Notes on epilithic and epigeic lichens from granite and
gneiss outcrops in mountains of Makedonia, Greece,
with emphasis on northern species
Author: Christensen, Steen N.
Source: Willdenowia, 44(3) : 399-405
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
URL: https://doi.org/10.3372/wi.44.44310
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Willdenowia 44 – 2014
399
STEEN N. CHRISTENSEN1
Notes on epilithic and epigeic lichens from granite and gneiss outcrops in mountains
of Makedonia, Greece, with emphasis on northern species
Abstract
Christensen S. N.: Notes on epilithic and epigeic lichens from granite and gneiss outcrops in mountains of Makedonia, Greece, with emphasis on northern species. – Willdenowia 44: 399 – 405. 2014. – Version of record first published
online on 10 November 2014 ahead of inclusion in December 2014 issue; ISSN 1868-6397; © 2014 BGBM BerlinDahlem.
DOI: http://dx.doi.org/10.3372/wi.44.44310
The epilithic and epigeic lichen flora of eight localities with granite and gneiss outcrops in the mountains of Makedonia, N Greece has been investigated. Of the 46 taxa reported, seven species are new to Greece, viz.: Brodoa oroarcti
ca, Candelariella coralliza, Cetraria ericetorum, Lecanora swartzii, Parmelia serrana, Pseudephebe pubescens and
Ramalina strepsilis. One species is new to the Greek mainland (Umbilicaria polyphylla) and four species are new to
the region of Makedonia (Bryoria chalybeiformis, Melanelia disjuncta, Polychidium muscicola and Xanthoparmelia
loxodes). Altitudinal range and abundance are briefly discussed for a number of species.
Additional key words: Arctic-Alpine lichens, crystalline acidic rocks, Mediterranean-montane lichens
Introduction
Lichenological investigations of Greece has a long history resulting in a total of 1296 taxa (Abbott 2009). Since
then the number has been raised by nine (Sipman & Ahti
2011), four (Sipman 2012), 27 (Christensen & Alstrup
2013), 13 (Christensen 2014) and seven (this paper)
bringing the total number up to 1356. Considering the
varied geology and topography and the number of vegetation zones present, this number is certainly far too low.
Historically lichenological papers have mainly focussed on floristic data, reporting the presence of species from different areas (e.g. Zahlbruckner 1906; more
references in Abbott 2009). Only few papers deal with
the lichen flora of specific substrates or biotopes (e.g.
Steiner 1893; Krause & Klement 1962). During the last
three or four decades a number of papers have focussed
on recording lichen floras of different substrates. Zoller
& al. (1977) recorded the epigeic and epiphytic lichens
and mosses in woodlands in the mountains of N Greece.
Christensen (1989, 1994a – b, 1995b, 2000, 2007, 2014);
Mucina & al. (2000); Pirintsos & al. (1993, 1995, 1998)
and Riga-Karandinos (2000) reported on epiphytic lichens from a number of trees and shrubs. Epigeic lichens
were treated by Papp & al. (1999) and Christensen &
Alstrup (2013). In contrast, the lichen floras of different
bedrocks have only been treated in a few papers (Christensen & Alstrup 2013; Sipman & Raus 1999).
Except for Mt Olymbos (e.g. Hayek 1928; Szatala
1959; Christensen 1995a), the high mountains of the
Greek mainland are under-collected compared to the
lowlands and the islands, and Kriti in particular. In the
lowlands and the islands and on Mt Olymbos hard limestone (including dolomite and marble) is the predominant bedrock. Limestone lichens are, therefore, relatively
well represented in the total list of Greek lichens (Abbott
2009). Serpentinite and acidic crystalline rocks, which
are found as intrusions in the limestone rocks of the
higher mountains and some islands, have not been much
studied lichenologically (Krause & Klement 1962; Christensen & Alstrup 2013). This applies particularly to the
plutonic and metamorphic rocks of the high mountains.
1 Natural History Museum of Denmark, Botanical Garden & Museum, University of Copenhagen, Sølvgade 83, Entr. S, DK-1307
Copenhagen K, Denmark; e-mail: steenc@snm.ku.dk
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400
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
In a study on lichens of different rock
types Christensen & Alstrup (2013) reported a number of species with northern
distribution patterns, many of which were
not previously published for Greece, e.g.
Cornicularia normoerica, Dermatocarpon
intestiniforme, Melanelia stygia, Rhizocar
pon eupetraeoides, R. ferax, Rhizoplaca
chrysoleuca, R. melanophthalma and Spo
rastadia testudinea. Such an array of species indicates that rock outcrops elsewhere
in the high mountains of Greece might harbour Arctic-Alpine species and that they
are in fact more widely distributed than
publications so far show.
A field trip to mountains with known
outcrops of acidic plutonic and acidic metamorphic rocks was conducted to test this
hypothesis. In the field, collecting focus
was on the northern element of the macrolichen flora. Crustose lichens were only occasionally collected, as were macrolichens
with other distribution patterns. The results
are presented below.
Fig. 1. Greece with the position of the localities in W Makedonia (1 – 7) and E
Makedonia (8).
Material and methods
During one week in September 2013 seven localities in W
Makedonia known to have outcrops of granite and gneiss
were visited – locality no. 8 in the list below is a supplement to Christensen & Alstrup (2013) with only one
species, Bryoria chalybeiformis, reported (Fig. 1). The
locality numbers are given in the species list in brackets,
preceding the corresponding collection numbers. Latitudes and longitudes of the localities are map readings.
