the most
primitive group of gastropods, with many
marine and freshwater species, and
pulmonates, the most advanced group,
which features a type of lung and has
developed along the evolutionary line of
the Stylommatophora even in subaerial
environments. Despite the fact that these
two groups have differing anatomy and
physiology, they are both exclusive to
limestone at all mountain altitudes. They
live on exposed surfaces, both in the
shade and in direct sunlight, generally
insinuating themselves into crevices,
cracks, underneath jutting stones,
mounds, between mosses, and indeed
wherever they can easily find food and
be protected from excessive heat and
cold. Their shells are of different sizes,
and their cryptic colours reflect the sun’s
rays and reduce excessive heat: they
may be greyish-white with yellow, violet
or pinkish hues. Shells can be ribbed, to
reflect sunlight better, and covered with
soil or hardened excrement, which
enhance protection against heat and
possible predators. Molluscs feed on
encrusting vegetation (moss, lichen) or
plant debris that accumulates inside
cracks or in the scarce soil on terraces.
Despite their resistance to drought, their
Paolo Audisio
foremost limiting factor is lack of water
supply, which they are able to
circumvent in various ways. They have
specially adapted excretory apparatuses
that retain water, and thick shells that
enable them to seal themselves from
within by means of mucus. They can
extend their dormancy for lengthy
periods, thus limiting their excretions to
a minimum, in order to survive on highly
exposed cliffs, where the only water
supply is provided by night-time
Pyramidula attached to a rocky surface
condensation. In addition, many mollusc
species are resistant to the cold, and
can colonise high-altitude cliffs.
From the faunal and biogeographic
viewpoints, Italian cliff molluscs are very
interesting, thanks to their great
diversity. Their fragmentation into typical
local populations and their isolation
have given rise to great morphological
diversification and taxonomic
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fragmentation, which have not yet been
thoroughly analysed.
Among the prosobranchs, typical cliff
species are those of the genus
Cochlostoma, with small, usually ribbed
conic shells (about 1 cm high),
occasionally smooth. The group lives
everywhere in mainland Italy and on the
major islands, and is divided into
various species, many of which are
typical of small areas. Among these is
Cochlostoma canestrinii, the largest
Italian species endemic to the Presolana
chain in the Bergamasque Alps;
C. porroi, endemic to the Lombard and
western Trentino Pre-Alps; C. villae in
the Euganean and Berici Hills and PreAlps near Vicenza and Verona;
C. philippianum and C. henricae in the
Carnic and Julian Pre-Alps respectively;
C. scalanum and C. tergestinum in the
Trieste Karst; C. crosseanum in the
Tuscan Apennines; C. sardoum in the
limestone chains of central-eastern
Cochlostoma villae
Sardinia, and C. paladilhianum in northwestern Sicily.
Stylommatophora species of differently
evolved genera are far more numerous.
Among the most primitive is Pyramidula
(pyramidulid family), with the species
P. pusilla, found at various altitudes all
over Italy. It has a very small, dark red
shell a few millimetres across, shaped
like a top, which is generally streaked
and calcified. The shells of the genera
Rupestrella and Chondrina, both
belonging to the chondrinid family, are
larger (3-12 mm high). These species
have similar ecology, although the
former are typically Mediterranean and
live at low altitudes, and the latter can
tolerate cold temperatures and may live
at higher altitudes. Chondrina avenacea
is very common in Italy in the Alps,
Apennines, and the largest Italian
islands, and has several endemic
subspecies. In central-eastern prealpine areas, it is usually replaced by
Chondrina avenacea
similar species, like C. clienta and C.
megacheilos. The Apuan Alps host an
endemic species, C. oligodonta. The
chondrinid family includes another
genus, Solatopupa, whose whitish shells
are similar to those of the Chondrina
species, but larger (8-15 mm high);
Solatopupa similis is widespread (from
northern Spain to Liguria); S. juliana is
typical of Tuscany, S. pallida is only
found in a restricted area of Liguria,
S. psarolena in the Maritime Alps and
S. guidoni in Corsica, Sardinia and Elba.
The clausilids include typical cliff genera
with thick, robust, fusiform shells about
10-20 mm long, smooth or ribbed,
generally whitish, with toothed exterior
apertures closed by a rodlike structure
called clausilium. Leucostigma
candidescens is found along the
Apennines, even at high altitudes.
Its shell may be of various sizes and is
generally white. It is easily confused
with species of the genus Medora, a
trans-Adriatic group living in the
Apennines from the Marches to Calabria
and on the Gargano (the “spur” of the
Italian peninsula). Many populations
similar to the Dalmatian Medora
albescens colonise the Apennines, and
other populations, which are more
similar to another Dalmatian species,
M. dalmatina, live on the eastern flank of
Monte Pollino.
Other cliff species of this family live in
southern Italy and Sicily, most of them
belonging to the genus Siciliaria. They
are not described in this volume
because they are found at low altitudes.
The family Helicidae (helix snails)
includes the largest number of montane
and sub-montane cliff species, like
Chilostoma and Macularia. Chilostoma
had several species - in the past
described as separate - which today are
all included under the name Chilostoma
cingulatum. They live between the
southern Alpine versants, Pre-Alpine
zone and Apennines as far as the
Matese massif. They probably reached
the Apennines during a Quaternary
glaciation and, when more favourable
climatic conditions set in, they migrated
to higher altitudes on the limestone
mountains, thus becoming fragmented
into isolated populations with local
differentiation. The situation of the
genus Marmorana is similar, although
this Tyrrhenian taxon is typical of
medium altitudes with Mediterranean or
sub-Mediterranean climate.
One of the most common species is the
xerophilous Marmorana muralis, which
was spread by man in several areas in
Italy and the Mediterranean basin.
Western Sicily hosts a number of
interesting snails with overlapping
distribution, like Marmorana globularis,
M. platychela and M. scabriuscula.
M. nebrodensis is found in northeastern Sicily, between the Madonie and
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Peloritani mountains, and
M. fuscolabiata in western Sicily (Monti
Peloritani) and in the Apennines
between Calabria and Campania.
M. signata is very similar to this last
species, and lives in the centralsouthern Apennines from Umbria to
Matese. A few small southern areas in
Tuscany (Monte Amiata, Monti
dell’Uccellina, Argentario, Isola del
Giglio) host Marmorana saxetana.
Crustaceans. Isopods are almost totally
absent on pebbly and rocky substrates
lacking true soil, and it is therefore no
surprise that, although they are
extremely common in other terrestrial
environments, especially at medium-low
altitudes, none of them typically lives on
rocky cliffs. A few species of the large
genus Armadillidium, together with other
Armadillidium sp.
nocturnal genera, may find temporary
refuge at the base of cliffs and rocky
walls, especially near crests and
terraces and in rock crevices, where soil
deposits favour their occasional
colonisation.
Millipedes. Millipedes (diplopods) are an
important group of terrestrial arthropods,
with many mountain species. Just like
several other groups which are not
exclusively phytophagous, there are no
rock-cliff specialists, although a few
orophilous species associated with dry,
rocky grasslands may also be found at
the foot of rocky cliffs, in crevices and
near crests. Among these is the pillworm
Glomeris helvetica (glomerids), a circumalpine species living in isolated locations
which, in the Italian Alps, is actually
frequently found on rocky and woody
debris in landslide deposits in narrow
valleys at the base of cliffs, from low
altitudes to 2500 m. Among
polydesmids, Brembosoma
castagnolense is an Alpine endemic
found in Switzerland and the Lombard
Alps, where it lives on limestone in steep
environments up to 2500 m.
The julid Hypsoiulus alpivagus, another
circum-alpine endemic found in isolated
locations, generally prefers alpine areas
to 2800 m, but it can also live at
montane levels in deep gorges and cave
entrances. It seems to be closely
associated with sedimentary rock, and
its petrophilous behaviour can be
inferred from its favourite locations:
screes, rock debris and boulder
deposits, slabs, cliffs and river valleys.
The julid Leptoiulus (Kolpophylacum)
helveticus has similar geographical and
altitudinal ecology and distribution, and
the Apennines host other species with
analogous ecological requirements.
Centipedes. Centipedes (chilopods), like
all walking predators, do not find
favourable living conditions on mountain
cliffs. The only species which often finds
Orthopterans. Although many
species of orthopterans may
occasionally reach mountain cliffs and
faces, very few are regular visitors.
Among them is Stenobothrodes
cotticus, endemic to the Western Alps,
which colonises the margins of screes
and bare rocky walls, and is often found
on cliffs with poor grass covers
between 2000 and 2800 m.
Several species of the genus
Glyptobothrus can also live in these
environments, with large populations of
G. mollis ignifer in the Piedmont Alps.
Antaxius difformis and Chopardius
pedestris (ensifers) also live at high
altitudes, on cliffs and landslide
deposits in the Alps. In the centraleastern Alps, A. difformis lives
exclusively in montane environments, in
sometimes large communities.
A subspecies of C. pedestris, C. p.
apuanus, is found in the Apuan Alps.
●
Lithobius lucifugus
refuge at the base of Alpine cliffs is the
common, euryecious Lithobius lucifugus,
which is far more frequent in Alpine
grasslands, grassy and rocky habitats,
and forests at medium-low altitudes.
Insects. Most insect species are
phytophagous, with numerous
microphages living on rough substrates,
and only a few of them are predators or
parasites.
Glyptobothrus mollis ignifer
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Chopardius pedestris
Although it may live at lower altitudes in
a wide range of habitats, it is typical of
mountain cliffs. Isolated individuals of
many other orophilous and
xerothermophilous species can be
found basking in the sun at high
altitudes on well-exposed cliffs, like
many tettigoniid long-horned
grasshoppers of the genus
Anonconotus and numerous acridid
grasshoppers.
Heteropterans. Their great
capacity for adapting to various
types of food enables many
phytophagous bug species to
colonise montane cliff
environments. The assassin bug
Coranus subapterus (reduvids,
see drawing) is a predator living
on the ground, in interstices
between stones and rocks.
It is widespread in Europe and in
the Mediterranean basin, and is
found on the Italian mountains.
Euryopicoris nitidus (mirids) is a
phytophagous species which develops
on cliff plants of the pea family.
●
Glyptobothrus alticola
It is a Euro-Siberian species that, in Italy,
is only found in a few montane locations
in the Alps, Emilia-Romagna, Lucania and
Calabria Apennines. Dimorphocoris
poggii is a recently discovered plant bug
endemic to the Ligurian Alps. It lives on
xerophilous cliff grasses between 1600
and 2000 m. Plagiotylus ruffoi belongs to
the same family, and is a Sicilian endemic
associated with the pulvini of Astragalus
siculus (pea family), a cliff species
growing above the treeline on the
Madonie cliffs. Another typical species is
the lace bug Acalypta musci (tingids),
which inhabits high-altitude mountains in
Europe, the Alps and a few Apennine
locations in Italy. It grows on mosses
which develop along cliffs and at their
base, generally in shady, humid areas up
to 2300 m. It may also be found at lower
altitudes on the mountains, on tree
mosses growing on the trunks of conifers.
Steep, sunny cliffs all over Italy host the
peculiar genus Copium, a tiny
phytophagous lace bug that produces
galls (excrescences) on the calyxes and
corollas of small cliff
germanders. These bugs can be
identified by the last two
segments of their antennae, which
are huge. Lastly, the alydid bug
Alydus rupestris, an Alpine
endemic typically found at 2200
m at the base of cliffs, grows on
Vaccinum and Empetrum.
● Coleopterans. Beetles, together
with lepidopterans, are the most
numerous insects stably colonising
rocky cliffs.
Ground beetles (Carabidae). Although
Italy hosts more than 1300 species of
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ground beetles, most of which prey on
other invertebrates, only a few of them
inhabit mountain cliffs. It is actually hard
to find any one species regularly visiting
these habitats, which are very
Carabus (Orinocarabus) pedemontanus
inhospitable to “walking” insects. Some
Carabus of the subgenus Orinocarabus,
especially C. bertolinii in the eastern
Alps, may be caught climbing rocky
walls. In the Alps and Apennines, a few
tiny species of the genera Microlestes
and Syntomus frequently explore the
surfaces of cliffs, walls and huge
boulders. A few rare Ocys may be found
in rock and cliff crevices.
Sap beetles (Nitidulidae) and shortwinged flower beetles (Cateretidae). Sap
beetles and short-winged flower beetles
are adapted to various types of food, and
may be phytosaprophagous and
mycetophagous as well as
anthophagous and spermatophagous.
Many of them are exclusive to
montane and sub-montane cliffs.
Many of the most important Italian
beetles (including some endemics
and sub-endemics) may be
considered typical of these
environments. Among the most
frequent species are some
anthophagous sap beetles of the large
genus Meligethes, like M. subfumatus
(see drawing), which is endemic to a
small montane and sub-montane area
near the Maritime Alps between France
and Italy. Its larvae feed exclusively on
Lavandula angustifolia (the same
lavender used in cosmetics), between
low altitudes and 2000 m. Another
species, M. nuragicus, is endemic to
Corsica and Sardinia, where it lives only
on silica cliffs. There, at medium
altitudes, it is monophagous on the
germander Teucrium massiliense.
Meligethes lindbergi, an Italian subendemic, is very xerothermophilous and
monophagous on the common Teucrium
flavum, especially on steep, sunny
limestone slopes of inland Liguria and
Trieste, along the Italian peninsula and in
Sardinia and Sicily, where it can be found
up to 1500 m. The rare Meligethes
fumatus is also xerothermophilous, and
lives on warm mountains and hills from
Val d’Aosta to Calabria. It is associated
with winter savory (Satureja montana),
which is colonised by larvae and adults
in periods that are unusual for other
members of the genus, most of which
have spring phenology. Its breeding
period stretches from the second half of
July to mid-October. In spring, other
Meligethes are associated with various
small cliff crucifers that typically grow in
cracks between cliffs and in shady areas
at the base of rocky slopes. Among
these is Meligethes subaeneus, an
oligophagous species locally
associated with Cardaminopsis
spp. in north-eastern areas, the
sub-endemic M. spornrafti, which
lives on Arabis spp. between the
Western Alps and Calabria, and
the rare sub-endemic M. lunariae,
monophagous on honesty (Lunaria
annua) in interfacing habitats between
steep cliffs and poor woodland between
central Italy and Calabria, up to 1500 m.
Another species associated with
xerophilous cliff crucifers is the rare,
recently described M. erysimicola which,
in southern Europe, lives on plants of the
genus Erysimum (wallflower). This beetle
is found in the Alps and Apennines on
cliffs and high ridges above 2300 m.
Montane cliff environments on the
eastern flank of the Gran Sasso (up to
1500 m) host the rare M. tener, an
Anatolian-Balkan species (also found at
lower altitudes in the Gargano), which is
oligophagous on xerophilous cliff species
of the genus Aurinia. Similar sunny cliff
habitats on serpentine are inhabited by
the very rare M. chlorocyaneus, whose
hosting plant is still unknown (it is
certainly a cliff plant of the cabbage
family) and which lives in isolated relict
areas on the Bulgarian mountains,
Bosnia, southern Austria, Slovenia and,
in Italy, only in the valley of Stura d’Ala in
western Piedmont, north-west of Turin,
to 1500 m. Montane belts in the southern
Apennines (from Cilento to Calabria) and
Sicily host one of the most typical
species of these habitats, the subendemic Meligethes scholzi, closely
associated with Ballota rupestris (mint
family) (from sea-level to 2000 m) on
limestone cliffs and in warm river valleys.
Among short-winged flower beetles
(Cateretidae), the most common cliff
inhabitants are anthophagous species of
the genus Brachypterolus (which are all
associated with plants of the figwort
family). Examples are B. vestitus, found
on cliffs in sunny montane and sub-
montane zones of north-western Italy on
the plant Antirrhinum latifolium, which
usually grows in southern France and
Spain, and the more frequent B. linariae,
found on Kickxia spp. and Linaria spp.
Soft-winged flower beetles (Dasytidae)
and malachiid beetles (Malachiidae).
These two small families of tiny beetles
(Cleroidea superfamily) include a large
number of anthophagous species living
in warm, sunny environments. However,
very few have adapted to montane cliff
habitats and, in Italy, only xerophilous
species may be found at low altitudes (in
southern Europe, a few species adapted
to medium-high altitudes live on Corsica
and in the Pyrenees). For example, the
soft-winged flower beetle Danacea
nigritarsis reaches the summits of the
Monte Pollino massif in the southern
Apennines, where it can be found on cliff
saxifrages, although it is generally
typical of coastal and sub-coastal
environments. The Bergamasque PreAlps (from Canton Ticino to Adamello)
host Dasytes lombardus, a high-altitude
species associated with arid meadows.
Numerous individuals may be found on
the yellow flowers of several plant
families, as well as on rocky slopes and
cliffs. Although the species is not usually
associated with steep substrates, it
seems to prefer these conditions.
Blister beetles and oil beetles (Meloidae).
Blister beetles include a few hundred
European species, half of which live in
Italian steppe environments, where their
larvae feed on grasshopper eggs. Some
of those belonging to the oil beetle genus
Meloe and to the nemognathine
subfamily, are associated with bees, in
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whose nests their larvae develop,
especially at low altitudes. The most
common is the nemognathine Sitaris
muralis, which is parasitoid on woodboring bees (Anthophoridae) and lives on
rocky-earthy cliffs and dry walls along
the coastline, reaching Italian
xerothermic mountains inland, up to
1000 m and colonising isolated, sunny
cliffs and river gorges. Occasional
visitors to the same environments are
two rare Italian species of the genus
Stenoria, S. analis and S. apicalis,
associated with xerothermophilous cliff
plants of the mint family. They seldom
reach high altitudes (only in southern
Italy) and are parasitoid on plasterer bees
(Colletidae). Zonitis and Euzonitis have
similar ecology and may be found
throughout Italy and Sicily. Z. nana,
Z. flava, Euzonitis terminata and
E. quadrimaculata are parasitoid on leafcutting bees (Megachilidae).
Buprestis splendens
Scarabaeoid beetles (Scarabeoidea). This
group includes a large number of beetles
with various feeding habits (generally
coprophagous and phytophagous).
