Le guide di Dryades 1 – Serie Licheni I (L-I)
THE LICHENS OF ITALY
I. TERRICOLOUS SPECIES
Pier Luigi N
& Stefano M
The term “terricolous” 9
The terricolous lichen vegetation of Italy 12
Identification and classification 17
Program FRIDA 19
The distributional-ecological filter: ITALIC 22
The rank of characters 31
The keys 34
Tips and limitatons 35
GENERAL KEY 37
Subkey 1 - Fruticose lichens 37
Subkey 2 – Foliose lichens 83
Subkey 3 – Squamulose lichens 111
Subkey 4 – Crustose lichens 159
Subkey 5 – Leprose lichens 243
SIMPLIFIED KEYS 253
Key A: subalpine-alpine, on acid to subacid substrata 253
Fruticose lichens: subkey A1 253
Foliose lichens: subkey A2 259
Squamulose lichens: subkey A3 261
Crustose lichens: subkey A4 262
Key B: subalpine-alpine, on subneutral to basic (calcareous)
Fruticose lichens: subkey B1 269
Foliose lichens: subkey B2 271
Squamulose lichens: subkey B3 274
Crustose lichens: subkey B4 277
Key C: mediterranean to montane, on acid to subacid substrata 283
Fruticose lichens: subkey C1 283
Foliose lichens: subkey C2 287
Squamulose lichens: subkey C3 289
Crustose lichens: subkey C4 290
Key D: mediterranean to montane, on subneutral to basic
Fruticose lichens: subkey D1 295
Foliose lichens: subkey D2 297
Squamulose lichens: subkey D3 299
Crustose lichens: subkey D4 304
INDEX TO TAXA 333
Abstract: This book contains keys to all terricolous ascolichens hitherto
known from Italy (439 species). The introduction includes a critical
discussion of the term “terricolous”. The general key, provided with
descriptions and maps which show the hitherto known distribution in Italy, is
followed by four smaller keys for beginners, limited to species found on
acid-siliceous and calciferous substrata, below and above treeline. A
glossary of 300 technical terms is included. The basic outline of the keys
was produced by program FRIDA, patented by the University of Trieste.
Riassunto: chiavi per l'identificazione di tutti i licheni terricoli italiani (439
specie, escludendo i basidiolicheni). Introduzione: discussione del temine
“terricolo”. Chiave generale, con descrizioni e carte di distribuzione. Quattro
chiavi semplificate per i licheni di substrati silicei e calcarei, al di sotto ed al
di sopra della fascia montana. Glossario di 300 termini tecnici. Le chiavi
sono state generate da un programma originale (FRIDA), brevettato
dall'Università di Trieste.
Acknowledgements: we thank Dr. Guido Incerti (Trieste) for technical
assistance in the preparation of the maps, and Prof. Mauro Tretiach (Trieste)
for critical comments to the keys. The publication of this book was partially
funded by a MIUR grant to the senior author.
This book, dedicated to “terricolous” species, is the first in a
series of practical guides to the lichens of Italy, conceived as a
baseline for a forthcoming flora of the country. The outline of the keys
was produced by program FRIDA, written by the junior author (see
later), while data input and descriptions are by the senior author.
The instruments which led to the production of this book
originate from two projects co-financed by the Italian Ministry for
Education in the period 2000-2004, under the direction of the senior
author. The first project has created ecological-distributional and
nomenclatural databases searchable on-line for most of the terrestrial
“plants” of Italy. These are:
1) Macrobasidiomycetes http://dbiodbs.univ.trieste.it/global/mush
2) Lichens http://dbiodbs.univ.trieste.it/
3) Mosses http://dbiodbs.univ.trieste.it/global/mosses1
4) Liverworts http://dbiodbs.univ.trieste.it/global/epat1
5) Vascular plants http://dbiodbs.univ.trieste.it/global/flora1
The second project (2003-2004) is presently implementing
morphological databases and interactive identification keys. It aims at
making easier the identification of organisms by joining ecological-
distributional information and morpho-anatomical data into a program
for producing identification tools available on-line.
The term “terricolous”
The term “terricolous” is ambiguous. It well deserves to be
discussed in an introduction.
“Terricolous” derives from the Latin “terra”, which originally
referred to land as opposed to water and air (e.g. coelum et terra = the
sky and the earth, terra marique = the land and the seas, terrestrae
navalesquae pugnae = battles on land and sea, terram attingere =
getting to the harbour). Later, it was used for solid ground in general,
and even for what we now call “soil” (terrae solum = the soil; ea qua
gignuntur a terra = those things which arouse from soil; terram
subigere: working the soil; gleba terrae = a heap of soil).
The term “soil”, as well, derives from Latin. Solum is
something we can trample on by the sole (Italian = suola) of our feet
or shoes [nullius ante trita solo = a ground never trampled by anybody
(Cicero); sederunt medio terra fretumque solo = they were sitting on
bare ground (Ovidium); terrae solum = the real soil (Lucretius!)]
The Greek counterpart for terra is Γη. In earlier times Γη
was a primeval Goddess, the Mother Earth (the Goddess Tellus of
Romans). The term Γη evolved like terra: it was used for anything
which is different from sea and the atmosphere, for something solid
under our feet, sometimes for what we call “soil” (as early as Omerus,
we find the expression τεν γην εραξεσται = working the soil).
The Greek counterpart of solum is πεδον. Like solum,
πεδον has to do with the German-(English) concept of “ground”
(from the Barbarian root “grnd”), e.g., εσ πεδον χεο = I let it fall to
ground, Aeschilus. Πεδον was often used as the Latin solum in the
widest sense, e.g. Ευρωπησ πεδον = the territory of Europe.
(Aeschilus). The Latin term for feet is pedes (Italian: piedi). For
Greeks and Romans the “ground” (πεδον, solum) is something we
trample with feet.
Solum and πεδον are perhaps nearer to the modern concept of
“soil” (see later) than the almost mythological terms Terra and Γη.
However, the term “solophilous” was never applied to lichens, being a
bad mixture of Latin and Greek, while the more correct, completely
Greek term pedophilous is not good for lichens. The fully Greek term
“epigaeous” (sometimes spelled epigaeic) can be used instead of
“terricolous”, with the same meaning, i.e. "growing on the ground". In
this book we use the term “terricolous”, because is more familiar to
Italians (terricolo) than the Greek “epigaeous”.
Apart from etymological matters, most lichenologists agree
that “terricolous” (“epigaeous”) lichens are those growing on the
ground, or on “soil”. In modern science, however, the term “soil”
enjoys a plethora of widely different definitions by pedologists, the
systematics of soil being even more complex and confused than that
of organisms. Definitions such as "material in the top layer of the
surface of the earth in which plants can grow", or "a variable mixture
of organic and inorganic materials, which contains life and
constitutes a complex biochemical system of solids, liquids and air"
are of no great help for defining the guild of "terricolous" lichens.