To distinguish between the Greek region of Makedonia and the Former Yugoslavian Republic of Macedonia
(F.Y.R.O.M.) the former is referred to as Makedonia and
the latter as Macedonia.
Names of provinces and larger landscapes follow The
Times Atlas of the World (1990). Geographical names for
Makedonia follow “Macedonia, 1: 200 000”. ROAD Editions, Athens Jul 2010 and “Freytag-Berndt Autokarte,
Griechenland, 1: 600 000”. Freytag-Berndt u. Artaria,
Wien. In the field the nomos maps of the Ethniki Statistiki
Ypiresia tis Ellados (National Statistical Office of Greece)
no 22: Nomos Kastorias, 1983 and no 46: Nomos Florinis,
1972 were used and some of the names of mountains and
summits are only to be found in these maps. Finally the forest map “, ”
. Y “” 1: 20 000” was
used for the Vrondous mountains.
The nomenclature is largely in accordance with Abbott
(2009). When other names are used, those used by Abbott
(2009) are given in synonymy. Whenever specimens are
labelled with older synonyms, those are also given.
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The Greek distribution of the species was checked
against the Greek checklist (Abbott 2009) and Christensen & Alstrup (2013). Asterisks are used to denote new
to Greece (***), new to the mainland of Greece (**), or
new to Makedonia (*).
The specimens are deposited in the Natural History
Museum of Denmark, Botanical Garden & Museum (C),
in the Botanical Museum in Berlin-Dahlem (B) and in the
author’s private herbarium.
List of localities
Nomos Florinas
1. S slope of Mt Varnous (), c. 14 km W of Florina, just above the village Pisoderi, 40°49'N, 21°15'E,
1450 m. – Open Juniperus communis thicket in opening
in Fagus woodland, S slope, incline 35°. – Bedrock augen gneiss. – On large boulders of gneiss. – 14 Sep 2013.
2. Mt Verno (), c. 1 km S of the summit Vitsi
(), at the monument, 40°39'N, 21°24'E, 1775 m. –
Granite outcrops in dry montane grassland. – 13 Sep 2013.
3. Mt Verno, c. 3.5 km S of the summit Vitsi, N slope
of the summit “1862”, 40°38'N, 21°23'E, 1750 – 1800 m.
– Granite outcrops in montane grassland on W slope, incline 20°. – 17 Sep 2013. – Fig. 2 & 3.
Nomos Kastorias
4. SW slope of Mt Verno, c. 12.5 km NE of Kastoria,
c. 1 km SE of the village Oxia, SW slope of the summit
“1653”, 40°37'N, 21°20'E, 1250 m. – Granite outcrops in
Willdenowia 44 – 2014
401
c. 17 km N of Serres, forest lot “II 8”, at the mountain
refuge, 41°14'N, 23°37'E, 1530–1550 m. – Pinus sylves
tris woodland, partly of scattered old tall trees, partly of
dense thickets of younger 7–8-m-tall trees, on SW-facing
slope. – Bedrock granite. – 14 Sep 1987. – This is the same
as loc. 27 of Christensen & Alstrup (2013).
Results
List of taxa
Fig. 2. Granite outcrops in montane grassland at 1750 – 1800 m
on Mt Vitsi (loc. 3). Note the population of Cornicularia nor
moerica in the foreground (many of the dark patches, which also
include specimens of Umbilicaria deusta and U. polyphylla).
montane grassland in large opening in Fagus woodland.
– 17 Sep 2013.
5. Mt Verno, c. 12 km NE of Kastoria, c. 1 km SE of the village Oxia, just N of the side road to the village Polykeraso,
40°36'N, 21°21'E, 1400 m. – Dry montane grassland in
large opening in Fagus woodland. – On granite boulder
and on the ground in patches of granite gravel. – 13 Sep
2013.
6. Mt Verno, SE slope of the summit “1505”, c. 8 km NNE
of Kastoria, NW of the village Tichio, along the Chloi-Vysinea road, 40°35'N, 21°18'E, 750 m. – Sun-exposed SSE
slope along road in Quercus woodland. – Weathered granite outcrops in Juniperus communis thicket. – 13 Sep 2013.
7. SE slope of Mt Verbista (), at the S part of
the Verno range, c. 17.5 km ENE of Kastoria, c. 1.5 km
NNE of the village Klisoura, 40°34'N, 21°29'E, 1200 m.
– Outcrops of augen gneiss in montane grassland on SSW
slope, incline 35°, in large opening in Quercus woodland.
– On S-facing gneiss rocks. – 16 Sep 2013.
Nomos Serron
8. Vrondous () Mts, Mt Lailia (),
SW slope of the summit “Ali Baba” ( ),
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Epilithic species
***Brodoa oroarctica (Krog) Goward – On top and on
N and E sides of granite boulders and granite rock
outcrops. (2) 14766; (3) 14873a, 14874, 14875. – “An
Arctic-Alpine, northern boreal, circumpolar species”
being “saxicolous on highly exposed siliceous …
rocks in Alpine areas or tundra heaths ...” (Thell &
Westberg 2011).
*Bryoria chalybeiformis (L.) Brodo & D. Hawksw. – On
N and E sides of granite rock outcrops. (3) 14871; (4)
14851; (8) 4448. – Distinguished from B. fuscescens
(Gyeln.) Brodo & D. Hawksw. by its uneven branches
and the substrate (Myllys & al. 2011). The sole previous record for Greece, from Thessalia, was regarded
as doubtful by Abbott (2009) due to the species having
a distinctly northern distribution. However, in light of
the recent reports of a relatively large number of species in the mountains of Greece with a northern affinity (Christensen & Svane 2009; Christensen & Alstrup
2013; and this paper) this argument is hardly valid.