They visit mountain cliffs only
occasionally, because these areas are
inhospitable for them. Except for a few
coprophagous aphodiids which follow
grazing goats, the only species that are
frequently found in these high areas are
a few leaf-chafers (Melolonthidae) of the
genera Rhizotrogus and Amphimallon
and, sometimes, Anoxia (e.g., A.
australis in warm cliff locations in the
Alps and western Pre-Alps). In summer,
they drone at twilight around shrubby
pines (Pinus spp.). There is also the
common, euryecious chafer Serica
brunnea.
Metallic wood-boring beetles
(Buprestidae). There are only a few
wood-boring beetles in montane cliff
environments, as their presence is
associated with the local availability of
trees and shrubby plants clinging to
exposed, steep cliffs, where these plants,
weakened by the harsh conditions, are
easily attacked by xylophagous beetles.
In addition to numerous species of small,
xerophilous and euryecious beetles of
the genus Anthaxia, often associated
with mountain pines, there is the larger,
shiny Latipalpis plana, which lives on
xerophilous oaks at medium altitudes on
sunny cliffs. Also worthy of mention is
the well-known, although rare Buprestis
splendens, a symbolic species of nature
conservation in Italy and Europe (being
of European Community interest). It lives
in relict locations in Europe, and in Italy
at montane levels in the National Park of
Pollino, between Basilicata and Calabria.
In Italy, this fascinating species is
associated with the rare, endangered
Bosnian pine (Pinus leucodermis), a
symbol of the Park itself and one of the
most typical tree of the CalabrianLucanian Apennines, where it grows on
sunny, rocky cliffs and crests.
Leaf beetles (Chrysomelidae). Leaf
beetles belong to an important family of
phytophagous beetles that do not
generally colonise mountain cliffs and
rocky slopes (they are usually found at
low altitudes on Mediterranean cliffs).
Among them are a few flea beetles
(Alticinae), such as Psylliodes toelgi,
which lives in southern Europe and in
the Alps (between Piedmont and Friuli)
and is associated with orophilous,
glareous or cliff crucifers of the genus
Biscutella. The same plants are
sometimes colonised by Phyllotreta like
P. atra and P. ganglbaueri. The congener
Psylliodes picipes is only found in the
eastern Alps, where it feeds on several
species of cliff crucifers.
Snout beetles (Curculionidae and
Apionidae). Snout beetles belong to a
large superfamily of phytophagous
beetles and include some of the most
important mountain cliff species. Among
long-beaked weevils (Apionidae),
Osellaeus bonvouloiri is typically found
on high-altitude Alpine ridges and cliffs,
feeding on saxifrages. It has two
subspecies, O. bonvouloiri bonvouloiri in
the western Alps (Italy and France), and
O. bonvouloiri baldensis in the Veneto.
Very important among snout beetles are
members of the genus Dichotrachelus,
with about 15 endemic and sub-endemic
species on various mountains and
valleys in the Alps. They are all
associated with Saxifraga spp., and
usually colonise steep slopes and cliffs
and medium-high altitudes. Another
species associated with yellow saxifrage
(Saxifraga aizoides) is the circum-alpine
Pelenomus hygrophilus. Cliffs at lower
altitudes in the Apennines host, among
many others, Ceutorhynchus pinguis, a
rare Apennine endemic that develops on
Alyssum diffusum (cabbage family), and
Ceutorhynchus verticalis (note the
specific name), an endemic of the
southern Apennines associated with
another plant of the cabbage family,
Aurinia saxatilis ssp. orientalis.
● Hymenopterans. Sunny cliffs and river
gorges are optimal breeding grounds for
many species of bees, especially boring
ones if the bedrock geology is limestone
or sedimentary rock, which is of more
recent age and less compact. Many
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species of leaf-cutting bees, woodboring and mining bees are regular
visitors to these habitats, although they
generally prefer lower altitudes.
Ants are not very common on mountain
cliffs, and only a few workers (especially
those of euryecious species) explore
rocky slopes. There are, however, the
tiny members of the genus Leptothorax,
which find favourable microhabitats in
cracks in cliffs, where soil is sufficient for
them to build their nests.
● Lepidopterans.
Both day- and nighttime flying lepidopterans include several
typical although not exclusive species
inhabiting mountain cliffs and slopes.
The papilionid Papilio alexanor (a
species of European interest), is found in
discontinuous Italian areas on xeric cliffs
up to 1000 m. Its larvae feed on
umbellifers of the genus Trinia. The wellknown parnassian Parnassius apollo (a
Erebia montana
species of European Community
interest) is often found in cliff
environments, and this is especially true
of populations of the Aspromonte
(P. apollo pumilus), as they are
associated - as larvae - with small plants
of the stonecrop family, especially those
of the genera Sedum and Sempervivum.
Among whites (pierids) there are species
of Anthocharis, like A. euphenoides,
found in rocky areas up to 2000 m both
in the Alps and Apennines, Pieris
callidice, which lives at high altitudes on
the Alps, and Euchloe species,
especially E. bellezina, found on coastal
hills in north-western Italy. Among blues
(lycaenids) are a few Agrodiaetus,
especially A. ripartii that lives on sunny
rocky slopes in the western Alps and is
associated with small cliff plants of the
pea family (genus Onobrychis), and A.
galloi, endemic to the Pollino group.
Satyrins are the best represented
nymphalid butterflies on mountain
slopes, and the larvae of many species
are associated with xerothermophilous
cliff grasses. Among the numerous,
typical species of these habitats, which
are also found in other montane
environments, there is Satyrus ferula
(Alps and Apennines), S. actaea (western
Alps), Chazara briseis (throughout Italy
except for Sardinia), Pseudotergumia
fidia and Erebia scipio (western Alps),
Hipparchia alcyone (north-western Italy
and central-northern Apennines), H.
statilinus (throughout Italy, except for
Sardinia), H. neomiris (Sardinia), Erebia
styx, E. styria and E. calcaria (eastern
Alps), E. christi (Piedmont), E. montana
and E. meolans (Alps and Apennines),
and the widely distributed Hyponephele
lycaon. The nocturnal species of this
order are ten times more numerous than
the daytime ones, the greatest family
being that of owlet moths (noctuids),
with several species and sub-families.
Among them are Euxoa decora,
Standfussiana lucernea, Epipsilia
grisescens, Rhyacia helvetina, Chersotis
ocellina, C. alpestris, C. oreina, Spaelotis
senna and Megasema ashworthii. Owlet
moths are closely associated with cliff
environments, and change colours to
camouflage on the substrate. They can
therefore be totally white when resting
on limestone rocks, grey on schist and
melanotic (black) when on lava.
Several Lychnis moths (Hadeninae) are
frequently found in mountain habitats,
like Hadula odontites, H. melanopa (a
daytime boreo-alpine species), Sideris
kitti, Antitype suda and many species of
Hadena, which are generally associated
with plants of the pink family, such as
Silene, Lychnis and Dianthus.
In the Alps above the treeline are plusiid
moths adapted to daytime flight, like
Syngrapha devergens, and
S. hochenwarthi. Although not exclusive to
the mountains, large numbers of
Bryophylinae are often found in these
areas (e.g., genus Cryphia). Their larvae
are lichenophagous, like those of
numerous tiger moths (arctiid moths) of the
sub-family Lithosiidae, such as Nudaria
mundana and species of the genus Setina.
The cryptic measuring worm moths
(geometrid moths) are also very numerous,
like the twig caterpillars (Ennominae) of
what used to be the genus Gnophos (e.g.,
G. obfuscatus) - now divided into several,
more specific genera – those of the genus
Dyscia (D. raunaria, D. sicanaria) and a few
Larentinae, especially those of the genus
Eupithecia, like E. venosata. Typical of the
mountains are also heliophilous (sunloving) pyralid moths of the genus
Metaxmeste. These black insects bask in
the sun with their wings open. Other
frequently found Alpine pyralid moths are
those of the genus Orenaia and Catharia
pyrenaealis.
Bagworms (psychids) of the genera
Dahlica, Pseudobankesia, Luffia,
Leptopterix and Oreopsyche build their
elaborate larval cases on the mountain
rocks just before pupating in them. There
are also clothes moths (tineids) of the
genus Eudarcia, the larvae of which feed
on green algae, and a few Chamaesphecia
- wasp moths (sesiids) associated with
typically montane spurges.
Lastly, the curious habit of the small
diurnal hawkmoth (sphingids)
Macroglossum stellatarum is worth
mentioning: it carefully inspects rocky
faces in search of tiny cracks to spend
the winter.
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■ Scree fauna
98
Molluscs. Terrestrial gastropods colonise
mountain screes according to the
morphological characteristics and
maturity of these environments. Unstable
screes with scarce vegetation, mainly
composed of small rocks (pebbles and
gravel) generally host very poor
malacofauna without typical species.
However, screes with alternating pebbles
and large boulders, especially if
colonised by sparse grass and shrubs,
are ideal habitats for molluscs, the
diversity of which increases with the
complexity of the environment.
Interstices between clasts, deposits of
debris in the cracks or at the base of
shrubs, niches where humidity collects
are all ideal places for the colonisation by
terrestrial gastropods. Of the very limited
few which, despite not being exclusive to
them, live in these ecosystems there are
Granaria stabilei and Candidula
unifasciata, found on grassy screes,
Tandonia simrothi and some species of
vitrinas of the genera Eucobresia,
Phenacolimax and Oligolimax, which are
usually found under the rocks of stable
screes. The species living in these
environments are also influenced by the
Oedipoda germanica
Chilostoma cingulatum colubrinum
microclimate of the local altitude, and by
bedrock (calcareous or siliceous). For
instance, a few subspecies of
Chilostoma cingulatum, C. alpinum,
Cepaea sylvatica and Arianta
chamaeleon are found on limestone at
high altitudes, whereas Chilostoma
millieri and C. zonatum prefer siliceous
substrates. The extension and location of
screes also affect malacofauna
components: small screes often host
populations of nearby ecotones like
rocky faces, pastures and even
woodland. Their populations are
therefore varied, although lacking
characteristic species of these habitats.
Crustaceans. Isopods are almost totally
absent from mountain screes, due to lack
of sufficient soil. However, a few
orophilous night species like those of the
genus Armadillidium, may occasionally
be found under bare rocks and at the
margins of debris slopes, where enough
soil accumulates.
Millipedes. Millipedes are an important
group of terrestrial arthropods, and many
of them live in montane habitats. Although
none of them is exclusive to screes, a few
orophilous species associated with arid,
rocky grassland may be found at the base
of debris slopes with scarce vegetation
and rough surfaces. Among these is
Pseudocraspedosoma grypsichium
(neoatractosomatids), a circum-alpine
endemic with wide ecological tolerance
that frequently colonises the margins of
screes and steep rocky-pebbly slopes up
to 3000 m. Bergamosoma canestrinii
(craspedosomatids) is also found in the
Alps on shallow soil and on rocky debris,
as well as on south-facing open areas
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(warm, rocky grassland on the mountains)
up to 3000 m, together with other
members of the same family with similar
distribution and ecology (e.g.,
Atractosoma meridionale,
Dactylophorosoma nivisatelles,
Rothenbuehleria minima), the polydesmid
Polydesmus monticola, and the julids
Leptoiulus alemannicus and L. riparius.
Julid millipede
Centipedes. The only species of
centipede that frequently colonise the
margins of Alpine screes with sufficient
soil are a few lithobiids, like Lithobius
lucifugus which, in the Eastern Alps, lives
together with L. schuleri. Although
L. lucifugus is typical of these areas, it is
also found in arid Alpine pastures and
montane forests. In the Dolomites, the
same habitats host L. muticus and
L. nodulipes, whose wider ecological
tolerance makes them less significant.
Insects. The most important orders of
insects colonising mountain screes are
coleopterans, lepidopterans,
orthopterans, heteropterans, dipterans
and hymenopterans. Although most of
them are phytophagous, there are
several microphagous species on rough
surfaces. There are only a few predators
and parasitoids.
● Orthopterans and closely related
orders. Although orthopterans, especially
grasshoppers, are very numerous on
mountain screes, few are exclusive to
this habitat. The group includes several
families (particularly Catantopidae,
grasshoppers and a few pygmy
grasshoppers) with species closely
associated with xeric, rocky substrates
colonised by the pulvini of pioneer plants
and orophilous, xerophilous grasses (the
traditional ecology of Italian orthopterans
includes them in the “field-lapidicolous”
community). The most frequent species
on the scarce grass growing on screes,
especially near Seslerietum (moor grass),
juniper and rhododendron, are the
pygmy grasshopper Tetrix bipunctata
(Tetrigidae) (southern Europe, Alps), very
difficult to see when it rests on greyishwhite rocks up to 3000 m, and
Catantopidae of various genera, like
Epipodisma pedemontana (western
Epipodisma pedemontana
Alps), many Italopodisma (all endemic to
small high-altitude areas in the central
Apennines), Chorthopodisma cobellii
(endemic to north-eastern Italy), species
of the genus Podisma (with Alpine and
Apennine species) and especially
Melanoplus frigidus, one of the most
cryophilic species best adapted to life on
Alpine rocky substrates. The common,
euryecious Calliptamus italicus (widely
2000 m), and the common Oedipoda
caerulescens and O. germanica (to 2000
Oedipoda caerulescens
Calliptamus italicus
distributed in southern Europe and Italy)
may reach mountains in southern Italy.
Sub-montane levels are colonised by
Calliptamus siciliae (Catantopidae) (in the
Mediterranean basin and as far as
Manchuria) and Pezzotettix giornai (a
European-Mediterranean species).
Although the two latter species may be
included in the “field-herbicolous”
community, they are frequently found on
the mountains along rocky paths and at
the margins of debris slopes, where they
cohabit with grasshoppers of the genus
Oedipoda (Acrididae).
Grasshoppers are frequent, although
not exclusive inhabitants of these areas,
and find favourable living conditions
on Alpine, pre-Alpine and Apennine
screes.
Among the many species are Psophus
stridulus (Alps and central Apennines, to
m), cryptic species whose typical brightly
coloured metathoracic wings open
suddenly during their brief flights,
disorienting potential predators and
trekkers, who are caught by surprise.
Stenobothrodes rubicundulus (southern
Europe) colonises the Alps and central
Apennines to 2000 m, together with the
similar S. cotticus and S. fischeri (in
isolated locations in the Alps, Pre-Alps
and Apennines as far as Calabria). Then
there are Aeropus sibiricus (found
throughout Italy), Glyptobothrus
binotatus daimai (Western Alps) and
G. alticola (in Italy, along the Alps and
central-eastern Pre-Alps), G. eisentrauti
(Alps) and Aeropedellus variegatus, one
of the most typical species in these
Alpine habitats (especially in central and
western Italy, to 2500 m). The common
Glyptobothrus brunneus brunneus may
also reach high altitudes, especially in
the central-southern Apennines, where it
is abundant in rocky, pebbly areas.
Although they are not regular inhabitants,
large numbers of Myrmeleotettix
maculatus may locally be found on
Apennine screes and in the Western
Alps. Among Ensifera, the katydids
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(Tettigoniidae) Antaxius difformis and
Chopardius pedestris typically colonise
mountain screes, together with species
of the genus Anonconotus. In Sardinia,
some katydids of the genus Rhacocleis
live in similar environments on the
highest summits of the island, like Monte
Corrasi, Bruncu Spina and Limbara. The
earwig Forficula apennina (dermapterans)
is an endemic species typically found on
the coldest Apennine screes.
Other colonising earwigs are Chelidura
aptera (Western Alps) and Anechura
bipunctata (Western Alps and the Gran
Sasso). Worthy of mention is the
cockroach Ectobius montanus, which
may live on the few grasses colonising
steep screes in the central-southern
Apennines.
● Heteropterans.
Mountain screes host
many heteropterans, although most are
associated with xeric, rocky montane
pastures and cliffs. The most typical are
the already mentioned lace bugs of the
genus Copium, especially C. clavicorne,
a central-southern European species
found throughout Italy, except for
Sardinia. They are associated with cliff
and scree germanders (especially
Teucrium chamaedrys, T. scorodonia
and, in Sicily, T. flavum). The congener
Copium teucrii (Europe and the
Mediterranean basin) is more closely
associated with mountain screes and, in
the Alps, lives on Teucrium montanum.
Among flat bugs (aradids) is Aradus
frigidus, a Euro-Siberian insect found in a
very few sites in the Alps and southern
Apennines. Unlike its congeners - which
develop under the bark of felled trees - it
lives under small pebbles in high-altitude
rocky debris and on cliffs.
Another mountain species is the false
chinch bug Geocoris grylloides (lygeids),
which is generally found under
germanders. This Euro-Siberian insect
colonises the Alps and central
Apennines. Another false chinch bug is
the predaceous Geocoris lapponicus,
which lives in a few Alpine locations.
● Coleopterans. Ground beetles. They
are the most numerous in montane
environments, especially in grassland at
medium-high altitudes and in montane
habitats. However, the number of species
regularly living on screes and debris
deposits is much smaller. The large
Carabus genus includes mainly snaileaters of the subgenus Platycarabus, like
Carabus creutzeri (Eastern Alps) and
Carabus creutzeri
Anechura bipunctata
C. depressus (Alps) - found on screes
even under small, flat stones - and
members of the subgenus Orinocarabus,
like Carabus concolor (Central-Western
Alps), C. baudii and C. heteromorphus
(Western Alps). Typical beetles of screes
and humid debris deposits are a few
interesting high-altitude snail-eaters of
the genus Cychrus, very unusual,
isolated and of conservational interest,
like C. graius (endemic to the Graian
Alps), C. cylindricollis (Lombard, endemic
to the Bergamasque Alps), C. angulicollis
(Maritime Alps) and C. schmidti (Julian
Cychrus schmidti
Alps). One of the most typical beetles of
Alpine screes is Patrobus septentrionalis,
a rare orophilous species of the Eastern
Alps, with boreo-alpine distribution in
isolated, relict locations. Various
orophilous species of the genera Amara,
Pterostichus, Harpalus and Cymindis live
on mountain screes, although they
cannot be considered typical of these
habitats.
More humid Alpine screes host several
orophilous Bembidiinae of the genus
Ocydromus, especially those of the
subgenus Testediolum, a few Trechinae
of the genus Trechus (particularly those
of the T. strigipennis group), many
Oreonebria, Nebria germari (Dolomites),
N. orsinii (central Apennines), Licinus
Nebria germari
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italicus, and a few Platynus and Leistus
glacialis (central Apennines).