Soils can be mainly organic, mainly mineral, or a mixture of
these two components. On the organic side, can a half-rotten, half-
living leaf of Saxifraga colonised by Caloplaca saxifragarum be
considered as a “soil”? The border between living-dead plant material
and a “soil” is a faint one: the degree of decomposition of organic
material in order for it to be considered as real “soil” is controversial.
However, a layer of recently fallen leaves in a forest is an integral
component of the soil profile according to most pedologists. On the
mineral side, things become even more confuse: lichens are cited in
many textbooks as the pioneers in the processes of pedogenesis. Thin
layers of strongly weathered rocks can act as a “soil” for several
lichens, but no phanerogamist would ever consider them as a “real
Like organisms, soils have had a long evolution on Earth.
According to Barreno (1998): “Terrestrial habitats in the Palaeozoic
consisted only of siliceous rocks and mineral soil: the first adaptive
radiation is likely to have occurred on these substrata…we should
distinguish between lichens of mineral siliceous soil and those
occurring on organic soil, the latter habitat certainly being much
younger….A relatively young genus, like Cladonia, finds its maximum
diversity on such substrata”. This sentence is worthy of consideration,
as a trigger for further research.
The present book deals with “terricolous” lichens. How do we
define this ecological guild?
Some “terricolous” lichens exclusively grow on predominantly
mineral soil (e.g. Solorinella asteriscus), others are found on living or
half-dead plant debris on the ground (e.g. Caloplaca tiroliensis), still
others only occur only on humic, organic soil (e.g. Cladonia
macroceras). In their checklist of Austrian lichens, Hafellner & Türk
(2001) were so brave as to distinguish between strictly “terricolous”
species (those growing only on “real soil”), and those which occur as
epiphytes on living or half-dead bryophytes, or on living plants found
on the ground.
In this book, however, the term “terricolous” has a very broad
sense, perhaps closer to the roots of the term “soil”: it refers to all
lichens found on the ground, irrespectively of whether they occur on
mineral or humous soil, strongly weathered rocks, dead bryophytes or
small, half-dead plants.
One could rightly wonder whether such a broad definition is
adequate for delimiting a clear-cut guild of ecologically different
Several lichens which usually dwell on rocks or bark
occasionally become “terricolous”. Such shifts in the main substrata
are most common in strongly oceanic or continental areas, e.g. along
the coasts of W Europe and in the steppes of C Asia. The senior
author has seen Ramalina farinacea and Lobaria pulmonaria growing
on soil in Portugal, and a hill near Lake Baikal whose “soil” was
covered by a lichen community dominated by Parmelia caperata and
other foliose lichens, very much resembling that which can be found
on isolated trees near Trieste. Several epiphytic or epilithic lichens
occasionally occur on “soil” in the most humid parts of the
Mediterranean, and in the driest valleys of the Alps. Not all of them
are included in these keys, the selection being restricted to the most
Italy, however, is the “temperate” country par excellence,
where most lichens are bound to a given type of substrate. Keys to the
“terricolous” species of Italy may have a practical sense.
The terricolous lichen vegetation of Italy
No study was ever published on terricolous lichen
communities in Italy. Northern and Central Europe - as the whole
circumboreal-subarctic zone - are better known, due to the prominent
role of terricolous lichens in these biomes (see e.g. Klement 1955,
James et al. 1977, Gilbert 1980, Daniëls 1982, Ahti & Oksanen 1990,
Paus 1997, Fryday 2001). The work of Kalb (1970), although
restricted to a narrow area, gives important insights on the terricolous
vegetation of the Alps. Southern Europe, on the contrary, is almost
unexplored, with relevant exceptions due to Spanish authors (e.g.
Crespo & Barreno 1978, Egea 1985, Burgaz & Ventureira 1990,
Barreno 1991), who described the terricolous vegetation of dry areas
of their Country. In other dry areas of the world, esp. America and
Australia, intensive research was devoted to “soil crusts”, often - not
always - without much detail about the species making up the so-
called “crusts” (for a review, see Rosentreter & Eldrige 2002).
At least in Europe, the scarce interest for “terricolous” lichen
communities could be due to methodological problems related to the
prevailing phytosociological approach (Nimis 1991). Terricolous
lichens grow together with vascular plants and bryophytes, and are
just one of the components of vegetation. Several authors wondered
whether it is correct to describe “terricolous lichen communities”
without considering the other organisms occurring together with
lichens (for a discussion see e.g. Barkman 1958, Nimis 1991).
However, it is difficult to put together three experts, a lichenologist, a
bryologist and a “botanist”, for studying a lichen-rich terricolous
vegetation. In the European phytosociological literature there is a huge
number of relevés produced by non-lichenologists. They mostly
contain records of species such as “Cladonia pyxydata, Cetraria
islandica, Toninia coeruleonigricans, Peltigera canina”, which tell
very little on lichen vegetation.
The best-known communities are those where lichens are so
prominent as to force serious phytosociologist to ask for assistance by
a lichenologists: the Toninion (the famous Bunte
Erdflechengemeinschaft), the carpets of Cladonia found under pine-
woods or among Rhododendron heaths in the Alps, the vegetation of
wind-swept ridges (Loiseleurio-Alectorietum).
As a result of this state of affairs, the senior author of this book
is not able to describe the “terricolous” lichen vegetation of Italy.
In order to provide a kind of “State of the Art” for the next
generation of Italian lichenologists, we present here, with a few
comments, the honest phytosociological scheme of Wirth (1995),
warning that it is based on a few papers only, that it is largely
incomplete, and that much interesting work is needed before to attain
a clear picture of the terricolous lichen vegetation of Italy.
Physcietea Tomaselli & De Micheli 1957
Xanthorion parietinae Oksner 1928 - Strongly nitrophytic, mostly
epiphytic communities of well-lit situations. Occasionally, at bird
perchings' sites several Xanthorion species may occur on the
ground, mostly on bryophytes, especially on calciferous subsrata.
Only a few of them are included in the present keys.
Cladonio-Lepidozietea Ječek & Vondr. 1962 – Communities of
lichens and bryophytes on rotting wood. Only a few species are
included in the present keys, the border between rotting wood and a
real “soil” being rather ambiguous.
Psoretea decipientis Mattick ex Follm. 1974 – Toninion sedifoliae
Hadač 1948 - Lichen-rich communities of calciferous substrata.
Toninio-Psoretum decipientis Stodieck 1937 –
Dominated by several species of Toninia, Fulgensia, Catapyrenium
s.lat., this vegetation of open calciferous soil, most frequent below the
alpine belt, includes different communities - several of which await
Cladonietum convolutae Kaiser 1926 – Dominated by
Cladonia convoluta, C. rangiformis, C. subrangiformis. Found in
slightly more sheltered and humid situations than the Toninio-
Psoretum. Frequent also in areas with sand dunes, mostly in intradunal
depressions, from the mediterranean to the submediterranean belts.