Buellia badia (Fr.) A. Massal. – On decomposed lichens
on W and E sides of granite outcrops. (4) 14847,
14848, 14850.
Caloplaca pelodella (Nyl.) Hasse (C. conglomerata
(Bagl.) Jatta) – On weathered granite outcrop on SSE
slope. (6) with Xanthoparmelia tinctina 14732. On
the Greek mainland this species is only known from
two localities in Makedonia: Mt Vourinos (Christensen
2000) and Mt Avgo (Christensen & Alstrup 2013).
Caloplaca saxicola (Hoffm.) Nordin – On N side below
overhanging part of large gneiss boulder. (1) 14781,
14782a. The two previous records for Makedonia are
from the W part: Mt Avgo and Mt Triklarion (Christensen & Alstrup 2013).
***Candelariella coralliza (Nyl.) H. Magn. – On weathered granite outcrop on SSE slope. (6) with Xantho
parmelia tinctina 14732. – Arctic-Mediterranean
montane (Wirth 1995). In the Balkans apparently
only reported from Macedonia (Murati 1992; Rohrer
& al. 2012).
Candelariella vitellina (Hoffm.) Müll. Arg. – On N side of
granite boulder. (5) 14757.
Cornicularia normoerica (Gunn.) Du Rietz – On W side
of large gneiss boulder and on N, W and S sides of
granite boulders and granite outcrops. (1) 14777; (2)
14769; (3) 14863, 14864; (5) 14752, 14754. – Known
402
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
from a single locality in the Vrondous Mts in
E Makedonia at 1530 – 1625 m (Christensen &
Alstrup 2013). Of the sites with acidic crystalline rocks studied during the field trip in 2013
the species occurred on all the rock outcrops
above 1400 m. It often occurred in abundance
(Fig. 2).
Hypogymnia farinacea Zopf – On W side of granite
rocks. (2) 14767.
Hypogymnia physodes (L.) Nyl. – On vertical N
side of large gneiss boulder. (1) 14786. – Epilithic occurrences of this normally epiphytic
species are relatively rare.
Lasallia pustulata (L.) Mérat – On top and W side
of large gneiss boulder and on top, NE, S and E
sides of granite rock outcrops. (1) 14776, 14779,
14788; (3) 14870; (4) 14845, 14849; (6) 14736.
Fig. 3. At 1750 – 1800 m the upper limit of the Fagus woodland with its
Lecanora bolcana (Pollini) Poelt – On granite sharp margin is clearly a result of the upper part of the woodland havrock outcrops and on pebbles on the ground in ing been cut and the area subsequently subjected to grazing, resulting
patches of granite gravel. (4) 14854; (5) 14764; in montane grassland. On the granite outcrop in the foreground a community mainly of Umbilicaria cylindrica and U. polyphylla is present.
(6) with Xanthoparmelia tinctina 14732.
***Lecanora swartzii (Ach.) Ach. – Below over- Mt Vitsi (loc. 3).
hanging part of S-facing gneiss rock. (7) with
Pleopsidium chlorophanum 14835. – This species is
here referred to P. serrana. “The world distribution is
(Arctic-) boreal to Mediterranean montane (Wirth
poorly known and it is hitherto known from southern,
1995). In the Balkans it has been published from
central and western Europe and the Canary Islands.
Bulgaria (Mayrhofer & al. 2005) and Macedonia
The Atlantic preference is less pronounced compared
(Mayrhofer & al. 2013).
with P. ernstiae” (Thell & al. 2011).
Lecidella carpathica Körb. – On weathered granite out- Parmelina tiliacea (Hoffm.) Hale – On S side and top of
crop on SSE slope. (6) with Xanthoparmelia tinctina
gneiss boulders and outcrops. (1) 14780, 14789; (7)
14732.
14836.
*Melanelia disjuncta (Erichsen) Essl. – On NE side of Pertusaria albescens (Huds.) M. Choisy & Werner – On
large gneiss boulder. (1) 14784. – Known from two
NW side of gneiss rock. (7) 14838. – Epilithic records
records, from Attiki (Attica) and Evvia, respectively
of this mainly epiphytic species constitute only a minor
(Abbott 2009).
portion of the many records cited by Abbott (2009).
Melanelixia fuliginosa (Fr. ex Duby) O. Blanco & al. Physcia dubia (Hoffm.) Lettau – On N side below over(Melanelia fuliginosa (Fr. ex Duby) Essl.) – Below
hanging part of large gneiss boulder. (1) 14782. – This
overhanging part of NW side of granite outcrop. (4)
species is only known from one record for Makedonia
with Xanthoparmelia tinctina 14860.
and one for Thessalia (Christensen 2000; Christensen
Parmelia saxatilis (L.) Ach. – On NE side of granite
& Svane 2009).
boulder shaded by Juniperus. (6) 14735.
Physcia tenella (Scop.) DC. – On top of granite boulder.