Rove beetles (Staphylinidae). In the Alps,
very interesting rove beetles are
associated with calcareous-dolomitic
screes, which are consolidated by
mountain avens (Dryas octopetala). These
environments contain turf with very deep
soil, the favourite habitat for many
apterous (wingless) and microphthalmic
(having small eyes) species of the genus
Leptusa, some of which live together in
the same areas, like the widely-distributed
L. piceata (along the Alps), as well as
many steno-endemics. In Firmetum, the
small flowers of mountain avens and the
white ones of mouse-ear (Cerastium) are
colonised by species of Eusphalerum of
restricted distribution: E. pulcherrimum in
the central Pre-Alps, E. albipile in the
Veneto Pre-Alps, E. annaerosae in the
Dolomites, and probably also
E. angusticolle in the Maritime Alps.
These species are associated with
mountain avens only in Firmetum,
because when the plant grows on more
Leptusa angustiarumberninae rosaorum
(Lombard Pre-Alps)
L. areraensis (Lombard Pre-Alps)
L. baldensis (Veneto Pre-Alps)
L. brachati (Veneto Pre-Alps)
L. cavallensis (Veneto Pre-Alps)
L. c. ceresoleana (Western Alps)
L. fauciumberninae
(Alps and Lombard Pre-Alps)
L. grignaensis (Lombard Pre-Alps)
L. knabli recticollis (Veneto Pre-Alps)
L. mandli (Lombard Pre-Alps)
L. manfredi (Dolomites)
Steno-endemic species of the genus Leptusa
evolved soils, these animals are totally
absent. Many other phytosaprophagous,
anthophagous and predaceous rove
beetles (e.g., Omaliinae subfamily) are
quite frequent between scree stones and
on the pulvini of glareous plants, although
they are orophilous generalists and are
more common in nearby montane
pastures and Seslerietum.
Sap beetles (Nitidulidae) and short-winged
flower beetles (Cateretidae). Many of
these beetles live on montane and submontane screes and rock deposits.
Among the most frequent are some
anthophagous sap beetles of the large
genus Meligethes, like the common M.
aeneus, associated with many plants of
the cabbage family (even cultivated ones),
found up to 2500 m. On high-altitude
screes, it is generally stenophagous on
plants of the same family (Biscutella spp.).
Other species are M. erysimicola and
M. fumatus (see drawing), already
described under cliff fauna, M. reyi and
M. solidus, which are associated with
orophilous plants of the rockrose family
L. montispasubii settei (Veneto Pre-Alps)
L. montiumcarnorum (Eastern Alps)
L. occulta (Veneto Pre-Alps)
L. piccata (Central-Eastern Alps)
L. portusnaoniensis (Veneto Pre-Alps)
L. pratensis (Eastern Alps)
L. rhaetoromanica (Rhaetian Alps)
L. rosai (Lombard Pre-Alps)
L. sudetica (Rhaetian Alps)
L. tirolensis tirolensis (Dolomites)
L. tridentina (Lagorai chain)
L. trumplinensis (Lombard Pre-Alps)
L. vallisvenyi (Western Alps)
(Helianthemum) growing in
xerothermic areas and sunny
screes in the Alps, Pre-Alps and
the Apennines as far as the Pollino
Massif, between 800 and 2500 m,
and M. oreophilus, an Italian
endemic found in the Alps and
Apennines exclusively on screes,
rock debris and Seslerietum on
medium-high altitude mountains
between 1200 and 2500 m,
associated with species of thyme. The
Eastern and Western Alps and an isolated
location in the Latium-Abruzzi Apennines
(Monte Elefante, in the Terminillo Massif)
host the rare M. devillei which, in southern
Europe, is exclusive to high altitudes and
associated with the rare orophilous
species of the genus Dracocephalum (mint
family), both in Seslerietum and on rock
debris. Another important species has
recently been discovered in several
southern European countries: this is
M. arankae, which in Italy is monophagous
on the rare crucifer Hesperis laciniata, a
typical mountain glareous plant. The
extremely rare Meligethes salvan may also
be associated with mountain screes. This
animal is a palaeo-endemic with a very
small distribution area and has only
recently been described. Its biology is still
unknown, and it is only identified by
specimens collected in the early 20th
century on Monte Argentera (Maritime
Alps). Among short-winged flower
beetles, the most typical inhabitant of
mountain screes is a member of the
genus Brachypterolus, the already
mentioned B. linariae, found in sunny,
rocky, xeric environments in Italian
montane and sub-montane belts and
associated with Kickxia spp. and Linaria
spp. (figworts).
Soft-winged flower beetles
(Dasytidae) and malachiid
beetles (Malachidae). These
two small families of minute
beetles include a large number of
anthophagous species living in
warm, sunny habitats.
However, very few are truly
adapted to life on mountain
screes, which are inhabited
by several species of softwinged flower beetles of the genus
Danacea, also found at much lower
altitudes. Very peculiar are the malachiid
beetles of the genus Malthodes belonging
to the group M. trifurcatus. Although they
normally live on conifers (Norway spruce,
larch), the populations colonising higher
altitudes above the treeline are
differentiated into “high-altitude” forms,
the taxonomy of which is little known.
They live on the ground, on screes and in
arid, rocky pastures. The females are
apterous and the males have poorly
developed posterior abdominal segments.
Malthodes trifurcatus atramentarius
(Western and Central Alps), M. penninus
icaricus (Central Alps), M. atratus atratus
(Maritime Alps) and M. atratus samniticus
(Gran Sasso) are perhaps “altitude
subspecies”, although little is known of
how gradients and substrate aridity affect
them. However, exposure to sunlight, the
duration and thickness of snow cover, and
other abiotic factors certainly play an
important role.
Blister beetles and oil beetles (Meloidae).
Although it is not typical, the oil beetle
Meloe erythrocnemus may occasionally
be found in xeric sub-montane screes at
medium altitudes. It has late winterspring phenology, and is generally
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associated with xeric low-altitude sites.
It lives in most of Italy and Sicily. Its
larvae typically feed on the leaf-cutting
bee Chalicodoma muraria. A Maghreb
species, Meloe aegyptius has similar
ecology and etology. It is very rare in
Italy, and can only be found in northern
Sicily in a few sites like Bosco di Ficuzza,
at the foot of the spectacular Rocca di
Busambra. A few orophilous mylabrines
of the genus Mylabris (especially those of
Mylabris variabilis
the subgenus Micrabris), such as
M. pusilla and M. flexuosa, which
typically live in steppe-like montane
grasslands, are sometimes found on the
scarce xerophilous grasses on Alpine
and Apennine screes, at even high
altitudes (2000-2500 m).
Darkling beetles (Tenebrionidae). Many of
these beetles live in rocky environments
at medium-high altitudes. The family
includes apterous phylogenetic groups
that can tolerate drought well. This makes
these evolutionary lines “pre-adapted” to
life on rocks, where strong winds favour
non-flying insects (flying ones would
easily be blown away). In addition, little
water - especially in summer - together
with the inability of soil to retain water
and lack of plant cover, all mean that
these insects survive in conditions of
extreme evapo-transpiration, temperature
variations and drought. Although many
insect species living near the snowline
have lost their wings to adapt to life at
high altitudes, these species of darkling
beetles were already apterous, which is
precisely why they were able to colonise
these inhospitable environments better
than others. There are therefore several
darkling beetle species which are
petrophilous (adapted to life under rocks)
in arid mountain habitats like screes and
rock deposits. Although none of them is
endemic to the Alps (which were recolonised by these animals only after the
last glaciation), the Apennine range has
two endemic species with subspecies
typical of mountain summits: Asida
pirazzolii and Colpotus strigosus
ganglbaueri. The former is characteristic
of the central Apennines and is divided
into two subspecies: the typical form is
frequently found under rocks in arid
environments, and A. pirazzolii
sardiniensis (which, despite its name,
does not live in Sardinia) colonises the
same habitats at lower altitudes.
C. strigosus strigosus and its subspecies
colonise the Apennines, especially along
the Tyrrhenian versant, from the TuscanEmilian range to Sicily. C. strigosus
ganglbaueri may be found under rocks in
a few arid sites in the central Apennines.
Another interesting darkling beetle is
Crypticus quisquilius, a species found all
over Europe, in the Caucasus, Siberia
and Mongolia. The ssp. aprutianus is
endemic to the central-eastern
Apennines, where it lives under rocks in
arid areas. Last come the sublapidicolous species Opatrum dahli,
endemic to Sardinia, and O. nivale
(Gennargentu mountains). The former is
xerophilous and colonises areas from
sea-level to 1000 m, and the latter lives
between 1200 and 1800 m.
Click beetles (Elateridae). Only a few
species of click beetles, most of which
are hygrophilous, live on mountain
screes, as they are better associated
with riparian alpine debris, near streams
and snowmelt rills. Hypnoidus
consobrinus lives in Scandinavia and
along the Alps, even at high altitudes.
The congeners H. rivularius and
H. riparius are more widely distributed,
and in Italy colonise the Alpine range and
the northern Apennines. These species
have root-eating larvae, and only require
minimal plant cover and relatively stable
soil surrounding plant pulvini in order to
colonise an area. Their presence on true
screes is therefore occasional. Another
species associated with rock debris is
Berninelsonius hyperboreus, of boreoalpine origin, which in Italy lives in the
Alps between Liguria (west) and Trentino
(east). Two species of Selatosomus, S.
amplicollis and S. aeneus, are sub-
Selatosomus sp.
lapidicolous mountain species that visit
steep slopes and rock debris.
S. amplicollis (southern Europe and
Turkey) is only found in the Ligurian and
Maritime Alps and along the Apennine
range as far as the Nebrodi and Madonie
mountains in Sicily. Selatosomus aeneus
(Siberia and Europe) lives in the Alps and
northern Apennines. The adults of a few
Ctenicera - with root-eating larvae that
require small quantities of soil to develop are frequently found on the few blades of
grass growing on screes (although they
are generally more numerous on xeric
grassland), especially
C. pectinicornis (Europe and Siberia),
which in Italy colonises the highest peaks
of the Alps and Apennines.
Scarabaeoid beetles (Scarabaeoidea).
Scarab beetles are occasional visitors to
mountain screes. Except for a few
unspecialised coprophagans that may
follow grazing animals along the margins
of these habitats, the only frequent
species are a few aphodiids of the
genera Agolius and Neagolius. Most of
them are typical of mountain zones
between 1800 and 2800 m (high
pastures, snow valleys, screes with
patches of snow). Agolius abdominalis
(Alps and Carpathian mountains) is
usually found in the excrement of
marmots, partridges, chamois, sheep
and sometimes cattle. Neagolius are
only found under rocks or flying.
Perhaps they are phytosaprophagous
rather than coprophagous. They include
Neagolius pollicatus (Eastern Alps and
Veneto Pre-Alps), N. amblyodon (Cottian
and Graian Alps), N. montanus (from the
Veneto Alps to the Balkan mountains),
N. limbolarius (from the Central Alps to
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mountains in northern Greece),
N. schlumbergeri (Pyrenees, Alps,
Apennines), N. liguricus (Ligurian and
Maritime Alps) and N. penninus (an Italian
endemic of the Pennine Alps, south of
Monte Rosa). The Neagolius are so
sexually dimorphic that males and
females were often described in the past
as distinct species. In some species, the
females are micropterous (smallwinged) or brachypterous (shortwinged) and depigmented. Some
species seem to be attracted to
light-coloured surfaces, and are
therefore found on calcareous
rocks and on screes.
Leaf beetles (Chrysomelidae).
Leaf beetles are not very
numerous on mountain screes,
and populations that typically
inhabit xeric pastures at medium-high
altitudes often mix with those living on
cliffs. The flea beetle Longitarsus
springeri (Alticinae) - a rare endemic at
medium-high altitudes in the Apennines is associated with Senecio rupestris
(daisy family), and the minuscule
congeners L. obliteratus and
L. obliteratoides, found throughout
Europe, are frequently found between
500 and 1500 m all over Italy in
association with xerophilous glareous
plants of the mint family, like winter
savory (Satureja montana) and
orophilous species of the genus Thymus.
Another flea beetle, Psylliodes instabilis,
of southern Europe and Italy, is
associated with plants of the cabbage
family, like those of the genera Alyssum,
Aurinia, Iberis and Eysimum. Dibolia
rugulosa of southern Europe and
northern Italy is often associated with
plants of the mint family, like annual
woundwort (Stachys annua). Among leaf
beetles is Oreina viridis, the larval
biology of which is little-known.
It is found at high altitudes in the Alps
(from the Maritime to the Julian Alps)
and in the central Apennines. Its
congener O. sibylla is endemic to the
central Apennines and is associated with
daisies of the genus Doronicum in
high-altitude pastures and nearby
screes.
Snout beetles (Curculionoidea).
Mountain screes host several snout
beetles. In addition to the alreadymentioned alpine elements of
the genus Dichotrachelus (see
drawing), which also live on rock
debris at medium-high altitudes,
there are four species of the genus
Oreorhynchaeus (O. baldensis,
O. focarilei, O. pacei, O. spectator), all
endemic to small areas of the Italian
versant of the Alps and Pre-Alps at high
altitudes and probably feeding on tiny
high-altitude plants of the pink family
that often grow between the rocks of
mountain screes. The same family
includes the rove beetle Trachystyphlus
alpinus (Staphylinidae), with many
subspecies living in the Alps and central
Apennines, and the rare weevil
Ceutorhynchus inaffectatus (mountain
areas in northern and central Italy) and
C. bifidus (mountains in central and
southern Italy), both associated with the
infrequent but typical glareous crucifer
Hesperis laciniata. In addition to several
other Ceutorhynchus, tychiines and
apionid weevils associated with other
high-altitude glareous plants (a few
Brachyodontus and Apion, several
Tychius, etc.), many other genera of
snout beetles live under sometimes very
small stones at the margins of mountain
screes, where the soil is more compact.
Among them are some high-altitude
orophilous Italian endemics of the
genera Otiorhynchus, subgenus
Nilepolemis, Neoplinthus and Leiosoma.
Otiorhynchus sp.
● Dipterans.
Hoverflies (Syrphidae).
Medium- or high-altitude screes and
rocky areas host members of this
important family, such as Rohdendorfia
alpina, a petrophilous species of the
Alps, typically found on rocky substrates
of moraines and screes, near streams,
and at the margins of glaciers (25002800 m). They skim surfaces swiftly and
rest on the highest jutting stones. As
soon as a cloud covers the sun, they
hide under pebbles. They feed on the
flowers of mouse-ear (Cerastium), Alpine
moon daisy (Leucanthemopsis alpina)
and stonecrop (Sedum). The females
hover over flat stones surrounded by
plant cushions, like those of mouse-ear,
and lay their eggs under the stones.
Cheilosia aristata (Alps) is a typical
orophilous fly of sunny siliceous rocky
environments facing south (2300-2400
m). It can be seen on the flowers of rock
catchfly (Silene rupestris). The males are
very territorial and rest on sunny flat
stones, from which they only fly off
occasionally to hover near the surface.
Ischyroptera bipilosa (Alps) is strictly
petrophilous and lives at over 2400 m on
compact, rocky screes and substrates
with little vegetation. The adults are
active between May and June, especially
when large patches of snow still remain.
The males, unlike most of the other
species, fly at 2-3 m above the ground.
Tachinid flies (Tachinidae). Among the
numerous members of this important
family of parasitoid dipterans is
Sarromyia nubigena (between the Central
Alps and the Pyrenées). It is a rare
species parasitic on the bagworm
Oreopsyche leschenaulti (Psychidae
moths). It flies in tiny swoops between
pebbles and creeping plants on highaltitude screes and rock debris, at over
2500 m, often on the flowers of
Rhododendron and creeping azalea
(Loiseleuria). Although the hosts of
Admontia cepelaki (similar distribution)
are unknown, this group is parasitic on
the larvae of crane flies (Tipulidae
dipterans). Just like the previous species,
it flies in a series of small swoops a few
centimetres above the ground on
high-altitude screes, often near perennial
snow, between 2700 and 3300 m.
Wagneria alpina (Alps and Pyrenées,
Scandinavia and European Russia) is
another xerophilous fly typical of southfacing rocky areas over 1200 m. Its hosts
are also unknown (perhaps caterpillars of
micro- and macrolepidopterans).
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110
● Hymenopterans.
Many hymenopterans
live on mountain screes, especially bees
(mining bees, leaf-cutting bees,
bumblebees). The vespoid wasp Polistes
biglumis bimaculatus, which lives and
nests in the Alps above 700-800 m,
behaves in a very peculiar way. Its nest,
made up of paper cells, is carefully
suspended under the rocks of screes so
that it does not touch the ground. In this
manner, the species can exploit the
particular microclimate, which is warmer
than the surrounding area, both because
the place is sheltered, and because the
stones are heated by the sun. This
enables the nest-builder to survive in the
mountains for longer periods than other
hymenopterans, except, perhaps, for
Bombus bumblebees.
There are many species of bumblebee in
arid grasslands and pastures at mediumhigh altitudes in the Alps and Apennines.
Although the adults do visit the flowers
and inflorescences of glareous plants,
none of the species is typical of screes
and rock debris, which are visited only to
collect pollen from flowers and exploit
the higher temperatures.
A few velvet ants (Mutillidae), especially
the common Mutilla europaea, parasitoid
on the underground nests of Bombus, is
Mutilla europaea
often found at the margins of the
warmest screes, where soil deposits
facilitate the construction of nests.
Many ants explore mountain screes,
although most of them cannot build
their nests on scree substrates for lack
of sufficient soil. The most frequent in
the Alps and Apennines are numerous
orophilous and xerophilous species of
the genera Lasius, Formica and
Tetramorium.