Cladonietum symphycarpae Doppelb. in Klem. 1955 –
Cladonia symphycarpia, C. polycarpoides, C. pocillum. A rather ill-
defined community of calciferous soil, somewhat intermediate
between the Cladonietum convolutae and the Toninion, with optimum
in the mediterranean-submediterranean belts.
Psoretum decipientis Frey 1922 (= Fulgensietum
alpinum Poelt) - The counterpart of the Toninio-Psoretum decipientis
above treeline, mostly on intermediate, weakly calciferous soil. A ill-
defined “community”, characterised by the presence of Fulgensia
bracteata and perhaps of Placidium lachneum. The vegetation with
Solorinella asteriscus still awaits description.
Endocarpetum pusilli Galle 1964 – A poorly defined
“community”, characterised by “Endocarpon pusillum” and perhaps
by Anaptychia bryorum, it colonises thin layers of soil on steeply
inclined faces of siliceous rocks with dripping of water in the
continental Alps (Kalb 1970).
Ceratodonto-Polytrichetea piliferi Mohan 1978 – Lichen-rich
vegetation of siliceous soils, often dominated by bryophytes.
Peltigeretalia Klem. 1950 – This “order” of the
phytosociological hierarchy is supposed to include all lichen-rich
communities on siliceous ground. After the monograph of Peltigera
by Vitikainen (1994) the name “Peltigeretalia” sounds like a garbage
can from old times. The many species of Peltigera have different,
nuanced, not always well-known ecological requirements. Many of
them could be used to distinguish among lichen-rich communities
which hardly can be subsumed under the same umbrella.
Baeomycion rosei Klem. 1955 – Lecideetum uliginosae
Langerf. ex Klem. 1955 - Pioneer communities of acid soil dominated
by crustose lichens.
Cladonion arbusculae Klem. 1950 – Subarctic-subalpine
communities of well-lit situations, occurring on acid soil, dominated
by species of Cladonia, such as C. arbuscula s.lat., C. crispata, C.
pleurota, C. rangiferina, C. uncialis.
Cladonietum mitis Krieger 1937 – This is the typical
“reindeer lichen community” - perhaps the only one which really
belongs to the Cladonion arbusculae - most frequent in the subalpine-
alpine belts, both in tundra-like vegetation and in the understory of
very open coniferous woodlands (esp. in the most continental parts of
the Alps, where it can descend to ca. 1000 m). It is not uncommon
throughout the Alps, and it occurs in fragmentary and impoverished
forms also in the Gran Sasso and Majella Massives (C-Apennines).
Cladonietum foliaceae Klem. 1955 – This
“community”, dominated by Cladonia foliacea and certainly not
related with the Cladonion arbusculae, is the silicicolous counterpart
of the Cladonietum convolutae. In Italy, Cladonia foliacea occurs
from the mediterranean belt to near treeline, and is likely to take part
into several, well-distinct communities which, however, still await
Cladonietum destrictae Krieger 1937 - This ill-defined
community is characterised by the dominance of Cetraria aculeata, a
species which, in Italy, is broad-ranging, and which is likely to be an
element of different vegetation types.
Cetrarion nivalis Klem. 1955 – Arctic-alpine communities of
wind-swept ridges above treeline, dominated by Flavocetraria
cucullata, F. nivalis and Thamnolia vermicularis s.lat.
Empetro-Cladonietum stellaris Du Rietz 1925 – A
well-defined, mainly subarctic-subalpine community of mostly
organic, acid soil in well-lit but wind-protected, rather humid
situations, characterised by the dominance of Cladonia stellaris, and
by the higher frequency of C. amaurocraea. This community - which
has little to do with the Cetrarion nivalis and is more related to the
Cladonietum mitis - needs a long ecological continuity. Hence, it is
very rare, and strictly limited to above treeline in the Alps of Italy.
Loiseleurio-Alectorietum ochroleucae Du Rietz 1925 –
The typical community of wind-swept ridges (short snow-lie) above
treeline, best developed on siliceous substrata but also occurring on
calciferous ground. It is dominated by Alectoria nigricans, A.
ochroleuca, Flavocetraria cucullata, F. nivalis, Thamnolia
vermicularis s.lat. Common throughout the Alps, it also occurs in an
impoverished form in the Gran Sasso and Majella Massives (C-
Solorinion croceae Klem. 1955 – Arctic-alpine to
subarctic-subalpine communities of siliceous, more or less acid soil,
near or above treeline. Best developed and mostly restricted to the
Alps in Italy.
Lecidomatetum demissae Frey 1923 – Characterised by
the dominance of Lecidoma demissum, this “community” of siliceous
substrata above treeline is actually dominated by higher plants: the
lichens (among them Cetraria islandica, Cladonia coccifera,
Protopannaria pezizoides, Solorina crocea, Stereocaulon alpinum,
etc.) colonise the bare soil amongst tufts of grasses. The soil is mostly
clay, with an organic component, hence rather humid. Restricted to
the Alps in Italy, with some species reaching south to the mountains
Lecideetum limosae Klem. 1955 – A pioneer
community of (mainly) crustose lichens, occurring on naked soil in
sites with a long snow-lie, e.g. in clearings of Alpine grasslands.
Stereocauletum alpini Frey 1937 – Characterised by the
dominance of Solorina crocea, Stereocaulon alpinum and Cladonia
macrophyllodes, this community very much resembles the
Lecidomatetum demissae, but tends to colonise mineral acid soil in
slightly drier, more illuminated situations. In Italy it is restricted to
above treeline in the Alps.
Caloplacetum nivalis Kalb 1970 – Characterised by
Arthrorhaphis alpina, Bryodina rhypariza, Bryonora castanea,
Caloplaca nivalis, this community colonises pockets of mostly
organic soil in depressions with a long snow-lie on siliceous substrata.
It is restricted to the Alps in Italy.
Megasporion verrucosae Kalb 1970 – Arctic-Alpine to
subarctic-subalpine communities of living or moribund bryophytes
and plant debris on calciferous substrata (Megaspora verrucosa,
Mycobilimbia hypnorum, Pertusaria geminipara, etc.).
Megasporetum verrucosae Frey 1927 – A rather ill-
defined, arctic-alpine community of more or less calciferous soil,
found in rather sheltered situations with a long snow-lie. It is
apparently dominated by Gyalecta foveolaris, Rinodina mniaraea, R.
turfacea, Solorina bispora, Vulpicida tubulosus, with several species
of the following community.
Caloplacetum tiroliensis Kalb 1970 – Dominated by
species such as Caloplaca ammiospila, C. cerina v. chloroleuca, C.
saxifragarum, C. tiroliensis, Lecanora hagenii v. fallax, Lecidella
wulfenii, Peltigera lepidophora, Phaeorrhiza nimbosa, this vegetation
is fairly common above treeline, on calciferous substrata, mostly on
plant debris and bryophytes in rather exposed, well-lit situations.