***Parmelia serrana A. Crespo & al. – On N and E sides
(5) 14762.
of boulders and outcrops of gneiss and granite. (1) Pleopsidium flavum (Bellardi) Körb. – On NE side of
14785; (4) 14846; (5) 14755. – Parmelia ernstiae,
gneiss boulder and below overhanging part of SP. saxatilis and P. serrana are rather similar spefacing gneiss rock. (1) 14784a; (7) 14835. – Known
cies. They share a number of characters, which they
from Kriti and from easternmost Makedonia (Kavala)
express to different degrees; they are therefore dis(Abbott 2009). It is an Arctic to Alpine, subcontinentinguished by a combination of characters, most of
tal species (Wirth 1995).
which are not exclusive to any of the species (Thell *Polychidium muscicola (Sw.) Gray – On moss on SW
& al. 2011). Specimens with rather broad and overside of large gneiss boulder. (1) 14778. – Previously
lapping marginal lobes, branched as well as simple
known only from one find in Thessalia (Abbott 2009).
rhizinae, marginal isidia in addition to laminal isidia ***Pseudephebe pubescens (L.) M. Choisy – On E side
(the latter mainly on ridges in peripheral parts but
of granite rock outcrop. (3) 14871a. – This species has
may also cover large patches of the central part of the
an Arctic-(montane)Alpine distribution (Wirth 1995).
thallus), no or occasionally little pruina, upper surface
In the Balkans it has been published from Bulgaria
± foveolate and no lobules or rarely with lobules de(Mayrhofer & al. 2005) and Macedonia (Mayrhofer
veloped from isidia in the older part of the thallus are
& al. 2013).
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Willdenowia 44 – 2014
Pseudevernia furfuracea (L.) Zopf – On N side of granite
boulder. (5) 14761. – This mainly epiphytic species is
widespread in Greece (Abbott 2009).
Ramalina capitata (Ach.) Nyl. – On top and W side of
granite boulders. (2) 14768; (5) 14751. – Known
from the Aegean island Naxos, Ipiros and E Makedonia (Abbott 2009; Christensen & Alstrup 2013).
***Ramalina strepsilis (Ach.) Zahlbr. – On top of granite
rock outcrops. (3) 14866; (4) 14853. – Ramalina strep
silis is sometimes included in R. capitata (Santesson
2011) or is treated as a variety of that taxon (Nimis
& Poelt 1987). In order to bring attention to the morphological variation in the present material of the R.
capitata complex, and as the taxonomy seems not to
be finally settled, the taxa are here treated at the species level. Thus, R. strepsilis is distinguished from R.
capitata by the labriform soralia along the margin of
the lobes in addition to the capitiform soralia apically
on the lobes (R. capitata in this sense only has the
latter type of soralia). As this taxon is often treated
as being conspecific with R. capitata, the geographical distribution is not clear. Ramalina strepsilis has,
however, been reported several times from Macedonia (Mayrhofer & al. 2013).
Umbilicaria cylindrica (L.) Delise – On top of granite
rock outcrop. (3) 14865.
Umbilicaria deusta (L.) Baumg. – On N, E and W sides
of granite boulders and outcrops. (2) 14770; (3)
14869, 14872. – The one previous record for Makedonia is from the Vrondous Mts E of Thessaloniki
(Christensen & Alstrup 2013).
**Umbilicaria polyphylla (L.) Baumg. – On S side of
granite rock outcrop. (3) 14868. – Hitherto only known
from the Aegean island Samothraki (Abbott 2009).
Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale – On top
of granite outcrop and on base of E side of granite
boulder. (4)14852; (5)14760.
*Xanthoparmelia loxodes (Nyl.) O. Blanco & al. (Neo
fuscelia loxodes (Nyl.) Essl.) – On top of large granite
boulder. (1) 14791, 14792. – Known from the Aegean
islands Aegina, Kriti, Lesvos and Kos and from Peloponnisos (Abbott 2009).
Xanthoparmelia pulla (Ach.) O. Blanco & al. (Neofusce
lia pulla (Ach.) Essl.) – On granite outcrop at ground
level in patches of granite gravel and on N and almost vertical W sides of granite boulders. (5) 14750,
14753, 14756; (6) 14734.
Xanthoparmelia stenophylla (Ach.) Ahti & D. Hawksw.
– On top of large gneiss boulder. (1) 14790.
Xanthoparmelia tinctina (Maheu & A. Gillet) Hale – Below overhanging part of NW side of granite outcrop
and on weathered granite outcrop on SSE slope. (4)
14860; (6) 14732.
Epigeic species
Cetraria aculeata (Schreb.) Fr. – On the ground among
gneiss boulders and granite outcrops and on the
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403
ground in patches of granite gravel. (1) 14793; (4)
14859, 14862; (5) 14737, 14742.
***Cetraria ericetorum Opiz – On the ground among
granite outcrops and on the ground in patches of
granite gravel. (4) 14856; (5) 14744. – In Europe it
is distributed from the Arctic to the temperate zone,
common in the Nordic countries and the Alps, scattered in E Europe and the Balkans, reaching Bulgaria
(Kärnefelt 1979: Fig. 42, Mayrhofer & al. 2005). In the
Balkans it is also reported from Slovenia (Mayrhofer
2006), Bosnia and Herzegovina (Bilovitz & Mayrhofer 2011), Montenegro (Kneževic & Mayrhofer 2009)
and Macedonia (Mayrhofer & al. 2012, 2013).
Cetraria islandica (L.) Ach. subsp. islandica – On the
ground among granite outcrops and on the ground in
patches of granite gravel. (4) 14861; (5) 14741.
Cladonia cervicornis (Ach.) Flot. subsp. cervicornis –
On shallow soil on granite outcrop and on the ground
in patches of granite gravel. (5) 14745, 14763. – Only
one previous record for Makedonia: Mt Cholomondas of Chalkidiki (Abbott 2009).