Lepidopterans. Both day- and nightflying lepidopterans have many typical,
although not exclusive species that
frequent mountain screes. Although
many of them are the same as those
generally found in the mountains, some
species actually prefer incoherent rocky
substrates, either because they are
strictly xerothermophilous, or because
their larvae are associated with glareous
plants. Screes host a kind of
xerophilous version of cliff biota, as
these areas feature great water drainage
and are therefore more arid. Satyrins
are, once again, one of the most
numerous butterfly groups on mountain
screes. Among them are Oeneis glacialis
(Alps) and several Erebia species, like E.
pluto and E. gorge (Alps and
Apennines). Generally speaking,
nocturnal lepidopterans on screes are
not very different from those already
described for montane and submontane cliffs. Among blues,
Scolitantides orion has several isolated
colonies on rocky, pebbly areas
especially in northern Italy, and its larvae
are associated with Sedum plants
(stonecrop family). Although not
exclusive to mountain screes, many
tiger moths like Chelis maculosa and
●
Arctia festiva find their ideal habitats in
these environments. Holoarctia cervini is
more closely associated with fissures
and glacial moraines, and lives in a very
few Alpine areas between 2500 and
3300 m. The well-known Arctia flavia, of
boreo-alpine distribution, lives in the
Alps at 3000 m. Among hawkmoths,
which are very large lepidopterans, the
most closely associated with glareous
plants is Hyles vespertilio, the
caterpillars of which feed on Epilobium.
In the large family of owlet moths, typical
elements are the rare, heliophilous
Sympistis, found at high altitudes on the
Alps, and Euxoa culminicola, which lives
in the same areas.
Erebia pluto
Screes also host very peculiar species,
the biology and distribution of which are
still little known, like the measuring worm
moths of the genus Elophos with two
Alpine species (E. caelibaria, E. zelleraria), whose females have very short
wings. However, the most characteristic
scree species are Sciadia tenebraria and
those of the genus Glacies, with many
diurnal Alpine species living at 4000 m.
G. alticolaria and G. coramica live in a
few Alpine sites, and G. canaliculata is
widespread in the Alps. Other typical
species are some Alpine species of bagworms of the genus Oreopsyche and
several gelechiid moths, especially of the
genus Caryocolum.
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112
Vertebrates: taxonomy
■ Amphibians
Few amphibians can survive in
mountain slope environments. Two
species, the large Alpine salamander
(Salamandra lanzai) and the Alpine
Lucio Bonato
individuals which have already
metamorphosed. Large Alpine
salamanders produce 5-6 juveniles, and
Alpine salamanders not more than two.
These species can therefore colonise
even the steepest flanks, and also karst
mountains, where water seldom
collects. Lack of competition from other
urodeles enables them to colonise these
areas in large numbers. The large Alpine
salamander is confined to a small area
in the Cottian Alps, and the Alpine
salamander lives in most of the Central
and Eastern Alps.
■ Reptiles
Alpine salamander (Salamandra atra)
Ibex (Capra ibex)
salamander (Salamandra atra), are the
only ones whose physiological
adaptations enable them to colonise
these areas.
These animals can withstand lengthy
periods of cold winter temperatures at
high altitudes, hibernating for several
months in the ground under the snow.
Unlike other amphibians, they do not
require surface water to reproduce and
develop.
Their entire larval stage is intrauterine
and the embryos receive nutrients not
only from their mothers, but also from
other embryos, which are doomed to
die. After a gestation that lasts 2-3
years, the females give birth to small
Only a few reptiles visit rocky cliffs and
screes, being restricted by their
temperature requirements. The harsh
meteorological conditions prevent them
from reaching the sufficient body
temperature that enables them to move,
feed and reproduce.
However, the viviparous lizard (Zootoca
vivipara), which is widespread in the
Alps, can tolerate the cold, humid
climate of high altitudes.
Although these lizards are generally
terrestrial and prefer grassy soil, they
can move on debris slopes and climb
rocky surfaces to warm themselves in
the sun and hunt the tiny arthropods on
which they feed.
Unlike oviparous populations living at
lower altitudes, most mountain lizards
have a particular way of reproducing:
the eggs are stored inside the mothers’
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114
Horvath’s rock lizard (Iberolacerta horvathi)
bodies, and emerge as small, active
lizards.
Unlike the viviparous lizard, Horvath’s
rock lizard (Iberolacerta horvathi) is less
tolerant of low temperatures but is even
more typical of slope habitats, and can
climb steep surfaces skilfully. This
species lives at intermediate heights,
almost exclusively on rocky slopes and
rock debris. It frequents small, still littleknown areas, mainly from the eastern
Carnic Alps towards the Dinaric Alps.
In the Carnic range in particular, it
regularly colonises screes and slopes up
to 2000 m, where it hunts spiders and
insects.
Lizards, small song birds and rodents
visiting these rocky habitats may be
stalked by the adder (Vipera berus).
Just like other vipers, the adder is a
strictly terrestrial snake that hunts by
exploiting its sensitivity to smell and
temperature. It approaches its prey
cautiously, pounces on it suddenly and
bites it. It then waits before eating it,
until its neurotoxic saliva has had its
effect. Unlike other vipers, adders can
better tolerate low temperatures and
considerable temperature variations
between day and night.
They have therefore colonised most of
the temperate and sub-arctic regions of
Eurasia, and in the south are restricted to
the mountains. In Italy, they live only in
the Alps, generally above 1000 m.
There, they are often to be seen on
sunny rock deposits, between shrubs
and Alpine pastures, which enable them
to regulate their body temperature,
especially in spring.
Their ovoviviparous reproduction is
particularly adapted for survival in the
montane climate: the embryos do not
develop inside laid eggs, but inside the
mother’s body, a warmer and stabler
environment. Every two years, the
females give birth to a dozen small, active
individuals.
Like the adder and even more closely
associated with rocky
substrates is the hose-horned viper
(Vipera ammodytes) which, despite being
■ Birds
Rocky slopes and jutting stones in the
Italian mountains are visited exclusively
by some bird species. Niches and deep
fissures in rocky slopes are perfect
shelters for the Alpine chough
(Pyrrhocorax graculus) and the Cornish
chough (Pyrrhocorax pyrrhocorax). In
winter, these areas shelter flocks of
dozens, even hundreds of birds and, in
summer, are used as breeding grounds
for single pairs or colonies. The pitch
black feathers of these birds contrast
with the red skin of their thin legs and
with the colour of their bills, which are
yellow and short in the Alpine chough,
bright red and slightly curved in the
Cornish chough. Their black feathers are
even more evident against the white
background of stones and snow, which
Hose-horned viper (Vipera ammodytes)
a mountain species, does not reach high
altitudes. It lives in the Balkan area as far
as the eastern Alps and the arid, rocky
areas of Vallagarina in Trentino.
Alpine chough (Pyrrhocorax graculus)
cover these areas for lengthy periods.
Choughs look for food on only partially
rocky grassland, on high-altitude
pastures and terraces with minimal soil
cover. Walking on the ground, they turn
stones over with their bills and search the
soil for insects and arthropods. Many
choughs often fly in noisy flocks to help
keep in contact. They emit hisses and
shrieks that echo on the rocky slopes
and cliffs. Climbers and trekkers are
familiar with these birds, because they
often approach summer pastures and
mountain huts seeking for man’s leftovers. In Italy, both species are restricted
to the highest Alpine and Apennine
summits, seldom nesting below 1500 m,
and occasionally reaching the snowline
looking for food. The Alpine chough is
widespread in the Alps, whereas the
Cornish chough lives only in the Western
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116
Alps, having become extinct in the other
areas of the chain. The two species may
cohabit in the Apennines, but only the
Cornish chough lives in southern Italy,
Sicily and Sardinia.
Crevices in mountain slopes are used as
nesting grounds by the raven (Corvus
corax), which lives in the Alps and on all
Italian mountains, including those on the
major islands. It is actually an
opportunistic generalist for both food
and breeding grounds. In its wide
distribution area, from America to
Eurasia, it visits various environments,
not only rocky mountains, and may even
nest in trees. In northern Italy, due to
man’s disturbance, it is restricted to the
most inaccessible slopes, although in the
south it is frequently found at low
altitudes and on sea cliffs. It has shiny
black feathers, a wide wingspan, a
strong bill and a wedged tail. Its body is
powerful, its flight is accompanied by
Raven (Corvus corax)
hoarse croaks and, at close quarters, by
the swishing sound of its wings. It lays its
eggs in February, when its surroundings
are still covered with snow. The nest is
large and complex, being a platform of
strong twisted twigs over 1 metre in
diameter, covered with fine vegetal
debris and lined with soil, moss and
other soft materials. Ravens feed on the
ground, even far from slopes, generally
hunting small animals, but also plant
material and organic waste.
The crag martin (Ptyonoprogne rupestris)
may nest on protected, not windy slopes.
Although the species is associated with
temperate or even warm, dry climatic
conditions, it has colonised rocky
montane environments of the pre-Alpine
and Apennine areas. In recent years, it
has also colonised cities in northern Italy,
where buildings, bridges and masonry
provide conditions similar to those of its
original breeding grounds. Its nest has a
117
Crag martin (Ptyonoprogne rupestris)
Kestrel (Falco tinnunculus)
semicircular jug-like shape, made of clay
material that is collected, carried and
moulded with its small bill. Nests are
constructed under ledges and jutting
stones on bare, almost vertical cliffs.
Crag martins are small, with tapering
bodies and sharply tipped wings. Skilful
fliers, they perform stunts like sudden
landings, accelerations and acrobatic
veers that enable them to flash past
rocky cliffs and faces, taking insects by
surprise and catching them in mid-air.
Ledges and crevices are inaccessible,
undisturbed places where species of
falcon may also nest. The most common
on Italian mountains are the kestrel
(Falco tinnunculus) and the peregrine
(Falco peregrinus). They do not build
nests, but lay their eggs directly on the
rocky surface or in the little sandy soil
that accumulates in protected areas.
They may occupy abandoned nests
previously used by ravens or other birds.
They are very good fliers and hunt in
open areas, exploiting their sharp sight.
The kestrel is smaller, slimmer, and its
wings are more curved than the
peregrine’s. It glides at considerable
heights, carefully searching the ground
below, often hanging poised in the air,
before plunging swiftly on large insects
and small rodents moving on the
ground.
The peregrine is a more robust bird, with
stronger claws and bill and a wider
wingspan. It hunts birds while airborne,
chasing them or catching them by
surprise by suddenly accelerating and
plunging down with closed wings along
inclined trajectories.
Kestrels nest from sea-level to above
2000 m, and regularly visit meadows
surrounding mountain slopes. Peregrines
are less frequent, but typical of jutting
118
The wallcreeper
The unusual wallcreeper (Tichodroma
muraria) is certainly the vertebrate
species best adapted to life on highaltitude slopes. Its strong feet, with
long digits and claws, are very
effective for climbing vertical, even
jutting surfaces, which the wallcreeper
ascends jerkily but swiftly.
With its stumpy tail and short wings,
it performs quick, butterfly-like
flights and makes safe landings
between cliffs.
It forages on rocky walls, feeding on
the insects and spiders that frequently
visit these areas. The wallcreeper
climbs the walls inspecting crevices,
niches and fissures, as well as scanty
soil deposits and blades of cliff
grasses clinging to the surface. Its
long, thin, slightly downward-curving
bill can enter even narrow interstices,
where its thin, forked tongue extracts
the small invertebrates hiding within.
Its acrobatic dexterity enables the bird
to catch even disturbed insects trying
to escape.
When clinging to jutting stones, its
wings folded, the wallcreeper appears
as a small, ash-coloured lump
between the cracks and rocky walls,
inconspicuous to raptors hovering
over the same cliffs in search of prey.
However, the bare stone is a
background against which this bird
can communicate with flashes of
colour. While performing its jerky hops,
the wallcreeper partially opens its
wings, flicking its rectrices for a split
second.
The visual effect is stunning: a flash of
crimson immediately followed by a
black-and-white pattern. The colours
are even more evident when the bird
flies, because the red patches and
Lucio Bonato
black-and-white mosaic on its back
are fully revealed.
In spring, during the breeding season,
the males exhibit their black throats to
the females by lifting their heads and
pointing their bills upwards. Mating
takes place on the same cliffs. Each
pair builds its nest in a protected
crevice within its territory, generally a
whole rocky wall or a smaller series of
jutting rocks. The male performs
typical circling flights to mark the area
it has chosen to build the nest,
emitting nasal whistles. The female
prepares the nest by twisting mossy
tufts and blades of grass together,
leaving two apertures, one entrance
and one exit. After the mother has sat
on her eggs for about 20 days and fed
them for another month, four or five
juveniles emerge from their nests,
stretching their necks to peep out into
the void.
stones in the pre-Alpine and
pre-Apennine foothills, to 1500 m.
Neither species is typical of montane
environments, as they generally colonise
rocky areas in hills, coastal cliffs, and
buildings and masonry in cities.
The golden eagle (Aquila chrysaetos) is a
mountain species of great ecological
importance on high-altitude meadows
and slopes. With a wingspan of more
than 2 metres, a strong head and neck
and a large curved bill, the golden eagle
glides over undulating, high-altitude
rocky pastures to stalk hares, marmots
and birds. The narrow valleys and rocky
pinnacles of these areas provide it with
safe nesting grounds.
The nest is a large, tall platform made up
of intertwined twigs, where the female
lays 2 eggs each year and weans only
one juvenile.
The same inaccessible cliffs can be
nesting grounds for the bearded vulture
(Gypaetus barbatus), a large bird about
1 metre long, with a wingspan of about
3 metres. As an adult, it has a very
particular aspect, because a strip of
black bristles rises from its tawny head
and stretches to under its “chin” to form
two beard-like tufts. It glides over
pastures and rocky slopes hunting for
carrion of sheep and goats, only
occasionally feeding on live animals. It
breaks bones by dropping them on
rocky wedges, and then extracts the
marrow with its tongue.
Distributed on the mountain ranges of
the Mediterranean basin, in the past the
bearded vulture was widespread on the
Italian mountains. However, direct
hunting by man and reduction of its food
sources, due to the declining numbers of
wild cattle on the mountains, have
gradually brought it to extinction.
Until the 16th century, it was still found
in the Apennines, in the 1850s it became
extinct in Sicily, in the Alps it gradually
disappeared from east to west in the
early 20th century, and the last pairs
were still living in Sardinia in the early
1960s. Recent re-introduction strategies
carried out in the Alps have had positive
results, but recovery times are very slow,
because these birds reach sexual
maturity after several years and their
reproduction rate is very low, because
the pairs only lay one egg each year.
The species which is best adapted to
living, breeding and feeding on rocky
faces is certainly the wallcreeper
(Tichodroma muraria). A rock-creeping
passerine, this bird skilfully climbs
vertical surfaces looking for food like tiny
arthropods. Cliffs are also its breeding
grounds and shelter. In summer, it
reaches the highest rocky mountains in
the Alps and northern-central
Apennines, preferably on limestone and
dolomite, to 3000 m, nesting in
protected areas far from predators. In
winter, when meteorological conditions
in the mountains become harsh and prey
is no longer available, the wallcreeper
flies to lower altitudes by means of a
series of short, mainly altitudinal
migrations. It winters in pre-Alpine hills,
along the margins of the Apennines and
also in the plains, as far as the sea. Its
environments are always similar, as it
prefers sunny, south-facing areas richer
in arthropods, like natural rocky cliffs,
cave or building walls. In summer, other
insectivorous birds visit rocky debris and
mountain screes, where they find food
between the rocks and patches of grass.
Among these are the black redstart
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120
Black-eared wheatear (Oenanthe oenanthe)
(Phoenicurus ochruros), the black-eared
wheatear (Oenanthe oenanthe) and the
Alpine accentor (Prunella collaris), often
found together in the same rocky areas
where they feed and breed. Although
their ecology, size and body structure are
very similar, their song and plumage are
different. The black redstart has uniform
dark head and thorax, and rust-brown
rectrices, the black-eared wheatear has
contrasting black-and-white patterns on
its head and tail, and the Alpine accentor
has uniformly grey plumage with brickred streaks along its sides. In the
summer, these species are commonly
found along the Alps. In the Apennines,
the Alpine accentor visits only the
central-northern area; the black redstart
and the black-eared wheatear in
particular colonise the Italian peninsula
and the major islands. Very agile on their
thin legs, they hop and walk on the
ground and stones, looking for prey,
which their thin bills peck up with sudden
yerky movements.
Black redstarts and black-eared
wheatears sometimes perch on rocky
spurs, peering into the distance and then
plunging on prey. In these open, irregular
environments, their flight is short and
low, often interrupted to check the
surroundings. If they sense danger, they
quickly hide between rocks and crevices.
In these conditions, their colours play an
important role in communication: when
they fly low, the rust-brown tail of the
black redstart and the black-and-white
pattern of the black-eared wheatear are
evident. In the mating season, the males
mark their territory by emitting short,
elaborate sounds - particularly sharp in
the black redstart - as they sit on the
tops of rocks and spurs.
Their nests, made up of moss and blades
of grass and lined with hair and feathers,
lie directly on the ground, in protected
crevices or niches in the cliffs. In winter,
when food is limited, they fly to lower
altitudes.
The Alpine accentor stays in Alpine and
Apennine areas, but on south-facing
flanks of mountains and hills; the black
redstart flies to the plains and coasts,
colonising even cultivated fields and
cities; black-eared wheatear migrates to
sub-Saharan regions.
The snow finch (Montifringilla nivalis) lives
on the summits of cliffs all year round.
This passerine species inhabits all the
mountain ranges from southern Europe
to the Himalayas.
In Italy, it is found in the Alps and central
Apennines. Female and male adults
share the same black-and-white
plumage, which is maintained throughout
the year. Snow finches peck up seeds
and arthropods on the ground on highaltitude grasslands and in winter, when
the extensive snow cover limits food
availability, they become opportunistic
commensals, benefiting by left-overs
from trekkers and skiers, and even
approach mountain huts and ski
facilities.
The same rocky environments
surrounding the Alpine summits are
visited by the rock ptarmigan (Lagopus
mutus), certainly the best adapted of the
grouses (tetraonids) to life in the Alpine
environment.
Throughout the year, this strictly
terrestrial bird lives at the margins of
screes and on rock debris carpeted by
shrubs and grass cushions in the Alps,
usually at over 2000 m and up to 3000
m. In winter, the thick snow is a uniform
background on which the perfectly white
Rock ptarmigan (Lagopus mutus)
rock ptarmigan walks, almost invisible.
The only exceptions to its candid attire
are the outer tail rectrices which,
however, are only visible when the bird
flies.
The males also have a thin black line
crossing their eyes. They move in flocks,
walking on the snow, and avoid sinking
into it thanks to the stiff feathers that
cover their feet and tarsi. They burrow
their nests, rest and shelter in the frozen
surface. In the short Alpine summer,
when the snow melts to reveal rocks and
plant patches, the plumage of rock
ptarmigans becomes dark, with
brownish-black spots, which enables
them to blend in perfectly against the
new, darker background. In this period,
the females lay their eggs, which are also
camouflaged and spotted, in hollows in
the ground.