Widespread throughout the Alps, well-developed also in the Gran
Sasso and Majella Massives (C-Apennines), it occurs also in the
highest calcareous peaks of the S Apennines (e.g. the Pollino Massif).
The senior author of this book is not a supporter of the so-
called “syntaxonomical” approach of phytosociologists. The existing
phytosociological schemes are based on old species concepts, and
often on a poor knowledge of species. Some examples: the recently-
revised genera Catapyrenium s.lat. (Breuss 1990) and Placidiopsis
(Breuss 1996) include many species which occur in the so-called
Toninion; in the phytosociological literature they were mostly
subsumed under the epithet “Dermatocarpon trapeziforme”. Before
the monograph of Toninia by Timdal (1991) most terricolous species
were called “Toninia coeruleonigricans”. Even for Peltigera
information is scanty: before the monograph by Vitikainen (1994)
most phytosociologists recognised a few species only (e.g. P.
aphthosa, P. canina, P. rufescens). The genus Fulgensia, one of the
most characteristic elements of the Toninion, like Endocarpon and
many other groups, is still in need of a thorough revision.
Much work is needed to appreciate the real role of lichens in
the “terricolous” vegetation of Italy.
Three suggestions from the senior author to the next generation
of Italian lichenologists: 1) be accurate in identifying the species (this
book might be of some help), 2) avoid the phytosociological
“orapronobis”, 3) use a rigorous sampling design in your studies.
Identification and classification
After Gutenberg, information useful for identifying organisms
was printed on paper, as in any classical Flora, and in the present
book. The constraints of a paper-printed text have forced most authors
to organise information according to the hierarchical scheme of
Classical keys first lead to families, then to genera, and finally
- if everything works - to species. Supraspecific taxa often need
“difficult” characters. Many of the easy-to-look-at characters, such as
the colour of the thallus, and those referring to ecology and
distribution, are alien to the hierarchical-taxonomic scheme. There is
an abysmal difference between the amount of information requested
by a classical flora, and that which would be enough for giving a name
to an organism. Odd options may be encountered, such as the
distinction between two species that never had the pleasure to meet
with each other, having completely different ecology and distribution:
that between two lichens, one with a yellow, the other with a black
thallus, using the character: “ascus Porpidia-type”. The old
hierarchical-taxonomic scheme still maintains a prominent position
for projects aiming at producing a “global” key for any group of
organisms. Until now - fortunately - no such keys were produced for
lichens (see later).
Classification and identification - albeit related - belong to two
different operational processes (Bridgman 1927). Classification is the
job of taxonomists, identification can be fun for anybody. The great
American lichenologist Mason Hale was aware of this fact when he
produced the classical “booklet” How to know the lichens (Hale1969,
Nowadays, information can fly on wings much stronger and
elastic than sheets of printed paper. Computer-based programs can
utilise, in a multi-dimensional way, a wealth of morphological-
anatomical data, plus the distributional-ecological information usually
hidden in the large ocean of scientific literature. A revolution!
Traditional floras had several drawbacks:
1) Being printed on paper, their content is "frozen". Nomenclatural-
taxonomic changes, progress in floristic exploration, the discovery of
new species, often render a flora outdated within a few decades.
Computerised systems, on the contrary, can be updated and corrected
in real time.
2) The larger a flora (or a taxon) is, the more difficult it is for the user
to identify an organism. Computerised tools permit to reduce the set of
organisms using different combinations of morphological, ecological,
3) Traditional keys are "rigid". They contain a huge amount of
information which is frozen into the format and the logical structure
selected by the author. Computerised floras, being “elastic”, can
generate products which would have required a huge amount of work
in the past. Some examples: a) regional-local floras (e.g. of a biotope,
a natural park, a province); b) keys for "virtual habitats", by
combining distributional data with ecological indicator values (see
Nimis & Martellos 2001), c) keys for special users.
In Italy, lichens are widely used in educational projects at all
levels, from the ground school to the university. FRIDA (see later) not
only permits to construct keys for the area in which the school is
located, but also to adapt the language to the educational level of
schoolchildren (see Nimis et al. 2003). Some characters can be
automatically downscored in the keys (e.g. the use of
Paraphenilendiamine for colour tests, which, being cancerogenic, is
not appropriate for children).
4) Databases are “accumulative”. A small database (e.g. limited to a
taxonomic group, or to a local flora) can be the starting point for
future expansions. For example, the production of a key of terricolous
lichens known from Slovenia has required the addition of 17 species
only to the Italian database.
5) Outputs can be edited in several different formats, from simple
texts to illustrated books (see Nimis et al., 2003). Any user can now
produce her/his "personalised books", à la carte, on-line. A warning
for the “global” publishing industry?
The present book is still printed on paper. One could wonder
why at the same time - it is made freely available in the internet
Databases are ephemeral: an economic crisis, the change of
policy in a Department, the sudden demise of The Professor, could
prevent the purchase of new servers, the payment of a Webmaster, any
On-line databases can disappear in the matter of a few hours.
Paper-printed information, while bulky and rigid, still may
enjoy a longer, safer life-span.
One of the first and best known packages for interactive
identification is IntKey, based on the DELTA format, developed in the
CSIRO Division of Entomology (Australia) starting from as early as
1971 (http://www.biodiversity.uno.edu/delta/). Its facilities include the
generation of descriptions and conventional keys, the conversion of
data for use by classification programs, and packages for interactive
identification and information retrieval. The system is capable of
producing high-quality printed descriptions. IntKey generates
conventional identification keys: in selecting characters; at any step,
the program determines how well they divide the remaining taxa, and
balances this information against subjectively determined weights
which specify the ease of use and reliability of the characters. The
system is in use worldwide for a wide array of organisms, including
viruses, corals, crustaceans, insects, fish, fungi, plants, and wood.
IntKey was tested in Trieste in 2000 on some subsets of
lichens. A trial was also made on the genera of Graminaceae occurring
in NE Italy (Ganis et al. 2000, and http://www.univ.trieste.it/
~biologia/gram/ indxgram.htm). The judgement of the package was
positive, but several negative aspects were also pointed out.
For lichens, IntKey was adopted by LIAS (A Global System for
Lichenized and Non-Lichenized Ascomycetes), an ambitious, well-
established, multi-authored information system for the collection and
distribution of descriptive and other biodiversity data on lichens and
non-lichenised ascomycetes coordinated by the Botanische
Staatssammlung München (Rambold, 1996–2004).
One could wonder why, instead of joining LIAS, we have
produced our keys using a different program. The main reason is the
critical attitude of the senior author toward “global” projects.