Cladonia foliacea (Huds.) Willd. – On the ground among
granite outcrops and on the ground in patches of granite gravel. (4) 14855; (5) 14747; (6) 14733.
Cladonia furcata (Huds.) Schrad. – On the ground among
granite outcrops and on the ground in patches of granite gravel. (4) with Cetraria aculeata 14862; (5) 14743.
Cladonia rangiformis Hoffm. – On the ground among
granite outcrops. (4) 14858.
Diploschistes scruposus (Schreb.) Norman var. scrupo
sus – On the ground in patches of granite gravel. (5)
14748. – Only two previous records for Makedonia,
both from Halkidiki (Abbott 2009).
Leptogium pulvinatum (Hoffm.) Otálora var. pulvinatum
– On litter. (6) 14731. – For Makedonia this taxon
is only reported from two localities on Mt Olymbos
(Szatala 1959).
Peltigera rufescens (Weiss) Humb. – On the ground among
outcrops of gneiss and granite and on the ground in
patches of granite gravel. (1) 14794; (4) 14857; (5)
14738.
Discussion
At Mt Verno, the principal sampling area, including localities 2 – 6, the vegetation zone of the deciduous oaks, the
Quercetalia pubescentis, range from 700 – 1000 m. From
850 – 1700 m the dominant tree species is Fagus sylvatica
of the Fagetalia zone (Fotiadis & al. 2005). In both zones
the woodlands have been cleared over large expanses giving room for montane grasslands used for the grazing of
livestock (Fig. 3). The lichens were sampled on granite
and gneiss outcrops and boulders in these grasslands (loc.
2, 3, 4, 5, 7, Fig. 2). On Mt Varnous (loc. 1) and on Mt
Verbista (loc. 7) the zonation is similar. At loc. 1 and 6 the
lichens were collected on gneiss outcrops and boulders in
404
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
Juniperus communis thickets. The environment of the localities is open, exposed to wind and sun (the grasslands,
Fig. 2), or semi-open (the Juniperus thickets). Juniperus
communis thickets are indicative of former grazing areas
– at least it is the case in N Europe outside fell-heaths and
other natural vegetation types. Thus, with the exception of
the outcrops and boulders, the biotopes are anthropogenic
in origin (Fig. 3).
Christensen & Svane (2009) reported a number of
species with northern affinities from mountains of N
Greece. Later Christensen & Alstrup (2013) published
species lists from different rock types from a number of
sites in the mountains of N Greece. They found that crystalline rocks at high altitude harbour a number of species
with boreal to Arctic and Alpine distribution. Allowing
for the few study sites in the present study, this seems
to confirm a connection between crystalline rocks, high
altitude and a host of species with a northern distribution
type. The outcrops and boulders of granite and gneiss
included in the present study were found at altitudes between 750 m and 1800 m. The three Umbilicaria species
(U. cylindrica, U. deusta and U. polyphylla) occurred
only above 1750 m, while Lasallia pustulata was found
as low as 750 m. Cetraria ericetorum and C. islandica
were not found below 1250 m, while C. aculeata was
found as low as 950 m (it occurs as low as sea level –
Linda in Arcadia 2013). Bryoria chalybeiformis was
found above 1250 m. Cornicularia normoerica occurred
above 1400 m. Pseudephebe pubescens was found above
1750 m. Ramalina capitata was found above 1400 m and
R. strepsilis above 1250 m. Brodoa oroarctica occurred
above 1750 m. Lecanora swartzii occurred at 1200 m.
The distribution of the above-mentioned species
along the altitude gradient represented by the seven
(eight) localities of the present study seems to mirror
the geographical distribution of the species: The species
found in the highest levels are those with the most pronounced bias toward north in their distribution patterns.
Of the two localities above 1750 m the Umbilicaria
species occurred on one (U. cylindrica, U. polyphylla) or
both (U. deusta) suggesting that the three species probably are very common on acidic rocks at this altitude.
Lasallia pustulata was found at all altitudes in four of the
seven localities and seems, therefore, to be rather common. Cornicularia normoerica was found at all four localities above 1400 m. It must be very common on acidic
rocks at this altitude. The three Cetraria species are very
sensitive to the high grazing intensity of these montane
grasslands. They were found as fragmented cushions
or as detached lobes patchily in the grass swards. The
species were, therefore, not an obvious element in the
grassland and could easily have been overseen in some of
the localities, as time did not allow for intensive searching. Bryoria chalybeiformis was found at two of the five
localities above 1250 m (or three of the six, if loc. 8 is
included). It is, therefore, likely to be rather common
on acidic rocks at this altitude. The ornithocoprophilous
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character of Ramalina capitata and R. strepsilis makes it
difficult to evaluate their occurrences. Together they occur at four of the seven localities. The presence of Brodoa
oroarctica at the two very highest localities underlines its
Arctic-Alpine affinity. It may be relatively rare. The same
applies to Pseudephebe pubescens, which was found at
one of these two localities (above 1750 m).
The findings of this small study, with emphasis on
macrolichens from a small number of sites, may indicate that the northern element of the Greek lichen flora is
much more abundant than suggested from previous publications. Macrolichens of acidic crystalline rocks of the
high mountains are certainly under-collected in Greece.
This applies even more to crustose lichens.
Acknowledgements
Thanks are due to Helmut Mayrhofer (GZU) and another,
anonymous reviewer for constructive suggestions on an
earlier draft of this paper. Nicholas Turland kindly prepared the map.