121
■ Mammals
122
The chamois (Rupicapra rupicapra) and
the Pyrenean chamois (Rupicapra
pyrenaica ornata) are among the largest
mammals regularly visiting debris slopes
at the base of rocky walls.
These ungulates can dexterously climb
unstable, steep rocky deposits in order
to reach undisturbed pastures, to avoid
man and occasional predators.
They can do this thanks to the special
characteristics of their legs and feet,
which have very strong tendons and
robust joints that tolerate remarkable
stress and torsion.
Their sharp, horny hoofs, with broad,
flexible pads, provide a firm grip on
unstable, slippery rocks and frozen
debris. Their internal organs (heart and
Croda Cimoliana (Carnic Pre-Alps, Friuli Venezia Giulia)
Pyrenean chamois (Rupicapra pyrenaica ornata)
lungs) are particularly well developed to
enable them to cope with fast running.
Chamois live in forests and cliff habitats
along the mountain versants above the
treeline, up to the small patches of
grassland on the highest rocky summits.
In summer they move to great
heights, and in winter wander down to
lower altitudes.
The chamois is currently found in the
Alpine range, from the Maritime to the
Julian Alps, and the Pyrenean chamois
lives in the Abruzzi mountains,
especially in the Camosciara and Meta
groups, on the Majella Massif and
Gran Sasso.
The Italian populations of the two
species are actually increasing, thanks
to recent conservation strategies of reintroduction in territories where their
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predators are almost totally absent.
Until a decade ago, their Italian
populations, especially those of the
Pyrenean chamois, were threatened by
excessive hunting.
In the past, the chamois, which probably
colonised Italy during a recent glaciation,
could be found from the Sibillini
mountains to Monte Pollino, and the
Pyrenean chamois, which migrated from
eastern Europe during the last Ice Age,
colonised the Alps and northern
Apennines.
The numbers of ibex (Capra ibex) have
fallen even more dramatically in recent
centuries. As soon as the climate
became milder after the Würm glaciation,
an event which had caused ibexes to
disappear in the rest of Europe, these
ungulates moved to the Alps. However,
Mountain hare (Lepus timidus) in summer
centuries of excessive hunting made
them almost extinct, and in the second
half of the 19th century, only a hundred
individuals survived in the Gran Paradiso
Massif (Graian Alps).
The creation of a royal game preserve
and, later, of a national park, enabled
ibexes to survive into the 20th century,
and repeated re-introduction has
produced several Alpine populations.
Ibexes live in alpine environments above
the tree- and shrublines.
They can climb slippery, steep, rocky
surfaces with great agility.
In summer, they reach the patches of
grassland growing above 3000 m and, in
winter, wander down to warmer altitudes
between 1500 and 2000 m.
Among other herbivores capable of
finding seasonal food in alpine belts
below the snow level, there is the
mountain hare (Lepus timidus). Although
this animal normally lives in the Alps well
above 2000 m, it can colonise areas with
scanty shrubs and forests at lower
altitudes. Its ever-growing, chiselshaped incisors are more curved than
those of other species of hare, and
extremely effective for gnawing on bark
and severing plant roots and the fibrous
stems of grasses.
Like other hares, its long hindlegs are
specially adapted for sudden
acceleration, enabling it to bound away
from predators in long leaps. Its ears are
proportionally shorter than those of other
hares, because they have to withstand
freezing temperatures and cold
mountain winds. The coat of mountain
hares undergoes seasonal colour
Stoat (Mustela erminea) in summer
changes, and becomes totally white in
winter to blend in with the regular,
lengthy snow cover.
The stoat (Mustela erminea) resorts to
similar camouflage. This carnivorous
mammal is adapted to moving and
hunting between stones and rocks. Its
pointed muzzle, elongated neck, slender,
flexible body and short, agile legs enable
it to climb rocky surfaces with great
dexterity, stalking birds and small
mammals which it catches with
lightning-swift leaps.
The summer coats of Alpine stoats are
reddish-brown and, in winter, turn
completely white - like those of Arctic
stoats - with a tuft of black hair at the tip
of the tail.
The snow vole (Chionomys nivalis) is the
most typical prey of stoats. This small,
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Alpine marmot (Marmota marmota)
Snow vole (Chionomys nivalis)
Alpine shrew (Sorex alpinus)
grey-coated rodent colonises stable
screes and rock deposits in the Alps and
Apennines. Although it generally seeks
refuge underground, in the interstices
between rocks, it emerges on the
surface even during the day, to feed on
plants and shrubs, and bask in the sun.
Among the numerous Eurasian voles, it
is certainly the best adapted to microthermic, high-altitude rocky
environments, and may reach the Alpine
peaks - up to 4000 m on the Mont
Blanc. During the last glaciation, snow
voles were common in Italy, even at low
altitudes.
Today, they live on the Alpine summits,
and a few relict populations may also be
found in the Apennines.
The larger Alpine marmot (Marmota
marmota) is another rodent which
spends part of its life underground.
The more or less stable ridges circling
Alpine glaciers provide optimal
conditions for the settlement of Alpine
marmot colonies. Although they forage
on grassy patches, rocks and stones are
ideal camouflaged entrances to their
underground burrows. They are also
surfaces on which marmots prefer to
bask in the sun and stand on guard. If
they sight a potential predator or an
approaching danger, marmots emit
sharp, penetrating cries of alarm and,
leaping between the rocks, hide in their
burrows. The narrowest interstices
inside debris deposits offer shelter and
passageways to the Alpine shrew
(Sorex alpinus).
Although this is the highest-reaching
species among insectivores, it is not
restricted to open habitats above the
treeline, and may be found at lower
altitudes, on the rocky surfaces of humid
woodland in small valleys.
Alpine shrews are active at night, when
they scurry in search of arthropods and
other terrestrial invertebrates which they
identify thanks to the great sensitivity of
their noses.
In north-eastern Italy, the Alpine shrew
can wander to the high plains, at 160 m
(Cornino, Friuli Venezia Giulia).
Conservation and management
PAOLO AUDISIO · LUCIO BONATO · MARCELLO TOMASELLI
■ Endangering factors
From several standpoints, mountain
slopes and screes are the best
preserved and least threatened
habitats in Italy.
Their uneven, steep surfaces,
geomorphological instability and
harsh climatic conditions have always
made them dangerous and almost
inaccessible. In the past, therefore,
humans only came to these cliff areas
occasionally, and then only for
hunting, quarrying or transportation
Protected mountain cottage at the base of a
purposes. Roads were unlikely to be
debris slope (Val Grosina, Lombardy)
built here, and gently sloping valleys
were obviously preferred. Recently, however, roads and railways have
increasingly invaded these habitats, with sometimes whole series of tunnels
through rock or long bridges over valleys.
These mountains have hosted human settlements for only temporary periods.
The absence of true soil and the high permeability of the debris substrates
hinders agriculture. Tree growing has also been very difficult, because the soil
cannot support any trees other than larch and patches of forests with
mountain pine and alder. Even sheep-rearing, which was a quite frequent
mountain activity in the past, could only rely on poor, steep, rocky
pastureland. However, local outcrops and debris slopes have sometimes
favoured the quarrying of various types of rocks and minerals for industrial
purposes.
Despite their overall good state of preservation, mountain slopes and screes
have recently been threatened by modern human activities which do not only
damage the beauty of the landscape, but also interfere with natural physical
dynamics and local biocoenoses.
Limestone scree at the foot of Monte Coglians (Carnic Alps, Friuli Venezia Giulia)
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Rock climbing, mountaineering, and other sports. The ever-growing
number of visitors to these environments poses a severe threat. Rock
climbing, mountaineering, hiking and other related sports are becoming more
and more popular - even true mass phenomena in Italy. The steepest slopes
are favourite runways for hang-gliders, and jutting rocks are used as platforms
for bungee-jumping. These types of sports are not performed only at high
altitudes, but also on hills lower down. As they are not generally regulated,
they jeopardise local animal life and may also damage the slope surfaces
irredeemably.
The essentials of these mountain sports are learnt on cliffs which are specially
equipped with pitons (metal spikes driven into rock cracks), ropes and
carabiners (metal loops that are snapped into pitons and through which ropes
are passed) in order to create permanent trails. This means that all cliff
vegetation has to be uprooted to provide footholds on jutting rocks which are
constantly trodden on by crampons or used as camp sites, and where rubbish
impossible to carry down is dumped.
These are only some of the factors producing a severe impact on the mountain
landscape. Pioneer plants and associated phytophagous insects are damaged,
and cliff raptors and other sensitive birds breeding in spring and summer are
A climber’s pathway with iron steps set in rock
jeopardised. Evidence shows that even
involuntary disturbance near nests
reduces successful reproduction, and
sometimes rare birds protected by
European laws, like the golden eagle
and peregrine, may even abandon their
nests. In the past, the eggs of these
species were taken by collectors and
chicks were stolen by falconers.
Unfortunately, lack of awareness of
the natural importance of these
environments enables these sports and
activities to be carried out freely
everywhere and at any time of year.
Protected areas where they are banned
have sometimes been defined, and
the Club Alpino Italiano and the
Rock climbers
Associazione Italiana Preparatori Itinerari
d’Arrampicata have produced codes of self-regulation, but these norms and
limitations are often not respected. Tourist activities on gentler slopes can also
jeopardise cliffs nearby. These areas are often crossed by cableways reaching
high-altitude mountain huts, and funiculars and cable-cars exploit the beauty of
the mountain landscape to carry tourists, rock climbers and hikers to even
higher altitudes. Many ski-lifts and cable-cars, especially in the Apennines, can
disturb the environment and provide endless sources of rubbish and various
objects which are either dumped or lost. Maintenance of tourist facilities, which
is carried out with tractors, severely damages scree vegetation, produces
erosion of nearby alpine meadows, and oils and other pollutants can leak out
into the porous substrates.
Roads and other infrastructures. When buildings and roads develop below
rocky versants and debris slopes, they must be protected against material
falling from above - the result of landslides and debris flows in rainy seasons
and avalanches in spring and winter. Metal meshes, barriers and periodic
removal of unstable material, either manually or with explosives, have visible
effects. Injections and covers of concrete irreparably compromise the original
ecological functions of these environments - even more so, when they are
inhabited by particularly important cliff plants and their associated
phytophagous insects. The construction of hydro-electric power plants often
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involves variations to the local
microclimate due to huge amounts of
water and associated infrastructures,
like water pipes under pressure,
drains and tunnel passage-ways, have
an even greater impact on rocky
slopes. Moreover, these structures are
used for mountain sports when they
are no longer exploited by hydroelectric companies, thus becoming a
kind of shared heritage of industrial
archaeology.
Pylons for power and telephone lines
and relay stations have been erected,
neglecting the natural and aesthetic
values of steep slopes. Their impact is
therefore severe, both for the
Relay station on a mountain summit (Sardinia)
alterations they create to the local
environment and for the consequences associated with maintenance.
Multiple power cables can electrocute large birds like raptors and ravens,
especially near the rocky faces and ridges where these birds nest. Little is
known about the effects on biocoenoses of so-called electromagnetic
pollution. As if all this were not enough, military installations have covered
extensive substrates with cement and roads have been built on barely
accessible slopes.
Caves and quarries. Local, destructive impact is produced by quarrying
mountains to extract materials for building roads and railways, or to obtain
particular types of rock for industrial and ornamental use. The effect is
severe not only when quarrying occurs in the open, but also when it is
carried out underground, as it gives rise to ever-accumulating debris,
acoustic disturbance, vibrations and transport of particulate matter
(sometimes toxic) by winds. Although today quarrying in the mountains is
less frequent than in the past and at lower altitudes, its consequences are
still felt high up. For example, in the Apuan Alps, centuries of marble
quarrying have removed and moulded entire mountains with devastating
effects. This has given rise to new rocky slopes and screes, which have
recently become popular destinations with tourists, who are attracted by the
residual environmental value of these areas.
Dumps. Small mountain towns, especially in central-southern Italy, Sicily and
Sardinia, have often used gently sloping mountain screes and dry cirques as
preferential dumping sites, thus destroying the original state of these areas.
They are even more deeply affected by illegal dumps, which here find ideal
locations due to the fact they are unmonitored, little frequented, barely visible
and almost inaccessible.
Grazing. Consolidated screes, moraines and rocky flanks covered with
vegetation are often overgrazed by sheep and goats, which can easily climb
these steep slopes. Excessive grazing has sometimes reduced the plant
cover, and has always modified its structure and floral composition, giving rise
to debris flows. Accumulation of excrement at the base of boulders, on natural
terraces and along gorges has enriched the local substrate with nutrients,
which in turn destroy sensitive chasmophytes and glareous plants, and favour
the development of nitrophytes and more tolerant plants. This also affects the
associated communities of epigean phytophagous arthropods.
■ Values and the current situation
Preservation of the natural conditions of cliff environments is recognised by
“Marble” quarry in the Carnic Alps (Friuli Venezia Giulia)
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the European Community, which has
included these areas among those of
European interest.
Among those listed in Annex I of the
Habitat Directive (92/43/CEE), the
conservation of which requires the
identification of special areas, are
mountain screes, with both siliceous
(Androsacetalia
alpinae
and
Galeopsietalia ladani) and calcareous
communities (Thlaspietea rotundifolii),
and siliceous and calcareous rocky
slopes with chasmophytic vegetation.
Central European calcareous screes
and limestone pavements deserve
particular attention.
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Slopes and debries in the Monte Pramaggiore
massif (Carnic Pre-Alps, Friuli Venezia Giulia)
Flora at risk. Although flora on screes
and mountain slopes is not greatly
endangered, some chasmophytes and glareous plants are considered
vulnerable and are included in the “Libro Rosso delle Piante d’Italia” (Red List
of Italian Plants). Among these are Berardia subacaulis, Rhamnus
glaucophyllus and Saxifraga tombeanensis.
Other species, which cannot yet be considered endangered but might
become at risk in the future, are classified as rare, like Achillea lucana, Adonis
distorta, Androsace mathildae, Aquilegia champagnatii, Armeria gussonei,
Ballota frutescens, Campanula morettiana, C. raineri, Daphne petraea,
Helichrysum montelinasanum, Linaria tonzingii, Moehringia dielsiana, M.
lebrunii, M. markgrafii, M. papulosa, M. sedifolia, Moltkia suffruticosa,
Papaver degenii, Physoplexis comosa, Potentilla saxifraga, Primula allionii,
Rhizobotrya alpina, Salix crataegifolia, Saxifraga arachnoidea, S. cochlearis,
S. florulenta, Sedum aetnense, Silene elisabethae, Viola comollia and V.
magellensis.
Devil’s claw (Physoplexis comosa)
Animal communities. Similarly, conservation of most animal species living in
these environments is not at risk. However, the numbers of some cliff raptor
species, whose breeding grounds are closely associated with these habitats,
have recently fallen due to illegal hunting, disturbance, and alteration of their
environment. For example, the bearded vulture became extinct in Italy in the
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20th century, and has only recently started reproducing again in a few areas,
thanks to initiatives of re-introduction and conservation. The peregrine and
golden eagle are less endangered. Among the birds living on mountain screes
and slopes, all raptors have been listed as threatened in the Bird Directive
(79/409/CEE), together with the Cornish chough and the rock ptarmigan,
occasional visitors to these habitats.
The numbers of chamois and particularly those of ibex have also fallen,
particularly as a consequence of excessive hunting in the 20th century. They
have recently been re-introduced and their communities are now increasing
slowly. The Habitat Directive has recognised the Pyrenean chamois as a
European priority species.
As regards invertebrates, the Ministry for the Environment and Territory
Protection is promoting the compilation of an objective, updated list of
endangered species. It will probably contain species that are typical or
exclusive to mountain slopes and screes, especially phytophagous insects
and molluscs, many of which are endemic to restricted areas.
Species of European Community interest which are already listed in the
Habitat Directive are Buprestis splendens and Parnassius apollo, typical of
mountain slopes, Rosalia alpina and Zerynthia polyxena, very occasional
visitors, and Erebia calcaria and E. christi, occasionally found on mountain
screes.
Many endemics, as well as several species of natural interest, colonise the
Alpine chain, especially the warmest calcareous areas. Other important
species are found, especially on the Latium-Abruzzi mountains in the
§Apennines, in Calabria and Lucania in the south, on the highest Sicilian
mountains (Nebrodi and Madonie chains) and in Sardinia (Gennargentu).
Erebia calcaria
Bearded vulture (Gypaetus barbatus)
■ Future prospects
One of the greatest obstacles to the conservation of natural life on mountain
slopes and screes is difficulty in arousing the interest of institutions, tourists
and rock climbers to the ecological, natural importance of these
environments. The severe impact of infrastructures, no matter how isolated,
is never evaluated in advance.
The misconception according to which these habitats are inhospitable and
therefore totally useless is probably the main cause of this situation, which
must be implemented with more carefully worded and detailed laws and
better education of the population.
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Suggestions for teaching
MARGHERITA SOLARI
■ Birdwatching in the mountains
Objectives: to recognise typical
species of mountain birds; to analyse
and compare the limiting factors of
various environments.
● Level: elementary or middle school
pupils (9-12 years).
● Equipment: suitable clothing for
excursions, reference books, manuals
for identifying birds, binoculars,
cameras.
● Collaborators: guides for excursions,
an expert ornithologist.
●
Alpine accentor (Prunella collaris)
PRELIMINARY STAGE
1. Discuss the characteristics of the mountain environment in class. Identify
the various mountain levels and their corresponding habitats: forests,
meadows and pastures, woodland, alpine belts, cliffs, screes, lakes, etc.
2. Divide the class into groups, each one of which should collect
photographs (photographs taken with the family, from books, Internet, etc.)
of slope and scree habitats. Discuss the characteristics of these
environments and the factors hindering the survival of animals and plants,
such as lack of water and soil, exposure to excessive sunlight and winds,
great temperature variations, instability and steepness of flanks, lack of
food, etc. Discuss the almost total absence of autophytes (plants capable of
synthesising their own food), the consequent lack of heterophytes (parasitic
plants) and why birds can reach these areas more easily than mammals,
amphibians and reptiles.