The Utopia of a unique, “global” system which could permit to
“give a name” to any organism collected on Earth is justified by the
progress of informatics, but not by the state-of-art of knowledge on
biodiversity. Such a Utopia was conceived for a World without
problems. The present World, however, is full of problems. Two
examples: 1) Does Helocarpon crassipes occur in the Alps?
According to some checklist, yes. Other authors maintain that the
populations of the Alps differ from those of N Europe; further study is
needed. 2) Is the continental Dirina stenhammari a “good” species or
just a morph of Dirina massiliensis f. sorediata? Tehler (1983) is for
the second option, other authors (e.g. Hafellner, in litt., and the senior
author) still have doubts. DNA data could perhaps solve the problem.
Contrasting opinions by different authors are powerful triggers
of research: they point to problems. Global systems are full - as the
present book, or any monograph, checklist, and flora - of hasty
synonymisations, wrong species concepts, and wrong records.
Nothing new, no problem. But a disaster if such systems will become
the “only” way to access biodiversity information worldwide. If the
next, probably small, generation of lichenologists will be fed on clicks
into “The Global System” to get inspiration for research.
A “global” repository of taxonomic-morphological information
is certainly possible, useful, and desirable. However, if this were the
only one on Earth (can we imagine more than one “global” system?),
it could turn into a nightmare: “The McDonaldization of Taxonomy”,
with swards of young lichenologists from Italy to Singapore eating the
same taxonomical BigMac. “Global” projects must live in the real
World. The real World is diverse, full of different systems, ideas,
taxonomic concepts, sometimes in agreement with each other,
sometimes not, as in the old “Floras”.
We did not join LIAS because we wanted to give a modest
contribution to the diversity of ideas and approaches in the “global”
In 2000, the junior author proposed to write a completely new
program. Work developed in a continuous, intense, often conflictual
interaction between the two authors. This was a good exercise for both
of them, which contributed in making the software digestable to
colleagues who know well the plants, but are not acquainted with
The result is program FRIDA, which attracted the interest of
AREA Science Park Trieste in the framework of project SISTER, that
financed the promotion of its products, and its patenting by the
University of Trieste
Procedures and functions of FRIDA are written in PL/SQL
language, running on a Oracle Database Server, which is connected to
the Web by an Oracle server. FRIDA is flexible, its use does not
require learning any special language nor using codes to input
information, and is able to export data in several other formats.
Partial on-line access to keys produced by FRIDA is available
since May 2003 through ITALIC (http:/dbiodbs.univ.trieste.it), limited
to terricolous lichens (Nimis & Martellos 2003). An interactive key of
vascular plants of the Trieste Karst, produced by FRIDA, is available
The keys printed in this book and those available on-line look
similar, but are basically different. In the on-line version, the user can
specify any set of ecological, distributional, morphological or
chemical characters he likes. These characters act as “filters” reducing
the set of species included in the key. For example, if one has found
Caloplaca cerina, a crustose lichen with a grey thallus and orange
apothecia reacting K+ red on a tree near Trieste, one can limit the
search to “crustose”, “thallus neither dark nor yellow-orange”,
“apothecia yellow to orange”, “on trees”, “Trieste”. The result is a key
to 6 species, without questions such as “Photobiont trentepohlioid?”,
“Spores polar-diblastic?”, “Cortex paraplechtenchymatous?”.
The users of paper-printed keys, such as those of this book,
cannot enjoy such privileges. However, informatics is developing
rapidly: multi-dimensional identification tools will be soon available
The distributional-ecological filter: ITALIC
The present book also includes four simplified keys for lichens
of calcareous and siliceous substrata, above and below the montane
belt. They were produced by joining FRIDA with information stored
into ITALIC, the information system on Italian lichens (Nimis &
ITALIC was created in 1999, when the junior author proposed
to the senior one to transform his bulky catalogue of Italian lichens
(Nimis 1993) into a database searchable on-line. Since then, ITALIC is
being developed as an organiser of several databases dealing with the
lichens of Italy, such as herbaria, regional checklists, archives of
pictures and maps, and presently FRIDA.
The first version of ITALIC was searchable in the internet since
October 2000, the second version was published on-line in 2001
(Nimis & Martellos 2002), a third version appeared in May 2003. In
2003, a thesaurus of more than 12.000 infrageneric epiteths was
incorporated into the system, which permits connection with other
databases following different nomenclatural standards (Nimis &
Lichens are broad-ranging; many species present in Italy also
occur in other European countries, several of them extend to the whole
holarctic region, sometimes to the Southern Hemisphere. Information
from ITALIC could be useful outside the narrow borders of Italy.
A description of ITALIC can be found in Nimis & Martellos
(2002). Here we only mention the data used for producing the
Geographic subdivisions – Maps (see later) and simplified keys are
based on two main subdivision of the Country:
1) Administrative regions. 21 Operational Geographic Units (OGUs,
see Fig. 1), the small Province of Trieste being separated from Friuli
due to its peculiar biogeographical features. For each species, ITALIC
gives references for each region, limited to papers published after
2) Phytoclimatic areas. 9 OGUs, delimited on the basis of several
GIS-maps (altitude, precipitation, etc.), also taking into account the
climatic difference between the Tyrrhenian (humid) and Adriatic (dry)
parts of the peninsula (Nimis & Tretiach 2004). Commonness-rarity
values (see later) were calculated for each taxon in each of the 9 areas.
A0) Nival area: well above treeline, in areas occupied by glaciers,
near the tops of the highest mountains (Alps only).
A) Alpine area (above treeline in the Alps and in the highest peaks
of the Apennines, Gran Sasso and Majella Massives in Abruzzi).
An area devoid of trees, where terricolous lichens give rise to a
diversified palette of communities, depending on the main
substrate, and on microhabitats (from fissures of the rocks to
organic soil in snow-beds). The so-called “arctic-alpine” element is
most common on acid siliceous substrata, whereas many species of
calciferous ground extend to the high mountains of Asia (Nimis
A1) Subalpine-oroboreal area (near treeline in the Alps, dominated
by Larix-Rhododendron, also including semi-natural, closed Picea
abies forests). Several so-called “boreal-montane” lichens form
distinct communities, such as those dominated by species of
Cladonia. In Italy, the limit of most arctic-boreal vascular plants
lies somewhere in the N Apennines, with the relevant exception of
the Gran Sasso-Majella Massives in the Central Apennines. As far
as lichens are concerned, however, the mountains of Calabria and
even those of Sicily do still host several so-called “boreal” lichens.
B) Oromediterranean area (above treeline outside the Alps and
Abruzzi). In southern and insular Italy only a few mountains attain
treeline. Some of them (e.g. the recent Etna Vulcano in Sicilia, and
the much older Gennargentu Massif in Sardegna) host a peculiar
vegetation, dominated by thorny-shrubs of the genus Astragalus,
Tragagantha-section. The thorny-shrubs formations of the
mediterranean mountains have an old history, perhaps dating back
to the Messinian period, when the Mediterranean was a semi-
desert, biogeographically connected with the Iranian-Turanian
region (Pignatti et al. 1997, Nimis 1981). The lichen flora of the
few truly Oromediterranean peaks of Italy still awaits exploration.