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Willdenowia 44 – 2014
399
STEEN N. CHRISTENSEN1
Notes on epilithic and epigeic lichens from granite and gneiss outcrops in mountains
of Makedonia, Greece, with emphasis on northern species
Abstract
Christensen S. N.: Notes on epilithic and epigeic lichens from granite and gneiss outcrops in mountains of Makedonia, Greece, with emphasis on northern species. – Willdenowia 44: 399 – 405. 2014. – Version of record first published
online on 10 November 2014 ahead of inclusion in December 2014 issue; ISSN 1868-6397; © 2014 BGBM BerlinDahlem.
DOI: http://dx.doi.org/10.3372/wi.44.44310
The epilithic and epigeic lichen flora of eight localities with granite and gneiss outcrops in the mountains of Makedonia, N Greece has been investigated. Of the 46 taxa reported, seven species are new to Greece, viz.: Brodoa oroarcti
ca, Candelariella coralliza, Cetraria ericetorum, Lecanora swartzii, Parmelia serrana, Pseudephebe pubescens and
Ramalina strepsilis. One species is new to the Greek mainland (Umbilicaria polyphylla) and four species are new to
the region of Makedonia (Bryoria chalybeiformis, Melanelia disjuncta, Polychidium muscicola and Xanthoparmelia
loxodes). Altitudinal range and abundance are briefly discussed for a number of species.
Additional key words: Arctic-Alpine lichens, crystalline acidic rocks, Mediterranean-montane lichens
Introduction
Lichenological investigations of Greece has a long history resulting in a total of 1296 taxa (Abbott 2009). Since
then the number has been raised by nine (Sipman & Ahti
2011), four (Sipman 2012), 27 (Christensen & Alstrup
2013), 13 (Christensen 2014) and seven (this paper)
bringing the total number up to 1356. Considering the
varied geology and topography and the number of vegetation zones present, this number is certainly far too low.
Historically lichenological papers have mainly focussed on floristic data, reporting the presence of species from different areas (e.g. Zahlbruckner 1906; more
references in Abbott 2009). Only few papers deal with
the lichen flora of specific substrates or biotopes (e.g.
Steiner 1893; Krause & Klement 1962). During the last
three or four decades a number of papers have focussed
on recording lichen floras of different substrates. Zoller
& al. (1977) recorded the epigeic and epiphytic lichens
and mosses in woodlands in the mountains of N Greece.
Christensen (1989, 1994a – b, 1995b, 2000, 2007, 2014);
Mucina & al. (2000); Pirintsos & al. (1993, 1995, 1998)
and Riga-Karandinos (2000) reported on epiphytic lichens from a number of trees and shrubs. Epigeic lichens
were treated by Papp & al. (1999) and Christensen &
Alstrup (2013). In contrast, the lichen floras of different
bedrocks have only been treated in a few papers (Christensen & Alstrup 2013; Sipman & Raus 1999).
Except for Mt Olymbos (e.g. Hayek 1928; Szatala
1959; Christensen 1995a), the high mountains of the
Greek mainland are under-collected compared to the
lowlands and the islands, and Kriti in particular. In the
lowlands and the islands and on Mt Olymbos hard limestone (including dolomite and marble) is the predominant bedrock. Limestone lichens are, therefore, relatively
well represented in the total list of Greek lichens (Abbott
2009). Serpentinite and acidic crystalline rocks, which
are found as intrusions in the limestone rocks of the
higher mountains and some islands, have not been much
studied lichenologically (Krause & Klement 1962; Christensen & Alstrup 2013). This applies particularly to the
plutonic and metamorphic rocks of the high mountains.
1 Natural History Museum of Denmark, Botanical Garden & Museum, University of Copenhagen, Sølvgade 83, Entr. S, DK-1307
Copenhagen K, Denmark; e-mail: steenc@snm.ku.dk
400
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
In a study on lichens of different rock
types Christensen & Alstrup (2013) reported a number of species with northern
distribution patterns, many of which were
not previously published for Greece, e.g.
Cornicularia normoerica, Dermatocarpon
intestiniforme, Melanelia stygia, Rhizocar
pon eupetraeoides, R. ferax, Rhizoplaca
chrysoleuca, R. melanophthalma and Spo
rastadia testudinea. Such an array of species indicates that rock outcrops elsewhere
in the high mountains of Greece might harbour Arctic-Alpine species and that they
are in fact more widely distributed than
publications so far show.
A field trip to mountains with known
outcrops of acidic plutonic and acidic metamorphic rocks was conducted to test this
hypothesis. In the field, collecting focus
was on the northern element of the macrolichen flora. Crustose lichens were only occasionally collected, as were macrolichens
with other distribution patterns. The results
are presented below.
Fig. 1. Greece with the position of the localities in W Makedonia (1 – 7) and E
Makedonia (8).
Material and methods
During one week in September 2013 seven localities in W
Makedonia known to have outcrops of granite and gneiss
were visited – locality no. 8 in the list below is a supplement to Christensen & Alstrup (2013) with only one
species, Bryoria chalybeiformis, reported (Fig. 1). The
locality numbers are given in the species list in brackets,
preceding the corresponding collection numbers. Latitudes and longitudes of the localities are map readings.
To distinguish between the Greek region of Makedonia and the Former Yugoslavian Republic of Macedonia
(F.Y.R.O.M.) the former is referred to as Makedonia and
the latter as Macedonia.