3 Collect references on the morphological features of Accipitridae, Corvidae
and Falconidae (colours of plumage, bill and feet, size, flight characteristics,
etc.). Examine the feeding habits of these birds. Compile entries enabling
Golden eagle (Aquila chrysaetos)
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140
pupils to recognise bearded vultures, eagles, peregrines, Alpine and Cornish
choughs and ravens. Describe in detail the objectives of the excursion, so that
pupils know how to behave responsibly.
141
EXCURSION
4. Identify a suitable mountain area for the excursion (which can be reached
by cable-car or after a short walk), along quiet paths far from crowded
mountain huts (the excursion should take place in early morning). Divide the
class into two or three groups, each one of which should have binoculars and
bird descriptions. Choose a panoramic spot for birdwatching.
5. Compile notes about the environment and types of birds (their flight,
behaviour, gregariousness, etc.) and take photographs.
FINAL PHASE
6. Each pupil should write a report, with the support of class discussion and
the notes taken during the excursion.
7. Follow-up in class: discuss the excursion and the difficulties involved in
birdwatching.
Screes in the Carnic Alps (Friuli Venezia Giulia)
■ Beware of screes !
Objectives: to analyse and describe the adaptations of high-altitude plants,
especially those living on screes, in order to arouse students’ awareness to
vegetation and its evolution.
● Level: middle or secondary school students (12-15 years).
● Equipment: references, stationery for board illustrations, suitable clothing for
excursions, cameras with zoom lenses, manuals to identify mountain flowers.
● Possible collaborators: excursion guides, or botanists, either prior to the
excursion or after it.
●
PRELIMINARY STAGE
Flight of a hunting eagle, which hovers in circles before plunging on its prey.
1. Class discussion on the elements defining mountain environments and their
typical ecological characteristics. Analysis of altitude levels. Identification of the
limiting ecological factors and characteristics of mountain screes (mean
temperature, variations due to altitude, annual and daily variations, together
with exposure to sunlight, freeze-thaw cycles, snow cover, lack of soil,
steepness and instability of flanks, reduced atmospheric pressure, lack of
water, etc.) through class discussion, also based on students’ personal
experience.
142
Migrating glareophytes
Long shoots penetrate well into the
substrate, parallel with the surface
(Campanula cochlearifolia,
Thlaspi rotundifolium, Viola calcarata)
Creeping glareous plants
Their small, foliate shoots rest on gravel
(Arenaria biflora, Linaria alpina,
Silene glareosa)
Rooting glareous plants
Their shoots extend in depth inside debris
(Doronicum, Hieracium intybaceum,
Oxyria digyna)
2. Analyse the adaptations of mountain plants due to the general ecological
factors and to the scree habitat in particular.
3. Divide the class into groups, each one of which should produce an
illustrated description of one of the adaptations examined (plants with rosettes
or cushion shapes, reduced size, roots, glossy leaves, presence of cuticles,
waxy or hairy surfaces, succulence, shortened vegetative periods, gaudy
flower colours).
4. Divide the class into five groups, ones different from the previous, to enable
exchange of information on the adaptations examined. Make charts (e.g.
100x70 cm) with drawings, diagrams and photographs showing the divisions
into migrating, creeping, rooting, mat-forming and stabilising glareous plants,
according to type of growth.
5. Explain the evolution of environmental and climax vegetation. Focus on the
role played by vegetation in stabilising screes.
6. With the help of a guide, choose a suitable itinerary for the excursion, which
should be easily accessible and within a short distance (one-and-a-half-hour
walk, according to students’ ability). Identify the objectives of the excursion
with the guide, avoiding excessive scientific information.
EXCURSION
7. Analyse mountain and scree environments (focusing on their limiting factors).
8. Identify the most typical plants according to adaptations, rare species, and
essential plants for the evolution of vegetation. Make observations with the
guide. Take photographs.
Mat-forming glareous plants
They root into debris and form extensive
covers
(Dryas octopetala, Gypsophila repens,
Saxifraga oppositifolia)
Stabilising glareous plants
They have very robust cushions and tufts
and thin roots
(Androsace alpina, Carex firma,
Ranunculus glacialis)
Subdivision of plants living on debris according to their type of growth
CONCLUSION OF CLASSWORK
9. Summarise the information collected. Complete the charts by attaching
photographs of the plants next to their group.
10.Discuss the characteristics of these environments, and the need to preserve
them.
Note: If the aim is also to introduce students to natural photography, essential
indications should be provided (unless a mini-course is feasible). When in the
mountains:
● use reflex cameras with different lenses, e.g. 50 mm, telephoto, wide-angle
and macro lens for flowers (modern digital cameras feature all these functions)
● with telephoto lenses, use short exposure times and closed shutters to focus
the whole flower.
● take photographs in diffused light, preferably at sunrise or sunset.
● use UV filters to avoid excessive UV radiation.
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■ Rocks, algae and lichens
Objectives: to arouse awareness of the diversity of plants and animals and
their adaptations to survival in differing environments. Identify possible
applications during the students’ course of study.
● Level: secondary school students (15-18 years).
● Equipment: references, photographs, school biology book, suitable
clothing, cameras.
● Collaborator: a guide for the excursion.
●
PRELIMINARY STAGE
1. Review study of prokaryotic and eukaryotic cells, autophytic and
heterophytic organisms and various types of metabolism. Analyse differing
kinds of unicellular organisms under the microscope and classify them.
2. Study the ways in which algae and epilithic plants adhere to the
substratum, whether they prefer siliceous or calcareous substrata
(petrography), limiting conditions (e.g. covered by sheets of water for lengthy
periods, even weeks, etc.), life-cycles and survival during drought.
3. Analyse the alga-fungus symbiosis in lichens and the role played by
phycobionts and mycobionts. Quote
examples of epilithic and endolithic
lichens.
4. Identify the systematic classification
of cyanobacteria in the school text
book. Analyse the subdivisions of
organisms into chemo-autophytes,
photo-heterophytes and chemoheterophytes according to their food
requirements,
and
review
the
classification of plants and bacteria
within the first category by studying
mechanisms of cellular structure and
metabolism.
5. Study karst phenomena and their
associated macro- and microforms,
the chemical balance regulating karst
Lichens
phenomena (role of temperature,
pressure, pH and CO2 in the reaction between carbonate and bicarbonate).
Discuss the phytokarstic phenomenon, the role of exolithic and endolithic
plants on limestone.
EXCURSION
6. Find a suitable area for the excursion. Divide the class into two or three
groups. Collect data and take notes.
7. Photograph lichens on rocky substrates.
CONTINUATION OF CLASSWORK
8. Collect the photographs and discuss data.
9. With the help of a guide or botanist, identify the lichens photographed.
This can also be done by logging on to special websites like dbiodbs.
univ.trieste.it of the Società Lichenologica Italiana (Italian Lichenological
Society), ospitiweb.indire.it, www.tamtamscuola.it/licheni, or www.dister.
unige.it/lablic.
10.Discuss the variety of living organisms and our ability to recognise forms of
life, like lichens, from what are seemingly insignificant elements.
Cliffs and rocky slopes can rise above towns (Valtellina, Lombardy)
Note: in order to arouse students’ interest in natural photography, see notes at
the end of the previous section.
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Select bibliography
Various Authors, 2004 - Italia. Atlante dei tipi geografici. (Italy. Atlas of Geographical Types). Istituto
Geografico Militare, Florence.
The atlas contains many illustrated sections of Italian geographical types, divided into 20 topics covering
both physical and human geography.
AESCHIMANN D., LAUBER K., MOSER D. M., THEURILLAT J.P., 2004 - Flora delle Alpi (Alpine Flora). 3 vols.
Zanichelli, Bologna.
The most recent, updated and complete guide to Alpine flora. The distribution and ecology of each
species are described with maps, diagrams and beautiful colour photographs. The volume contains
essential information for those wishing to study Alpine flora, which includes several cliff and debris
species of great phytogeographical interest.
BRICHETTI P., 1987 - Atlante degli uccelli delle Alpi italiane (Atlas of Italian Alpine birds). Ramperto, Brescia.
Although its information on distribution is dated, this is still the main textbook dedicated to montane birds
of the Italian Alps.
CARRETTA L., 1988 - Rapaci in volo (Raptors in flight). Pirella editore. Genoa.
The volume contains general information on the habits of raptors, especially their flight. With several
drawings based on site observation.
CARTON A., PELFINI M., 1988 - Forme del paesaggio d’alta montagna (High-altitude Landscape Forms).
Zanichelli, Bologna
A rigorously scientific textbook. Examines high-altitude lanscape forms and processes, with many
photographs and diagrams of the Alpine region.
CASTIGLIONI G. B., 1986 - Geomorfologia (Geomorphology). UTET, Turin.
A treatise on general geomorphology, analysing the morphogenetic systems of the world, with many
references to the geomorphology of Italy. Contains an extensive bibliography of Italian and international
works and journals of geomorphology.
FEOLI-CHIAPELLA L., 1983 - Prodromo numerico della vegetazione dei brecciai appenninici (“Warning
signals of reduction in numbers of plants of the Apennine chain”). “Collana del Progetto Finalizzato
“Promozione della Qualità dell’Ambiente” (series published by Italian Council for Research; project
“Promotion of Environmental Quality”), AQ/5/40, C.N.R., Rome.
An extensive and detailed scientific monograph on the phytosociological characterisation of vegetation
of scree slopes throughtout the Apennine chain.
FONTANA P., BUZZETTI F. M., COGO A., ODÈ B., 2002 - Cavallette, grilli, mantidi e insetti affini del Veneto
(“Grasshoppers, crickets, mantises and similar insects in the Veneto Region”). Museo Naturalistico
Archeologico di Vicenza, Vicenza.
A volume devoted to orthopterans, accompanied by a useful CD. Although the description of these
particular groups is limited to the Veneto, the publication may in fact be conveniently used for the whole
of Italy.
GENSBOL B., 1992 - Guida dei rapaci diurni d’Europa (Guide to diurnal European raptors). Zanichelli,
Bologna.
An authoritative work which synthesises information on the morphology, biology and ecology of several
Italian raptor species.
GRABHERR G., MUCINA L., 1993 - Die Pflanzengesellschaften Österreichs. Teil II. Natürliche waldfreie
Vegetation. Fischer Verlag, Jena.
A scientific guide in German to Austrian vegetation, also useful for classifying and describing cliff and
debris slope plants of the south-eastern Alps.
MINELLI A., CHEMINI C., ARGANO A., LA POSTA S., RUFFO A. (a cura di), 2002 - La fauna in Italia (Italian
fauna). Touring Club Italiano, Milan, and Italian Ministry of the Environment and Territory Protection,
Rome.
An advanced textbook containing fundamental information on Italian fauna, its legislative and
conservational aspects.
147
148
MINELLI A., RUFFO S., LA POSTA S., 1993-1995 - Checklist delle specie della fauna italiana (Checklist of
Italian fauna). Calderini, Bologna.
Lists all known species of Italian fauna, with authoritative, standard nomenclature. The series is in 110
parts. Material also available on website http://checklist.faunaitalia.it
149
Glossary
PINNA M., 1977 - Climatologia (Climatology). UTET, Turin.
General climatology, with analysis and classification of the world’s climates.
POLI MARCHESE E., 1965 - La vegetazione altomontana dell’Etna (Alpine vegetation on Mount Etna). Flora
et Vegetatio Italica, Mem. 5, Sondrio.
A historic scientific monograph on high-altitude vegetation of the largest Italian volcano, with essential
information on pioneer vegetation growing on lava.
REISIGL H., KELLER R., 1990 - Fiori e ambienti delle Alpi (Alpine flowers and habitats). Museo Tridentino di
Scienze Naturali, Trento.
A scientific textbook. Cliff and debris slope flora are clearly described with photographs and drawings.
Some of the drawings are used in this volume.
SPAGNESI M., TOSO S. (eds.), 1999 - Iconografia dei Mammiferi d’Italia (Iconography of Italian mammals).
Italian Ministry of the Environment and Territory Protection, National Institute for Wild Fauna A. Ghigi,
Rome.
The most complete and updated work on Italian mammals, with information on the morphology, ecology
and taxonomy of Italian populations, and detailed illustrations and distribution maps of all species.
> Alliance: in phytosociology, a grouping of
closely related associations.
> Angiosperm: flowering plant, generally
terrestrial.
> Antophagous: animal feeding on angiosperm
flowers.
> Apodal: footless animal (snakes, larvae of
certain insects, etc.).
> Association: basic unit of vegetation
characterised or differentiated by a given floristic
composition, with a precise association with the
environment; the basic phytosociological category.
> Biocoenosis: animal and plant community
sharing the same habitat.
> Biomass: body mass (dry weight) of one or
more organisms.
> Chasmophyte: a plant adapted to living in cliff
crevices.
> Class: the highest category of phytosociological
or syntaxonomic taxonomy; it includes one or
more orders.
> Clast: a single fragment of debris produced by
the disintegration of a larger rocky mass.
> Cryophilic: organism that develops in
microclimates with temperatures below 10°C.
> Diploid: organism having two identical sets of
chromosomes in each cell.
> Division, or phylum: in botany, the taxonomic
category between kingdom and class.
> Endemic: species with a distribution area
restricted to a particular geographical region.
> Entomocoenoses: the insect communities living
in a particular ecosystem.
> Euryecious: an organism that tolerates great
variations in the main parameters of the
environment in which it lives.
> Generalist: a non-specialised organism.
> Glareous plant: plant species living on debris
slopes.
> Imago: an insect in its final adult stage, sexually
mature and usually winged.
> Infacing slope: a landform commonly found in
regions of gently tilted sedimentary rocks and
consisting of an inclined upland, the slope of
which conforms with the dip of a resistant bed or
series of beds and a relatively steep escarpment
descending abruptly from its crest.
> Lithophilous: an organism adapted to living in
rocky environments.
> Mesophilous: an organism growing or thriving
best in an intermediate environment (e.g. one of
moderate temperature).
> Microphagous: an organism feeding on minute
particles.
> Microthermic: of, involving, or relating to very
small quantities of heat or changes of temperature.
> Monoploid: organism having only one set of
chromosomes in each cell.
> Monophagous: an organism feeding on or
utilising a single kind of food (generally a single
hosting plant).
> Neurotoxic: chemical substances that are
harmful or poisonous to the nerves or nervous
tissues.
> Order: a category of taxonomic classification
ranking above the family and below the class and,
in botany, characteristically having a name ending
in -ales (e.g. Rosales) and often made up of several
families. In phytosociological taxonomy, it includes
one or more alliances.
> Parasitoid: of, or relating to the relation existing
between various insect larvae and their hosts, in
which the larva feeds upon the living host tissues
in an orderly sequence, so that the host is not
killed until larval development is complete.
> Periglacial processes: typical processes in
areas with a cold climate, where snow and ice play
an important role, but where glaciers are not
involved.
> Phenology: in botany, the relation between
climate and periodic biological phenomena
culminating with the flowering of plants.
> Pheromonal:
relating
to
intraspecies
communication by means of chemical substances
which are perceived from a distance.
> Phyllophagous: an organism feeding on plant
leaves.
> Phytocoenosis (or plant communities): a whole
body of plants occupying a particular habitat.
> Phytophagous: an organism feeding on plants.
> Phytosaprophagous: an organism feeding on
decomposing plant parts.
> Phytosociology: the branch of ecology dealing
with the interrelations among the flora of particular
areas and especially with plant communities.
Plants are hierarchically classified in classes,
orders, alliances and associations.
> Pleistocene: the first of two periods of the
Quaternary era, from approx. 1.8 million to 10,000
years ago and characterised by several glaciations
and the first significative evidence of man’s
presence.
> Polyploid: organism having a chromosome
number that is a multiple greater than two of the
monoploid number.
> Productivity:
the
quantity
of
energy
accumulating in an ecosystem in the form of
biomass.
> Quaternary deposits: deposits of various nature
(fluvial, glacial, etc.), which accumulated in the last
2 million years (Quaternary epoch).
> Relict: a persistent remnant of an otherwise
extinct flora or fauna or kind of organism.
> Scapus: leafless stem bearing a flower or
inflorescence at its apex.
> Sciophilous: an organism living in shade.
> Shoot: the aerial part of a plant; a stem with its
leaves.
150
> Silicicolous: growing or thriving in soil rich in
silicon dioxide.
> Spermatophagous: an organism living on plant
seeds.
> Sub-Mediterranean: an organism that colonises
inland Mediterranean environments.
> Symbiosis: interaction of two dissimilar
organisms in any of several mutually beneficial
relationships.
> Tectonics: the study of regional structural and
deformational features of the Earth’s crust caused
by deep forces.
> Thallus: a plant body that shows no
differentiation into distinct tissue systems or
members (such as stems, leaves, or roots).
> Thermal capacity: quantity of heat required to
increase the temperature of an object specifically, of rock and debris - by 1°C.
> Transpiration: the emission or exhalation of
water vapour from the surfaces of leaves or other
parts of plants.
> Vascular plant: a plant having a specialised
conductive system that includes xylem and
phloem. Vascular plants include ferns and seeding
plants.
> Vicariate: relating to two taxonomic or
syntaxonomic organisms that replace each other
in bordering geographical areas or similar
ecological environments.
> Xerophilous: an organism living in habitats with
a dry microclimate.
> Xerothermophilous: an organism living in
habitats with a dry, warm microclimate.
> Xylophagous: an organism feeding on or in
wood.
> Zoocoenosis: the animal community occupying
a particular ecosystem.