C) Montane area (beech forests). Beech (Fagus sylvatica) is the
dominant tree in the mountains of Italy. In the Alps it forms pure to
mixed forests (with Abies alba in cool-humid situations), with a
broad altitudinal range, from ca. 600 m, in contact with the
submediterranean belt, to ca. 1800 m, in contact with the oroboreal
belt. Along the Apennines, down to Sicily, albeit twarted and
shrub-like, beech marks treeline, but it does not occur is Sardegna.
During the glacial periods, Fagus-forests and their flora were
confined to refuges in southern Europe, esp. in the Balkan and
Italian peninsulas, and later expanded northwards, the vascular
flora of beech forests becoming poorer from southern Europe to S-
Scandinavia (Nimis & Bolognini 1993). Beech forests are the
“national” forest of many C-European writers and poets, but those
of Italy and the Balkan Peninsula are the “original” ones, those
with the highest diversity. The beech forests of Italy, however, are
still a mystery for lichenologists: some of them (e.g. those in the W
part of Calabria), host an unusually rich, interesting, luxuriant
lichen vegetation (e.g. with mediterranean-montane species such as
Melanelia laciniatula, Physconia venusta and Ochrolechia
balcanica), others - even those located in rainy areas (e.g. parts of
Liguria and Friuli) - are almost a lichen desert. The balance
between air humidity and precipitation in the liquid form can
perhaps explain the dramatic differences. Commonness-rarity
values (see later) for beech forests wrongly consider the beech-belt
as a continuum from the Alps to the mountains of Sicily. A clever
exploration of the lichens in Italian beech-forests could give rise to
interesting results. Interestingly, no terricolous lichen of Italy has
an optimum on mineral soil in this belt, but the distribution of
many species of Peltigera still awaits further study.
D) Submediterranean area (deciduous oaks, excluding E and F):
the submediterranean area, lying between the montane and the
mediterranean belts, covers most of the lowlands and hills of Italy.
The potential vegetation is dominated by deciduous trees, esp.
Quercus and Carpinus, most forests having being substituted by
coppices dominated by Ostrya and Fraxinus ornus, urban areas and
cultivations. The glacial and post-glacial history of
submediterranean forests is similar to that of beech forests, with a
difference related to the thermic requirements of the dominant
trees, which survived in warmer sites, mostly in lowland areas, esp.
in S Italy. The vascular flora is richer in the south, poorer in the
north (see Bolognini & Nimis 1993), while lichen richness mainly
depends on air humidity. In ITALIC, the submediterranean area is
split into three different OGUs: 1) Padanian (the Po-plain, mostly
occupied by urban areas or intensive cultivations, almost devoid of
forests), 2) Tyrrhenian (exposed to humid air masses), and, 3) the
present OGU, which includes what is left excluding areas nr. 1 and
2. This area includes the hills and the alpine valleys of N Italy, and
the eastern part of the Peninsula (see map in ITALIC).
E) Padanian area (the plains of the North, plus a narrow strip
along the eastern side of peninsular Italy): this is the most heavily
anthropised part of Italy, where several species do not occur
because of pollution and/or almost total deforestation. Before
Roman colonisation, most of the Po-plain was covered by dense
deciduous forests. Presently, it is an agricultural, industrial or
densely urbanised area. This OGU has no biogeographic meaning
(it formally belongs to the previous OGU): it was established only
because of its peculiar features, which are not favourable to lichens.
F) Humid submediterranean area (Tyrrhenian), (as D, but
restricted to areas with a warm-humid climate: see Nimis &
Tretiach 2004). The western part of the Italian Peninsula has a
mild-humid climate generated by Tyrrhenian maritime air masses,
while the Adriatic coast, on the lee-side of the Apennines, is
subject to cold-dry winds from Eurasia during winter. Tyrrhenian
Italy is characterised by several suboceanic species, some of which
have clear tropical-subtropical affinities. The inland extent of
maritime influence differs according to the presence-absence of
high mountains stretching parallel to the coasts, being more
pronounced in Tuscany and Latium, where the humid Tyrrhenian
air masses can reach the watershed of the Apennines. The
submediterranean part of Tyrrhenian Italy was delimited taking the
arbitrary threshold of 100 m as the limit between mediterranean
and submediterranean vegetation (like in the map of potential
vegetation by Tomaselli et al. 1973), and considering the main
geomorphological features. For further details see Nimis &
G) Humid Mediterranean area (Tyrrhenian): see previous point.
H) Dry Mediterranean area. Contrary to the Iberian Peninsula - a
small continent in itself - the narrow Italian Peninsula and its
islands, bathed on all sides by the Mediterranean, rarely experience
extreme climates. There are, however, a few parts of Italy which
have a really dry-mediterranean climate. These are mostly located
in S Sicily, SE Sardegna, and in parts of Puglia. Such areas are
important for terricolous lichens: they host several species also
occurring in N-Africa, and sometimes in the steppes and semi-
deserts of C-Asia (Barreno 1991). After Nimis & Poelt (1987), no
thorough exploration of dry-mediterranean Italy was undertaken: in
our opinion, an exploration of this area is likely to produce
Fig. 1 – The geographic subdivisions of Italy adopted in this book: Abr: Abruzzi,
Bas: Basilicata, Cal: Calabria, Camp: Campania, Emil: Emilia-Romagna, Frl:
Friuli, Laz: Lazio, Lig: Liguria, Lomb: Lombardia, Mar: Marche, Mol: Molise,
Piem: Piemonte, Sar: Sardegna, Si: Sicilia, TAA: Trentino-AltoAdige, Tosc:
Toscana, Umbr: Umbria, VA: Val d’Aosta, Ven: Veneto, VG: Venezia Giulia.
a b c
d e f
Fig. 2 – The main bioclimatic
subdivisions of Italy adopted in this
book: a: alpine, subalpine and
oromediterranean, b: montane, c:
submediterranean, d: padanian, e:
humid submediterranean (Tyrrhenian),
f: humid mediterranean (Tyrrhenian),
g: dry mediterranean.
Altitudinal distribution - This parameter refers to potential
vegetation only, on a 6-class ordinal scale:
1) eu-Mediterranean belt (potential vegetation: evergreen
Quercus ilex forest),
2) submediterranean belt (deciduous Quercus-Carpinus
3) montane belt (Fagus forests),
4) Oroboreal belt of the Alps (natural Picea abies, and Larix-
Pinus cembra stands),
5) Alpine and oromediterranean (above treeline in the
The island of Sardegna is a special case: Fagus is missing, and
the few remains of the “montane” belt, largely destroyed by cattle-
grazing, consists of scattered forests dominated by Ilex and Taxus. In
this book, the altitudinal belts of Sardegna were adjusted as to fit those
of Arrigoni (1968), but the "submediterranean" belt may be dominated
by evergreen forests, the "montane" belt by deciduous oaks, or by
remnants of ancient woodlands. On the whole, the island of Sardegna
does not fit in the phytoclimatic scheme adopted in this book.