Names of provinces and larger landscapes follow The
Times Atlas of the World (1990). Geographical names for
Makedonia follow “Macedonia, 1: 200 000”. ROAD Editions, Athens Jul 2010 and “Freytag-Berndt Autokarte,
Griechenland, 1: 600 000”. Freytag-Berndt u. Artaria,
Wien. In the field the nomos maps of the Ethniki Statistiki
Ypiresia tis Ellados (National Statistical Office of Greece)
no 22: Nomos Kastorias, 1983 and no 46: Nomos Florinis,
1972 were used and some of the names of mountains and
summits are only to be found in these maps. Finally the forest map “, ”
. Y “” 1: 20 000” was
used for the Vrondous mountains.
The nomenclature is largely in accordance with Abbott
(2009). When other names are used, those used by Abbott
(2009) are given in synonymy. Whenever specimens are
labelled with older synonyms, those are also given.
The Greek distribution of the species was checked
against the Greek checklist (Abbott 2009) and Christensen & Alstrup (2013). Asterisks are used to denote new
to Greece (***), new to the mainland of Greece (**), or
new to Makedonia (*).
The specimens are deposited in the Natural History
Museum of Denmark, Botanical Garden & Museum (C),
in the Botanical Museum in Berlin-Dahlem (B) and in the
author’s private herbarium.
List of localities
Nomos Florinas
1. S slope of Mt Varnous (), c. 14 km W of Florina, just above the village Pisoderi, 40°49'N, 21°15'E,
1450 m. – Open Juniperus communis thicket in opening
in Fagus woodland, S slope, incline 35°. – Bedrock augen gneiss. – On large boulders of gneiss. – 14 Sep 2013.
2. Mt Verno (), c. 1 km S of the summit Vitsi
(), at the monument, 40°39'N, 21°24'E, 1775 m. –
Granite outcrops in dry montane grassland. – 13 Sep 2013.
3. Mt Verno, c. 3.5 km S of the summit Vitsi, N slope
of the summit “1862”, 40°38'N, 21°23'E, 1750 – 1800 m.
– Granite outcrops in montane grassland on W slope, incline 20°. – 17 Sep 2013. – Fig. 2 & 3.
Nomos Kastorias
4. SW slope of Mt Verno, c. 12.5 km NE of Kastoria,
c. 1 km SE of the village Oxia, SW slope of the summit
“1653”, 40°37'N, 21°20'E, 1250 m. – Granite outcrops in
Willdenowia 44 – 2014
401
c. 17 km N of Serres, forest lot “II 8”, at the mountain
refuge, 41°14'N, 23°37'E, 1530–1550 m. – Pinus sylves
tris woodland, partly of scattered old tall trees, partly of
dense thickets of younger 7–8-m-tall trees, on SW-facing
slope. – Bedrock granite. – 14 Sep 1987. – This is the same
as loc. 27 of Christensen & Alstrup (2013).
Results
List of taxa
Fig. 2. Granite outcrops in montane grassland at 1750 – 1800 m
on Mt Vitsi (loc. 3). Note the population of Cornicularia nor
moerica in the foreground (many of the dark patches, which also
include specimens of Umbilicaria deusta and U. polyphylla).
montane grassland in large opening in Fagus woodland.
– 17 Sep 2013.
5. Mt Verno, c. 12 km NE of Kastoria, c. 1 km SE of the village Oxia, just N of the side road to the village Polykeraso,
40°36'N, 21°21'E, 1400 m. – Dry montane grassland in
large opening in Fagus woodland. – On granite boulder
and on the ground in patches of granite gravel. – 13 Sep
2013.
6. Mt Verno, SE slope of the summit “1505”, c. 8 km NNE
of Kastoria, NW of the village Tichio, along the Chloi-Vysinea road, 40°35'N, 21°18'E, 750 m. – Sun-exposed SSE
slope along road in Quercus woodland. – Weathered granite outcrops in Juniperus communis thicket. – 13 Sep 2013.
7. SE slope of Mt Verbista (), at the S part of
the Verno range, c. 17.5 km ENE of Kastoria, c. 1.5 km
NNE of the village Klisoura, 40°34'N, 21°29'E, 1200 m.
– Outcrops of augen gneiss in montane grassland on SSW
slope, incline 35°, in large opening in Quercus woodland.
– On S-facing gneiss rocks. – 16 Sep 2013.
Nomos Serron
8. Vrondous () Mts, Mt Lailia (),
SW slope of the summit “Ali Baba” ( ),
Epilithic species
***Brodoa oroarctica (Krog) Goward – On top and on
N and E sides of granite boulders and granite rock
outcrops. (2) 14766; (3) 14873a, 14874, 14875. – “An
Arctic-Alpine, northern boreal, circumpolar species”
being “saxicolous on highly exposed siliceous …
rocks in Alpine areas or tundra heaths ...” (Thell &
Westberg 2011).
*Bryoria chalybeiformis (L.) Brodo & D. Hawksw. – On
N and E sides of granite rock outcrops. (3) 14871; (4)
14851; (8) 4448. – Distinguished from B. fuscescens
(Gyeln.) Brodo & D. Hawksw. by its uneven branches
and the substrate (Myllys & al. 2011). The sole previous record for Greece, from Thessalia, was regarded
as doubtful by Abbott (2009) due to the species having
a distinctly northern distribution. However, in light of
the recent reports of a relatively large number of species in the mountains of Greece with a northern affinity (Christensen & Svane 2009; Christensen & Alstrup
2013; and this paper) this argument is hardly valid.