151
List of species
Acalypta musci - 91
Acarospora - 30
Achillea atrata - 70
Achillea barrelieri - 67, 71
Achillea erba-rotta - 63, 69
Achillea erba-rotta ssp. ambigua
- 69
Achillea lucana - 51, 71, 135
Achillea mucronulata - 71
Achillea nana - 69
Achillea rupestris - 71
Adder - 114
Adenostyles leucophylla - 69
Adiantum - 45
Adiantum capillus-veneris - 34,
45
Admontia cepelaki - 110
Adonis distorta - 64, 71, 135
Adriatic bellflower - 49
Aeropedellus variegatus - 101
Aeropus sibiricus - 101
Agolius - 107
Agolius abdominalis - 107
Agrodiaetus - 96
Agrodiaetus galloi - 96
Agrodiaetus ripartii - 96
Alder - 129
Allioni’s primrose - 47, 53, 81
Allium narcissiflorum - 69
Alpine accentor - 120, 139
Alpine bladder fern - 47
Alpine chough - 115, 115, 140
Alpine fescue - 44
Alpine gypsophila - 62, 63
Alpine marmot - 107, 126, 127
Alpine rock-jasmine - 43
Alpine salamander - 113
Alpine scurvy-grass - 64
Alpine shrew - 127
Alpine toadflax - 62, 63
Alydus rupestris - 91
Alyssoides - 26
Alyssoides utriculata - 26
Alyssum - 63, 108
Alyssum argenteum - 69
Alyssum cuneifolium - 71
Alyssum diffusum - 81, 95
Alyssum ligusticum - 68
Alyssum ovirense - 70
Alyssum wulfenianum - 70
Amara - 103
Amphimallon - 94
Anaglyptus gibbosus - 82
Ancient king - 42, 47, 48, 49, 55
Andreaea - 33
Androsace alpina - 43, 142
Androsace brevis - 69
Androsace hausmannii - 53, 70
Androsace helvetica - 43
Androsace mathildae - 51, 71,
135
Androsace pubescens - 69
Androsace vandellii - 43, 55
Androsace vitaliana ssp.
praetutiana - 71
Androsace vitaliana ssp. sesleri 70
Androsace wulfeniana - 70
Anechura bipunctata - 102
Anonconotus - 91, 102
Anoxia - 94
Anoxia australis - 94
Antaxius difformis - 89, 102
Anthaxia - 95
Anthemis aetnensis - 71
Anthocaris - 96
Anthocaris euphenoides - 96
Antirrhinum latifolium - 93
Antitype - 97
Apennean pennycress - 64
Apennine cat’s ear - 66
Apennine fescue - 67
Apennine pheasant’s-eye - 64,
67
Apennine primrose - 50, 51, 55
Aphanothece - 28
Apion - 108
Apuan globularia - 39, 50
Aquila chrysaetos - 119, 138
Aquilegia champagnatii - 71, 135
Aquilegia einseleana - 70
Aquilegia magellensis - 71
Aquilegia ottonis - 70, 81
Arabis - 92
Aradus frigidus - 103
Arctia festiva - 110
Arctia flavia - 111
Arenaria biflora - 142
Arenaria huteri - 69, 81
Argentera pansy - 63
Arhopalus ferus - 82
Arianta chamaeleon - 99
Armadillidium - 88, 99
Armeria gussonei - 71, 135
Armeria morisii - 51, 71
Armeria sulcitana - 71
Artemisia glacialis - 69
Artemisia nitida - 70
Artemisia petrosa ssp. eriantha 70
Asida pirazzolii - 106
Asida pirazzolii ssp. sardiniensis 106
Asperula gussonei - 51, 71
Asperula hexaphylla - 68
Asperula pumila - 55, 71
Asplenium - 45, 46, 53
Asplenium cuneifolium - 46
Asplenium fontanum - 46
Asplenium lepidum - 46
Asplenium ruta-muraria - 46
Asplenium seelosii - 46
Asplenium septentrionale - 46
Asplenium trichomanes - 46
Asplenium viride - 46
Astragalus siculus - 91
Athamanta - 66
Athamanta cretensis - 66
Athamanta turbith - 69, 70
Athamanta vestina - 69
Atractosoma meridionale - 100
Aubrieta deltoidea - 71
Aurinia - 93, 108
Aurinia saxatilis - 81
Aurinia saxatilis ssp. orientalis 95
Axinopalpis gracilis - 82
Ballota frutescens - 68, 135
Ballota rupestris - 93
Barbula bicolor - 33
Barbula crocea - 35
Barrelieri’s sneezewort - 67
Bearded vulture - 119, 135, 137
Beech - 80, 82
Berardia subacaulis - 63, 65, 135
Berardie - 63, 65
Bergamosoma canestrinii - 99
Berninelsonius hyperboreus 107
Biscutella - 95, 104
Black Forest spleenwort - 46
Black redstart - 119, 120
Black-eared wheatear - 120
Black-eared wheatear - 83
Bladder campion - 62
Blue rock-thrush - 83
Blue-leaved buckthorn - 54
Bombus - 110
Bonelli’s eagle - 83
Bosnian pine - 95
Brachyodontus - 108
Brachypterolus - 93, 105
Brachypterolus linariae - 93, 105
Brachypterolus vestitus - 93
Brassica repanda - 69
Brembosoma castagnolense 88
Brittle bladder fern - 47, 53
Bumblebee - 110
Buprestis splendens - 94, 95, 136
Busambra cornflower - 51
Calliptamus italicus - 101
Calliptamus siciliae - 101
Caloplaca - 30
Calothrix - 28
Campanula - 47, 50, 63
Campanula albicans - 68
152
Campanula alpestris - 63, 69
Campanula carnica - 70
Campanula cenisia - 69
Campanula cochleariifolia - 27,
61, 62, 142
Campanula elatines - 49, 69
Campanula elatinoides - 69
Campanula excisa - 69
Campanula forsythii - 71
Campanula fragilis ssp. cavolinii
- 51, 70
Campanula macrorrhiza - 68
Campanula morettiana - 49, 50,
69, 135
Campanula petraea - 69
Campanula pollinensis - 71
Campanula raineri - 50, 69, 135
Campanula tanfanii - 70
Campanula thyrsoides ssp.
carniolica - 70
Campanula zoysii - 39, 40, 70
Candidula unifasciata - 99
Capra ibex - 112, 124
Carabus - 92, 103
Carabus (Orinocarabus)
pedemontanus - 92
Carabus baudii - 103
Carabus bertolinii - 92
Carabus concolor - 103
Carabus creutzeri - 103
Carabus depressus - 103
Carabus heteromorphus - 103
Cardaminopsis - 92
Carduus chrysacanthus - 71
Carex firma - 142
Carinthian mouse-ear - 70
Carnic sandwort - 81
Caryocolum - 111
Catharia pyrenaealis - 97
Catterpillar - 111
Centaurea busambarensis - 51,
71
Centaurea scannensis - 70
Centranthus trinervis - 71
Cepaea sylvatica - 99
Cephalaria mediterranea - 71
Cerambyx scopolii - 82
Cerastium - 104, 109
Cerastium carinthiacum - 70
Cerastium subtriflorum - 70
Cerastium thomasii - 70
Ceterach - 45
Ceterach officinarum - 45
Ceutorhynchus - 108
Ceutorhynchus bifidus - 108
Ceutorhynchus inaffectatus - 108
Ceutorhynchus pinguis - 81, 95
Ceutorhynchus verticalis - 81, 95
Chalicodoma muraria - 106
Chamaesphecia - 97
Chamois - 72, 107, 123, 124, 136
Charpentieria - 83
Chazara briseis - 97
Cheilosia aristata - 109
Chelidura aptera - 102
Chelidurella - 102
Chelidurella thaleri - 102
Chelidurella vignai - 102
Chelis maculosa - 110
Chersotis alpestris - 97
Chersotis ocellina - 97
Chilostoma - 83, 87
Chilostoma alpinum - 99
Chilostoma cingulatum - 87, 99
Chilostoma cingulatum
colubrinum - 99
Chilostoma millieri - 99
Chilostoma zonatum - 99
Chionomys nivalis - 125, 127
Chondrina - 87
Chondrina avenacea - 86, 87
Chondrina clienta - 87
Chondrina megacheilos - 87
Chondrina oligodonta - 87
Chopardius pedestris - 91, 102
Chopardius pedestris ssp.
apuanus - 89
Chorthopodisma cobellii - 101
Chough - 115
Cirsium bertolonii - 64, 70
Clausilia - 83
Cloak fern - 45
Cobweb houseleek - 44
Cobweb saxifrage - 41, 50
Cochlostoma - 83, 86
Cochlostoma canestrinii - 86
Cochlostoma crosseanum - 86
Cochlostoma henricae - 86
Cochlostoma paladilhianum - 86
Cochlostoma philippianum - 86
Cochlostoma porroi - 86
Cochlostoma sardoum - 86
Cochlostoma scalanum - 86
Cochlostoma tergestinum - 86
Cochlostoma villae - 86
Coincya richeri - 69
Colpotus strigosus - 106
Colpotus strigosus ssp.
ganglbaueri - 106
Columbine - 81
Comolli’s pansy - 63
Condrina - 83
Conocephalum conicum - 34
Copium - 82, 91, 102
Copium clavicorne - 102
Copium teucrii - 102
Coranus subapterus - 91
Cornish chough - 115, 116, 136,
140
Corvus corax - 83, 116
Crag martin - 116, 117
Creeping avens - 61
Crepis terglouensis - 70
Crimped bellflower - 39, 40, 50
Cryphia - 97
Crypticus quisquilius - 106
Crypticus quisquilius ssp.
aprutianus - 106
Cryptogramma crispa - 61, 62
Ctenicera - 107
Ctenicera pectinicornis - 107
Cychrus - 103
Cychrus angulicollis - 103
Cychrus cylindricollis - 103
Cychrus graius - 103
Cychrus schmidti - 103
Cymbalaria pallida - 71
Cymindis - 103
Cystopteris - 45, 47, 53
Cystopteris alpina - 47
Cystopteris dickiaeana - 47
Cystopteris fragilis - 47, 53
Dactylophorosoma nivisatelles 100
Dahlica - 97
Danacea - 105
Danacea nigritarsis - 93
Daphne petraea - 53, 69, 135
Dasytes lombardus - 93
Deroplia genei - 82
Devil’s claw - 134
Dianthus - 97
Dibolia rugulosa - 108
Dichotrachelus - 95, 108
Dickie’s fern - 47
Didymodon tophaceus - 34
Dimorphocoris poggii - 91
Dolomite cinquefoil - 54
Dolomite spleenwort - 46
Doronicum - 108, 142
Doronicum clusii - 62
Doronicum glaciale - 70
Doronicum grandiflorum - 62
Draba aizoides - 55
Draba dolomitica - 69
Draba olympicoides - 71
Dracocephalum - 105
Dryas - 104
Dryas octopetala - 62, 104, 142
Dryopteris villarii - 66
Drypis spinosa - 67, 71
Dwarf buckthorn - 52
Dwarf rampion - 49
Dyscia - 97
Dyscia raunaria - 97
Dyscia sicanaria - 97
Eagle - 140
Ectobius montanus - 102
Egyptian vulture - 83
Elophos - 111
Elophos caelibaria - 111
Elophos zelleraria - 111
Empetrum - 91
Ensifera - 101
Ephedra major - 26, 81, 81
Epilobium - 111
Epipodisma pedemontana - 100
Epipsilia grisescens - 97
Erebia - 110
Erebia calcaria - 97, 136, 137
Erebia christi - 97, 137
Erebia gorge - 110
Erebia meolans - 96
Erebia montana - 96, 97
Erebia pluto - 110, 111
Erebia scipio - 96
Erebia styria - 96
Erebia styx - 96
Erysimum - 93, 108
Euchloe - 96
Euchloe bellezina - 96
Eucladium verticillatum - 34
Eucobresia - 99
Eudarcia - 97
Eumenes - 82
Eupithecia - 97
Eupithecia venosata - 97
Euryopicoris nitidus - 91
Eusphalerum - 104
Eusphalerum albipile - 104
Eusphalerum angusticolle - 104
Eusphalerum annaerosae - 104
Eusphalerum pulcherrimum - 104
Euxoa culminicola - 111
Euxoa decora - 97
Euzonitis - 94
Euzonitis quadrimaculata - 94
Euzonitis terminata - 94
Evergreen oak - 80, 82, 95
Exocentrus lusitanus - 82
Fagus sylvatica - 80
Fairy’s thimble - 27, 61, 62
Falco biarmicus - 83
Falco peregrinus - 117
Falco tinnunculus - 117
Falcon - 117
Festuca alpina - 44
Festuca austrodolomitica - 69
Festuca dimorpha - 67, 71
Festuca laxa - 70
Festuca stenantha - 70
Fiori’s butterwort - 81
Forficula apennina - 102
Forked spleenwort - 46
Formica - 110
French sorrel - 61, 67
Galeopsis reuteri - 68
Galium magellense - 71
Galium margaritaceum - 70
Galium montis-arerae - 69
Galium noricum - 70
Galium pseudohelveticum - 69
Galium saxosum - 68
Galium tendae - 68
Geocoris grylloides - 102
Geocoris lapponicus - 103
Geum reptans - 61
Glacier buttercup - 62
Glacies - 111
Glacies alticolaria - 111
Glacies canaliculata - 111
Glacies coramica - 111
Globularia incanescens - 39, 70
Globularia neapolitana - 71
Gloeocapsa - 28
Gloeocapsa ralfsiana - 28
Gloeocapsa sanguinea - 28
Glomeris helvetica - 88
Glyptobothrus - 89
Glyptobothrus alticola - 90, 101
Glyptobothrus binotatus daimai 101
Glyptobothrus brunneus
brunneus - 101
Glyptobothrus eisentrauti - 101
Glyptobothrus mollis ignifer - 89
Gnophos - 97
Gnophos obfuscatus - 97
Golden eagle - 119, 131, 136,
138
Granaria stabilei - 99
Green spleenwort - 46
Grigne primrose - 50
Grimmia - 32, 33, 35
Grimmia alpestris - 32
Grimmia anodon - 32
Grimmia anomala - 32
Grimmia apiculata - 32
Grimmia arenaria - 32
Grimmia atrata - 32
Grimmia caespiticia - 32
Grimmia crinita - 32
Grimmia curviseta - 32
Grimmia limprichtii - 32
Grimmia montana - 32
Grimmia ovalis - 32
Grimmia pilosissima - 32
Grimmia pitardii - 32
Grimmia sessitana - 32
Grimmia teretinervis - 32
Grimmia tergestina - 32
Grimmia torquata - 32
Gussone’s woodruff - 51, 54
Gypaetus barbatus - 119, 137
Gypsophila repens - 62, 63, 142
Hadula melanocapa - 97
Hadula odontites - 97
Hairy pearlwort - 64
Hairy primrose - 55
Hairy spiraea - 81
Hare - 125
Harpalus - 103
Hausmann’s rock-jasmine - 53
Heart-leaved micromeria - 55
Helianthemum - 104
Helianthemum lunulatum - 68
Helichrysum - 51
Helichrysum frigidum - 52, 71
Helichrysum montelinasanum 51, 52, 71, 135, 135
Helichrysum nebrodense - 54, 71
Heracleum pyrenaicum ssp.
orsinii - 71
Herniaria litardierei - 71
Hesperis laciniata - 105, 108
Hieraaetus fasciatus - 83
Hieracium intybaceum - 142
Hieracium portanum - 71
Hipparchia alcyone - 96
Hipparchia neomiris - 96
Hipparchia statilinus - 96
Holoarctia cervini - 110
Hornbeam - 82
Horvath’s rock lizard - 114
Hose-horned viper - 114
Hyles vespertilio - 111
Hymenostylium recurvirostre - 34
Hypnoidus consobrinus - 107
Hypnoidus riparius - 107
Hypnoidus rivularius - 107
Hypnum dolomiticum - 33
Hyponephele lycaon - 96
Hypsoiulus alpivagus - 88
Iberis - 108
Iberis nana - 68
Iberolacerta horvathi - 114
Ibex - 112, 124, 136
Isatis alpina - 68
Ischyroptera bipilosa - 109
Italopodisma - 101
Jovibarba - 43
Jovibarba allionii - 68
Jovibarba arenaria - 70
Juniper - 100
Kestrel - 117, 140
Kickxia - 93, 105
Lagopus mutus - 121
Lanner - 83
Larch - 105
Large alpine salamander - 113
Large-flowered bellflower - 63
Laserpitium garganicum - 55
Laserpitium gaudinii - 70
Lasius - 110
Latipalpis plana - 82, 95
Lavandula angustifolia - 92
Lax potentilla - 52
Lecanora - 30
Lecidea - 30
Leiosoma - 109
Leistus glacialis - 104
Leontodon anomalus - 70
Leontodon montanus - 62
Leopardsbane - 62
Leptoiulus (Kolpophylacum)
helveticus - 88
Leptoiulus alemannicus - 100
Leptoiulus riparius - 100
Leptopterix - 97
Leptothorax - 95
Leptusa - 104
Leptusa angustiarumberninae
rosaorum - 104
Leptusa areraensis - 104
Leptusa baldensis - 104
Leptusa braccati - 104
Leptusa cavallensis - 104
Leptusa ceresoleana
ceresoleana - 104
Leptusa fauciumberninae - 104
Leptusa grignaensis - 104
Leptusa knabli recticollis - 104
Leptusa mandli - 104
Leptusa manfredi - 104
Leptusa montispasubii settei 104
Leptusa montiumcarnorum - 104
Leptusa occulta - 104
Leptusa piceata - 104
Leptusa portusnaoniensis - 104
Leptusa pratensis - 104
Leptusa rhaetoromanica - 104
Leptusa rosai - 104
Leptusa sudetica - 104
Leptusa tirolensis tirolensis - 104
Leptusa tridentina - 104
153
154
Leptusa trumplinensis - 104
Leptusa vallisvenyi - 104
Lepus timidus - 124, 125
Leucanthemopsis alpina - 109
Leucanthemum atratum ssp.
ceratophylloides - 68
Leucanthemum atratum ssp.
coronopifolium - 69
Leucanthemum atratum ssp.