Commonness-rarity - The related concepts of “commonness” and
“rarity” are difficult to define, and hence intrinsically fuzzy ones. A
given species might be fairly “common” in a narrow OGU, while it
may be extremely rare when the OGU is made broader. For example,
Cetraria islandica is fairly common in the Alps, rare along the
Apennines, extremely rare in the mountains of Sicily, and certainly
absent in the Po-Plain. There is obviously no sense in specifying its
“commonness” nationwide. In ITALIC, commonness-rarity - as a first
approximation - was calculated separately for each of the 9
phytoclimatic areas, on the basis of three main criteria: a) number of
samples in the TSB lichen herbarium (% on the total for each
phytoclimatic area), b) number of citations in the literature (taken
from ITALIC), c) an expert judgement used in particular cases (e.g.
that of recently-described taxa). A detailed explanation of the
methodology will be presented in a forthcoming paper. Commonness-
rarity was expressed on a 9-class scale, as follows: a: absent, er:
extremely rare, vr: very rare, r: rare, rr: rather rare, rc: rather
common, c: common, vc: very common, ec: extremely common. The
"er" class was adopted for lichens which are known from less than
five stations, and/or were not found in recent times, excluding most
recently-described species, and taxonomically very poorly known
taxa. This parameter was used to produce the distribution maps (see
Maps - Distributional maps appear at the left side of the description of
each taxon. They show - with different shadings - the probability of
encountering a given species in a given part of the country. The maps
are automatically produced by crossing information from ITALIC with
a GIS-based map subdividing Italy into phytoclimatic areas,
administrative regions, and altitudinal belts. ITALIC provides
information on: 1) commonness/rarity in the 9 phytoclimatical areas,
2) Presence/absence in the administrative regions, and in the
altitudinal belts. The overlapping of the 9 phytoclimatic areas and of 7
altitudinal belts upon the regional borders (the nival belt being merged
with the alpine-oromediterranean belts) produces a map
(downloadable from ITALIC) with 97 OGUs, each of which has
different shadings, related to commonness-rarity, with a single
constraint: if a species was never reported from a region, all of its area
remains white. These maps are likely to change at every updating of
ITALIC , e.g. when a species is found as new for a region. Their
ephemeral nature, however, can be beneficial for lichenologists: it
gives an added value to good floristic papers which rarely find a space
in high-impact journals. Any record of a species found as new to a
region immediately modifies its map, contributing to a better image of
its country-wide distribution. Due to their small size, the maps
included in this book utilise only 4 commonness-rarity classes: 1)
extremely to very rare, 2) rare to rather rare, 3) rather common to
common, 4) very to extremely common. More detailed maps are
available on-line through ITALIC.
Indicator values - Ecological Indicator Values (German:
Zeigerwerte) are expert assessments which express the ecological
range of organisms with respect to different ecological factors. They
are, at least in Europe, one of the most frequently used tools for
summarising the complex body of knowledge on the ecology of
organisms. Ecologists are rarely able to produce predictive models.
However, we have now a huge amount of information on the
ecological requirements of organisms, mostly hidden in many
scattered publications. For vascular plants, the best-known indicator
values are those proposed by Landolt (1977) for the Swiss flora, and
by Ellenberg (1974, 1991) for that of Central Europe. Indicator values
are presently available for several other organisms, e.g. diatoms (Van
Dam et al., 1994), bryophytes (Düll 1991) and lichens (Wirth 1991,
1995: Central Europe, Nimis 1999b: Italy). In this book, indicator
values (taken from the latest version of ITALIC, Nimis & Martellos
2003) were used for producing the simplified keys. They specify, for
each factor and for each species, a range on a 5-class ordinal scale.
Only one value, that relative to pH was used, to separate lichens of
siliceous and calcareous substrata. The indicator values for pH are as
follows: 1) on very acid substrata, such as very acid organic soil,
rotting wood, mineral soil deriving from acid siliceous rocks, 2) on
acid substrata (a buffer between 1 and 3), 3) on subacid to subneutral
substrata, such as soils deriving from base-rich siliceous rocks
(weakly calciferous sandstone, basic volcanic rocks, basalt, some
types of brick and tiles, etc.) 4) on slightly basic substrata, such as
soils deriving from calciferous sandstone, 5) on basic substrata, e.g.
more or less mineral soils deriving from pure limestone.
Notes - The notes which appear at the end of descriptions are basically
taken from the 3
version of ITALIC (Nimis 2003), with several
modifications, which will be incorporated in the next version. In order
to automatically produce notes in paper-printed form, those of ITALIC
were split into two distinct fields: a) information of general interest,
which can live independently from that provided by the system (e.g.
ecology and general distribution), b) information which is bound to
literature records of no general interest, or which cannot live
independently from the system (e.g. “the record from Piedmont by XY
is dubious, because...etc”, “see also note to species X”, etc.). Only the
former were included in this book. The notes of ITALIC, however,
were written in a very short format, excluding information provided
elsewhere in the system, e.g. in the ecological indicator values, the
altitudinal distribution, etc. When isolated from ITALIC, many of
them may appear as poor and dry. For this reason, several notes were
re-worked for this book, to include in a textual form at least the most
important information provided by ITALIC.
Pictures - We originally thought to complement this book with a CD-
Rom containing an illustrated version of the general key. Our system
can produce illustrated keys in a few minutes, thanks to a program
written by Dr. G. Incerti (Trieste) which automatically connects taxon
name, its description, the keys produced by FRIDA, and the archives
of ITALIC with distributional maps and pictures. The CD-Rom,
however, was not included, because of different reasons, some of
which have to do with complicate regulations concerning its
distribution together with a paper-printed book. Illustrated keys for
any subset of species specified by the user, can be purchased from the
Department of Biology of the University of Trieste. Photographs of
most taxa are freely available on-line through ITALIC (at the moment,
ca. 4000 images of ca. 2000 species). They show the general habitus
of the lichen, and mostly refer to samples preserved in the TSB
herbarium (with a few samples from GZU). These pictures have no
“aesthetic-artistic” pretension, having been mostly made by the senior
author with the idea of rapidly providing users with a first, rough set
of images. With all of its faults, the iconographical archive of ITALIC
contains the only extant image of many taxa. Input from colleagues
about “wrong” pictures - those referring to a different taxon - is most
welcome. Work on images is progressing rapidly: many more will be
added after the publication of this book, incl. those of anatomical
The rank of characters
As with most programs for interactive identification, the keys
produced by FRIDA are based on a hierarchy of characters, taxa being
separated on the basis of those which come first in the hierarchy.