halleri - 70
Leucanthemum laciniatum - 71
Leucostigma - 83
Leucostigma candidescens - 87
Lichen - 29, 145
Licinus italicus - 103, 104
Ligusticum ferulaceum - 68
Limbara everlasting - 52
Limonium morisianum - 71
Linaria - 93, 105
Linaria alpina - 62, 63, 142
Linaria tonzigii - 64, 69, 135
Linden - 82
Lithobius lucifugus - 89, 100
Lithobius muticus - 100
Lithobius nodulipes - 100
Lithobius schuleri - 100
Livelong saxifrage - 42
Lizard - 113, 114
Loiseleuria - 110
Longitarsus obliteratoides - 108
Longitarsus obliteratus - 108
Longitarsus springeri - 108
Lonicera stabiana - 71
Luffia - 97
Lunaria annua - 92
Lychnis - 97
Macroglossum stellatarum - 97
Macularia - 87
Madonie everlasting - 54
Maidenhair - 34, 45
Maidenhair spleenwort - 46
Malcolmia orsiniana - 71
Malthodes - 105
Malthodes atratus - 105
Malthodes atratus samniticus 105
Malthodes penninus icaricus 105
Malthodes trifurcatus - 105
Malthodes trifurcatus
atramentarius - 105
Maritime Alps pansy - 63
Marmorana - 83, 87
Marmorana fuscolabiata - 88
Marmorana globularis - 87
Marmorana muralis - 87
Marmorana nebrodensis - 87
Marmorana platychela - 87
Marmorana saxetana - 88
Marmorana scabriuscula - 87
Marmorana signata - 87
Marmota marmota - 126, 127
Marsupella - 33
Mathilda’s rock-jasmine - 51
Medora - 83, 87
Medora albescens - 87
Medora dalmatina - 87
Megasema ashworthii - 97
Melanoplus frigidus - 101
Meligethes - 92, 104
Meligethes aeneus - 104
Meligethes arankae - 105
Meligethes chlorocyaneus - 93
Meligethes devillei - 105
Meligethes erysimicola - 93, 104
Meligethes fumatus - 92
Meligethes lindbergi - 81, 92
Meligethes lunariae - 92
Meligethes nuragicus - 82, 92
Meligethes oreophilus - 105
Meligethes reyi - 104
Meligethes salvan - 105
Meligethes scholzi - 82, 93
Meligethes solidus - 104
Meligethes spornrafti - 92
Meligethes subaeneus - 92
Meligethes subfumatus - 82, 92,
105
Meligethes tener - 93
Meloe aegyptius - 106
Meloe erythrocnemus - 105
Melosira - 28
Mesoxyonyx osellanus - 81
Metaxmeste - 97
Metzgeria - 33
Micrabris - 106
Micrabris flexuosa - 106
Micrabris pusilla - 106
Microlestes - 92
Micromeria cordata - 55
Micromeria marginata - 68
Minuartia austriaca - 70
Minuartia cherlerioides - 70
Minuartia graminifolia - 69
Minuartia grignensis - 69
Minuartia rupestris ssp.
clementei - 69
Minuartia verna ssp. grandiflora 71
Mitopus morio - 77
Moehringia bavarica - 53
Moehringia bavarica ssp.
bavarica - 70
Moehringia bavarica ssp.
insubrica - 69
Moehringia concarenae - 69
Moehringia dielsiana - 69, 135
Moehringia glaucovirens - 69
Moehringia lebrunii - 68, 135
Moehringia markgrafi - 69, 135
Moehringia papulosa - 71, 81,
135
Moehringia sedifolia - 68, 135
Moltkia - 54
Moltkia suffruticosa - 54, 70, 135
Monte Alben primrose - 50
Monticola solitarius - 83
Montifringilla nivalis - 120
Moretti’s bellflower - 49, 50
Moris’s thrift - 51
Moss campion - 42, 43
Mount Linas everlasting - 51
Mountain avens - 62
Mountain hare - 124, 125
Mountain hawkbit - 62, 66
Mountain pine - 129
Mountain sorrel - 62
Mt. Cenis pansy - 69
Mt. Nebrodi joint pine - 26, 81
Murbeckiella zanonii - 64, 70
Mustela erminea - 125
Muticaria - 83
Mutilla europaea - 110
Mylabris - 106
Mylabris variabilis - 106
Myrmeleotettix maculatus - 101
Neagolius - 107
Neagolius amblyodon - 107
Neagolius liguricus - 108
Neagolius limbolarius - 107
Neagolius montanus - 107
Neagolius penninus - 108
Neagolius pollicatus - 107
Neagolius schlumbergeri - 108
Neapolitan bellflower - 51, 54
Nebria germari - 103
Nebria orsinii - 103
Neophron percnopterus - 83
Neoplinthus - 109
Nilepolemis - 109
Norway spruce - 105
Nostoc - 28
Notholaena - 45
Notholaena marantae - 45
Nudaria mundana - 97
Ocydromus - 103
Ocys - 92
Oedipoda - 101
Oedipoda caerulescens - 101
Oedipoda germanica - 98, 101
Oenanthe hispanica - 83
Oenanthe oenanthe - 120
Oeneis - 110
Oligomax - 99
Onobrychis - 96
Opatrum dahli - 107
Opatrum nivale - 107
Oreas martiana - 33
Oreina sibylla - 108
Oreina viridis - 108
Orenaia - 97
Oreonebria - 103
Oreopsyche - 97, 111
Oreopsyche leschenaulti - 109
Oreorhynchaeus - 108
Oreorhynchaeus baldensis - 108
Oreorhynchaeus focarilei - 108
Oreorhynchaeus pacei - 108
Oreorhynchaeus spectator - 108
Orinocarabus - 92, 103
Osellaeus bonvouloiri - 95
Osellaeus bonvouloiri ssp.
baldensis - 95
Osellaeus bonvouloiri ssp.
bonvouloiri - 95
Osmunda regalis - 34
Otiorhynchus - 109
Oxyria digyna - 62, 142
Oxytropis fetida - 69
Paederota bonarota - 70
Paederota lutea - 70
Papaver degenii - 71, 135
Papaver ernesti-mayeri - 70
Papaver kerneri - 70
Papaver rhaeticum - 66
Papilio alexanor - 83, 96
Papillifera - 83
Parietaria - 83
Parnassius apollo - 75, 84, 96,
136
Parnassius apollo pumilus - 96
Parsley fern - 61, 62, 67
Partridges - 107
Patrobus septentrionalis - 103
Pedicularis aspleniifolia - 70
Pelenomus hygrophilus - 95
Pellia - 33
Pellia endiviifolia - 34
Peregrine - 78, 117, 131, 136, 140
Petractis clausa - 30
Pezzotettix giornai - 101
Phenacolimax - 99
Phoenicurus ochruros - 120
Phyllotreta - 95
Phyllotreta atra - 95
Phyllotreta ganglbaueri - 95
Physoplexis comosa - 53, 70,
134, 135
Phyteuma cordatum - 68
Phyteuma hedraianthifolium 49, 69
Phyteuma humile - 49, 69
Phyteuma sieberi - 70
Piedmont saxifrage - 55, 68
Pieris callidice - 96
Pinguicula fiorii - 71, 81
Pinguicula hirtiflora - 71
Pinguicula poldinii - 70, 81
Pinguicula reichenbachiana - 81
Pink cinquefoil - 52, 53
Pinus - 94
Pinus leucodermis - 95
Plagiotylus ruffoi - 91
Platycarabus - 103
Platynus - 104
Poa laxa - 62
Podisma - 101
Poldini’s butterwort - 81
Polistes biglumis bimaculatus 110
Polyblastia - 30
Polydesmus monticola - 100
Polygala carueliana - 70
Polygonia egea - 83
Potentilla apennina - 54, 71
Potentilla caulescens - 52, 53
Potentilla clusiana - 70
Potentilla crassinervia - 71
Potentilla grammopetala - 69
Potentilla nitida - 52, 53, 70
Potentilla saxifraga - 68, 135
Preissia - 33
Preissia quadrata - 34
Primula - 47, 50
Primula albenensis - 50, 69
Primula allionii - 47, 68, 135
Primula apennina - 50, 51, 70
Primula grignensis - 50, 69
Primula hirsuta - 55
Primula marginata - 69
Primula recubariensis - 50, 69
Primula spectabilis - 53
Primula tyrolensis - 69
Pritzelago alpina ssp.
austroalpina - 70
Protoblastenia - 30
Protoblastenia immersa - 30
Protoblastenia incrustans - 30
Prunella collaris - 120, 139
Pseudobankesia - 97
Pseudocraspedosoma
grypischium - 99
Pseudotergumia fidia - 96
Psophus stridulus - 101
Psylliodes instabilis - 108
Psylliodes picipes - 95
Psylliodes toelgi - 95
Pterostichus - 103
Ptilostemon niveus - 64, 71
Ptilotrichum cyclocarpum - 54
Ptyonoprogne rupestris - 116,
117
Purple saxifrage - 7
Pyramidal saxifrage - 55
Pyramidula - 85, 86
Pyramidula pusilla - 83, 86
Pyramidula rupestris - 83
Pyrenean chamois - 123, 124,
136
Pyrrhocorax graculus - 115
Pyrrhocorax pyrrhocorax - 115
Quercus ilex - 80
Racomitrium - 33
Rainer’s bellflower - 50
Ranunculus glacialis - 62, 142
Ranunculus magellensis - 71
Ranunculus traunfellneri - 64, 70
Raven - 83, 116, 140
Recoaro primrose - 50
Reichenbach’s butterwort - 81
Rhacocleis - 102
Rhaetian poppy - 66
Rhaetian rampion - 49
Rhamnus glaucophyllus - 54, 70,
135
Rhamnus pumilus - 52
Rhizobotrya alpina - 64, 69, 135
Rhizocarpon - 30
Rhizocarpon geographicum - 31
Rhizotrogus - 94
Rhododendron - 100
Rhododendron - 110
Rhyacia helvetina - 97
Rigid buckler fern - 66
Robertia taraxacoides - 66
Rock alyssum - 54
Rock catchfly - 55
Rock mezereon - 69
Rock ptarmigan - 121, 136
Rohdendorfia alpina - 109
Rosalia alpina - 82, 137
Rothenbuehleria minima - 100
Round-leaved pennycress - 61,
63
Royal fern - 34
Rumex scutatus - 61
Rupestrella - 83, 86
Rupicapra pyrenaica ornata - 123
Rupicapra rupicapra - 72, 123
Rustyback - 45
Sagina pilifera - 64, 71
Salamandra atra - 113
Salamandra lanzai - 113
Salix crataegifolia - 70, 135
Saponaria sicula - 64, 71
Sardinian woodruff - 55
Sarromyia nubigena - 109
Satureja montana - 92, 108
Satyrus ferula - 97
Saussurea alpina ssp. depressa
- 69
Saxifraga - 47, 63, 95
Saxifraga aizoides - 27, 95
Saxifraga ampullacea - 71
Saxifraga aphylla - 70
Saxifraga arachnoidea - 41, 53,
69, 135
Saxifraga biflora - 65
Saxifraga burseriana - 70
Saxifraga callosa - 53, 54, 69
Saxifraga cervicornis - 71
Saxifraga cochlearis - 68, 135
Saxifraga cotyledon - 55
Saxifraga crustata - 70
Saxifraga depressa - 67, 69
Saxifraga diapensioides - 69
Saxifraga facchinii - 69
Saxifraga florulenta - 42, 48, 68,
135
Saxifraga glabella - 71
Saxifraga hostii - 70
Saxifraga italica - 71
Saxifraga lingulata see Saxifraga
callosa - 53
Saxifraga oppositifolia - 7, 142
Saxifraga paniculata - 42
Saxifraga paniculata ssp.
stabiana - 71
Saxifraga pedemontana - 55, 68
Saxifraga petraea - 70
Saxifraga porophylla - 71
Saxifraga presolanensis - 69
Saxifraga retusa ssp. augustana
- 69
Saxifraga sedoides - 70
Saxifraga speciosa - 71
Saxifraga squarrosa - 70
Saxifraga tenella - 70
Saxifraga tombeanensis - 69, 135
Saxifraga valdensis - 49, 69
Saxifraga vandellii - 53, 69
Sceliphron - 82
Schistidium - 33
Sciadia tenebraria - 111
Scleranthus annuus ssp.
aetnensis - 71
155
156
Scleranthus vulcanicus - 71
Scolitantides orion - 110
Scytonema - 28
Sedum - 43, 96, 109, 110
Sedum aetnense - 71, 135
Sedum alsinefolium - 68
Sedum brevifolium - 71
Sedum fragrans - 69
Segestria - 83
Segestria bavarica - 83
Segestria florentina - 83
Segestria senoculata - 83
Selatosomus - 107
Selatosomus aeneus - 107
Selatosomus amplicollis - 107
Sempervivum - 43, 96
Sempervivum arachnoideum - 44
Senecio aetnensis - 71
Senecio ambiguus - 71
Senecio candidus - 71
Senecio rupestris - 108
Serica brunnea - 94
Serpentine spleenwort - 46
Sesleria ovata - 70
Setina - 97
Sheep - 107
Short-haired sandwort - 81
Shrubby horehound - 81
Sicilian soapwort - 64, 71
Siciliaria - 83, 87
Sideritis kitti — 97
Silene acaulis - 42, 43
Silene campanula - 68
Silene cordifolia - 68
Silene elisabethae - 69, 135
Silene glareosa - 142
Silene lanuginosa - 70
Silene quadrifida - 70
Silene requienii - 71
Silene rupestris - 55, 109
Silene saxifraga var. lojaconoi - 71
Silene veselskyi - 70
Silene vulgaris ssp. glareosa - 62
Silver oat-grass - 62
Simple-leaved milfoil - 63
Sitaris muralis - 94
Snow finch - 120
Snow vole - 125, 127
Snowny thistle - 64, 67
Solatopupa - 83, 87
Solatopupa guidoni - 87
Solatopupa juliana - 87
Solatopupa pallida - 87
Solatopupa psarolena - 87
Solatopupa similis - 87
Soldanella minima - 70
Sorex alpinus - 127
Southern sermountain - 55
Spaelotis senna - 97
Spiny carnation - 67
Spiraea decumbens - 81
Spiraea decumbens ssp.
decumbens - 70
Spiraea decumbens ssp.
hacquetii - 69
Stachys annua - 108
Standfussiana lucernea - 97
Staurothele - 30
Staurothele immersa - 30
Stenhomalus bicolor - 82
Stenobothrodes cotticus - 89,
101
Stenobothrodes rubicundulus 101
Stenobothrus apenninus - 102
Stenobothrus fischeri - 101
Stenoria - 94
Stenoria analis - 94
Stenoria apicalis - 94
Stigonema - 28
Stoat - 125
Stromatium unicolor - 82
Swiss rock-jasmine - 43
Sympistis - 111
Syngrapha devergens - 97
Syngrapha hochenwarthi - 97
Syntomus - 92
Tabellaria - 28
Tandonia simrothi - 99
Telekia speciosissima - 53, 69
Testediolum - 103
Tetramorium - 110
Tetrix bipunctata - 100
Teucrium - 82, 91, 102, 103
Teucrium chamaedrys - 102
Teucrium flavum - 81, 92, 102
Teucrium massiliense - 92
Teucrium montanum - 102, 103
Teucrium scorodonia - 102
Thelidium - 30
Thick-leaved saxifrage - 53, 54
Thlaspi alpestre - 70
Thlaspi rotundifolium - 61, 63,
64, 142
Thlaspi rotundifolium ssp.
cepaeifolium - 70
Thlaspi rotundifolium ssp.
corymbosum - 69
Thlaspi rotundifolium ssp.
grignense - 69
Thlaspi stylosum - 64, 71
Thymus - 105, 108
Tichodroma muraria - 118, 119
Tonzig’s toadflax - 63, 64
Tortella - 33
Tortella tortuosa - 33
Tortula - 33
Trachystyphlus alpinus - 108
Traunfellner’s buttercup - 64, 70
Trechus - 103
Trechus strigipennis - 103
Trentepohlia - 28
Trichoferus holosericeus - 82
Trichoferus spartii - 82
Trinia - 96
Trisetum argenteum - 62, 70
Trisetum distichophyllum - 62
Tufted meadow-grass - 62
Two-flowered saxifrage - 65
Tychius - 108
Tyrrheniberus - 83
Umbilicaria - 30
Vaccinum - 91
Val di Fassa saxifrage - 67
Valeriana elongata - 70
Valeriana saxatilis - 70
Valeriana supina - 70
Vandelli’s rock-jasmine - 43, 55
Vaud saxifrage - 49
Verrucaria - 30
Vicia cusnae - 68
Viola - 63
Viola argenteria - 63, 68
Viola calcarata - 142
Viola cenisia - 69
Viola comollia - 63, 69, 135
Viola magellensis - 71, 135
Viola valderia - 63, 68
Vipera ammodytes - 114
Vipera berus - 114
Viviparous lizard - 113, 114
Vole - 127
Wagneria alpina - 110
Wallcreeper - 118, 119
Wall-rue - 46
White alpine poppy - 66
Woodsia - 45, 47
Woodsia alpina - 47
Woodsia glabella ssp. pulchella 47
Woodsia ilvensis - 47
Woodwardia radicans - 34
Yarrow - 51
Yellow saxifrage - 27
Yellow thistle - 64
Yellow whitlow grass - 55
Zanoni’s murbeckiella - 64
Zerynthia polyxena - 137
Zonitis - 94
Zonitis flava - 94
Zonitis nana - 94
Zootoca vivipara - 113
The authors wish to thank:
Roberto Argano (isopods)
Maurizio Biondi (leaf beetles)
Alessandro Biscaccianti (longhorn beetles)
Marco Bodon and Folco Giusti (molluscs)
Marco Alberto Bologna (oil beetles)
Attilio Carapezza (bugs)
Giuseppe Maria Carpaneto and Emanuele
Piattella (scarabeoid beetles)
Enzo Colonnelli (snout beetles)
Alessio De Biase (shining flower beetles)
Massimiliano Di Giovanni (millipedes)
Simone Fattorini (darkling beetles)
Paolo Fontana, Bruno Massa and Fabio
Collepardo Coccia (orthopterans)
Renato Gerdol (scree vegetation)
Cesare Lasen (endemics of the Eastern Alps)
Gianfranco Liberti (soft-winged flower beetles)
Andrea Liberto (metallic wood-boring beetles
and click beetles)
Paolo Maltzeff (beetles)
Iuri Nascimbene (lichens)
Guido Pagliano (hymenopterans)
Graziano Rossi (Insubrian endemics)
Stefano Scalercio and Alberto Zilli (lepidopterans)
Augusto Vigna Taglianti (ground beetles and
earwigs)
Adriano Zanetti (rove beetles)
Marzio Zapparoli (centipedes)
Thanks are also due to
Maria Manuela Giovannelli, Erika Gozzi,
Paola Sergo and Maura Tavano
The authors assume full responsibility for any
errors or omissions in the text.
This volume was produced with funds from the
Italian Ministry of the Environment and Territory
Protection
Printed in June 2006
by Graphic linea print factory - Udine, Italy