In the present keys, characters are ranked according to the price of
instruments which are necessary to appreciate them, beginners - and
people from less wealthy countries as Italy - coming first: a) bare
eyes, b) a lens helping the eyes, c) a few easy-to-get chemicals, d) a
cheap microscope (such as those for children), e) a UV-lamp, f) a
professional microscope, g) access to chromatography, scanning
electron microscopy, etc.
Bare eyes - Eyes are extremely sophisticated instruments. They
can reveal whether our lichen looks as a flat crust or a small shrub,
whether it is dark-coloured or bright-yellow, whether it drastically
changes of colour after the application of a chemical, etc.
The lens - Most lichens are small, and many of them look alike
under the eye. If they were several meters large, they would look
much different. A cheap lens can help the eye to appreciate such
differences. With this help one can decide whether pseudocyphellae
(see glossary) are linear or punctiform, whether the lichen has soredia
or isidia (see glossary), whether the latter are flattened or cylindrical
in section, etc.
Chemicals – Lichens produce a wide palette of complex,
sometimes unique substances, whose importance for identification is
often fundamental. Some of these substances change the colour of the
thallus or of the medulla under the effect of some reagents. The
colour-tests are often reliable enough to distinguish among species. A
table of the main reactions caused by the main lichen substances can
be found in Wirth (1995).
A cheap microscope - Several lichens look alike both under the
eye and the lens, they have the same chemical reactions, but they are
completely different if seen - in section - under a cheap microscope.
For example, Lecidella elaeochroma and Arthrosporum accline look
almost identical under a lens: they both have a grey, thin crustose
thallus, and black apothecia with a thin black margin. However, a
section of the apothecia under a cheap microscope reveals
fundamental differences: Lecidella has one-celled, oval spores,
Arthrosporum has kidney-shaped, four-celled spores.
The UV-lamp. - A UV-lamp may be useful to identify some
lichens, because the colour of a thallus or its medulla under UV-rays
can tell a lot about their chemistry. UV-lamps are rather cheap, and
not difficult to get..but be careful about your eyes!
The professional microscope - Many lichens look similar under
the eye, the lens, the chemicals, the cheap microscope, and the UV-
lamp...but they look different when observed under a professional
microscope. This instrument can reveal such things as the exact size of
the spores, the presence-absence of warts on their walls, the form of
the asci and of their apical structures, the presence, size and shape of
crystals in the hymenium seen under polarised light, etc. We have
tried to give a low profile to such characters in these keys, but they are
often indispensable for giving a correct name to many lichens.
Professional microscopes are expensive. If you cannot afford to buy
one, you can contact the nearest Museum of Natural History, the
Departments of Biology of the nearest University, perhaps the
Association of Amateurs in your village: they are likely to let you use
their precious instruments.
Chromatography - Chemicals and UV lamps are helpful for
understanding the chemistry of lichens. There are, however, several
cases in which chromatography is the only means to identify
substances which are fundamental for distinguishing among species.
Thin-layer chromatography (TLC), or - even worst - gas-
chromatography, are neither easy to get nor to use, although we know
several dedicated beginners which were able to set up a small TLC-lab
in their private houses. The main chemicals produced by a species,
while often mentioned in the descriptions, have a low profile in the
keys, unless they are the only means for distinguishing among species.
The order of the main characters adopted in the keys is as
1) Growth form: fruticose, foliose, squamulose, crustose, leprose
2) General colour of thallus: dark, bright yellow to red, of any other
colour (bare eyes).
3) Photobiont (cyanobacterial, chlorococcoid, trentepohlial) (bare
eyes, sometimes helped by a lens or a cheap microscope).
4) Macroscopic characters such as the presence of pseudocyphellae,
cilia, rhizines, whether a fruticose thallus is filamentous or not,
whether the lower surface of foliose lichens is dark or pale, etc.
(bare eyes, sometimes helped by a lens).
5) Presence/absence of vegetative propagules, like isidia and soredia,
and their general features (bare eyes, sometimes helped by a lens).
6) Type of ascomata: apothecia (lecanorine, non-lecanorine) or
perithecia (bare eyes, sometimes helped by a lens, or a cheap
7) General colour of the ascomata: dark, bright yellow to red, of any
other colour (bare eyes).
8) Colour reactions of (first) thallus and (secondly) medulla, under
chemicals (bare eyes, chemicals, sometimes a cheap microscope).
9) Anatomical characters such as spore shape and colour, or chemical
reactions of thallus of ascocarp sections (chemicals, cheap
10) Colour of thallus and medulla as seen under a UV-lamp (UV-
11) Several, rather variable general features of the thallus, or of the
ascomata, e.g. isotomic versus anisotomic ramification, form, size
and shape of the areolae in crustose lichens (bare eyes, sometimes
helped by a lens).
12) Detailed anatomical characters, such as chemical reactions of
microscopic sections, presence size and shape of crystals in
apothecial sections, fine structure of spore walls, spore size, etc.
13) Presence of specific lichen substances (chromatography).
The present keys are not organised according to systematic
criteria, and especially not by genera. FRIDA, however, can generate
more “ortodox” keys, just by changing the order of characters.
Five keys are included into this book:
There is a general key to all “terricolous” lichens known from
Italy, also including some species not known from Italy but occurring
in neighbouring areas, whose presence in Italy is possible. This is the
most complete, but also the most difficult key. It contains dichotomies
between species which will be never found together, such as lichens
from the driest parts of Sicily and those restricted to the nival belt of
the Alps. The total number of taxa is 439.
The general key is followed by four smaller keys (ca. 200 taxa
each) obtained by connecting FRIDA with ITALIC, using two
ecological parameters: a) altitudinal distribution, b) type of substrate
(siliceous versus calcareous substrata).
Keys 1 and 2 include lichens occurring in upland areas, from
the subalpine belt of the Alps (Larix-stands) to above treeline
(Northern Italy only).
1) On acid to subneutral (acid to basic siliceous) substrata.
2) On subneutral to basic (basic siliceous to calciferous)
Keys 3 and 4 include lichens occurring from the mediterranean
to the montane belt (beech forests).
3) On acid to subneutral (acid to basic siliceous) substrata.
4) On subneutral to basic, (basic siliceous to calciferous)
An ecological buffer was invoked in the production of the
smaller keys: all of them include lichens occurring on intermediate,
subneutral substrata. The most common and ecologically wide-
ranging species were included in most keys: for example, Squamarina
cartilaginea, very common on calciferous substrata below treeline,
very rarely also occurs in the subalpine belt on basic siliceous
substrata. Formally, it should occur only in keys 2 and 4. However,
due to its commonness, and to the practical difficulty for beginners in
distinguishing among different types of siliceous substrata, it is found
in all of the simplified keys.