Buellia badia (Fr.) A. Massal. – On decomposed lichens
on W and E sides of granite outcrops. (4) 14847,
14848, 14850.
Caloplaca pelodella (Nyl.) Hasse (C. conglomerata
(Bagl.) Jatta) – On weathered granite outcrop on SSE
slope. (6) with Xanthoparmelia tinctina 14732. On
the Greek mainland this species is only known from
two localities in Makedonia: Mt Vourinos (Christensen
2000) and Mt Avgo (Christensen & Alstrup 2013).
Caloplaca saxicola (Hoffm.) Nordin – On N side below
overhanging part of large gneiss boulder. (1) 14781,
14782a. The two previous records for Makedonia are
from the W part: Mt Avgo and Mt Triklarion (Christensen & Alstrup 2013).
***Candelariella coralliza (Nyl.) H. Magn. – On weathered granite outcrop on SSE slope. (6) with Xantho
parmelia tinctina 14732. – Arctic-Mediterranean
montane (Wirth 1995). In the Balkans apparently
only reported from Macedonia (Murati 1992; Rohrer
& al. 2012).
Candelariella vitellina (Hoffm.) Müll. Arg. – On N side of
granite boulder. (5) 14757.
Cornicularia normoerica (Gunn.) Du Rietz – On W side
of large gneiss boulder and on N, W and S sides of
granite boulders and granite outcrops. (1) 14777; (2)
14769; (3) 14863, 14864; (5) 14752, 14754. – Known
402
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
from a single locality in the Vrondous Mts in
E Makedonia at 1530 – 1625 m (Christensen &
Alstrup 2013). Of the sites with acidic crystalline rocks studied during the field trip in 2013
the species occurred on all the rock outcrops
above 1400 m. It often occurred in abundance
(Fig. 2).
Hypogymnia farinacea Zopf – On W side of granite
rocks. (2) 14767.
Hypogymnia physodes (L.) Nyl. – On vertical N
side of large gneiss boulder. (1) 14786. – Epilithic occurrences of this normally epiphytic
species are relatively rare.
Lasallia pustulata (L.) Mérat – On top and W side
of large gneiss boulder and on top, NE, S and E
sides of granite rock outcrops. (1) 14776, 14779,
14788; (3) 14870; (4) 14845, 14849; (6) 14736.
Fig. 3. At 1750 – 1800 m the upper limit of the Fagus woodland with its
Lecanora bolcana (Pollini) Poelt – On granite sharp margin is clearly a result of the upper part of the woodland havrock outcrops and on pebbles on the ground in ing been cut and the area subsequently subjected to grazing, resulting
patches of granite gravel. (4) 14854; (5) 14764; in montane grassland. On the granite outcrop in the foreground a community mainly of Umbilicaria cylindrica and U. polyphylla is present.
(6) with Xanthoparmelia tinctina 14732.
***Lecanora swartzii (Ach.) Ach. – Below over- Mt Vitsi (loc. 3).
hanging part of S-facing gneiss rock. (7) with
Pleopsidium chlorophanum 14835. – This species is
here referred to P. serrana. “The world distribution is
(Arctic-) boreal to Mediterranean montane (Wirth
poorly known and it is hitherto known from southern,
1995). In the Balkans it has been published from
central and western Europe and the Canary Islands.
Bulgaria (Mayrhofer & al. 2005) and Macedonia
The Atlantic preference is less pronounced compared
(Mayrhofer & al. 2013).
with P. ernstiae” (Thell & al. 2011).
Lecidella carpathica Körb. – On weathered granite out- Parmelina tiliacea (Hoffm.) Hale – On S side and top of
crop on SSE slope. (6) with Xanthoparmelia tinctina
gneiss boulders and outcrops. (1) 14780, 14789; (7)
14732.
14836.
*Melanelia disjuncta (Erichsen) Essl. – On NE side of Pertusaria albescens (Huds.) M. Choisy & Werner – On
large gneiss boulder. (1) 14784. – Known from two
NW side of gneiss rock. (7) 14838. – Epilithic records
records, from Attiki (Attica) and Evvia, respectively
of this mainly epiphytic species constitute only a minor
(Abbott 2009).
portion of the many records cited by Abbott (2009).
Melanelixia fuliginosa (Fr. ex Duby) O. Blanco & al. Physcia dubia (Hoffm.) Lettau – On N side below over(Melanelia fuliginosa (Fr. ex Duby) Essl.) – Below
hanging part of large gneiss boulder. (1) 14782. – This
overhanging part of NW side of granite outcrop. (4)
species is only known from one record for Makedonia
with Xanthoparmelia tinctina 14860.
and one for Thessalia (Christensen 2000; Christensen
Parmelia saxatilis (L.) Ach. – On NE side of granite
& Svane 2009).
boulder shaded by Juniperus. (6) 14735.
Physcia tenella (Scop.) DC. – On top of granite boulder.
***Parmelia serrana A. Crespo & al. – On N and E sides
(5) 14762.
of boulders and outcrops of gneiss and granite. (1) Pleopsidium flavum (Bellardi) Körb. – On NE side of
14785; (4) 14846; (5) 14755. – Parmelia ernstiae,
gneiss boulder and below overhanging part of SP. saxatilis and P. serrana are rather similar spefacing gneiss rock. (1) 14784a; (7) 14835. – Known
cies. They share a number of characters, which they
from Kriti and from easternmost Makedonia (Kavala)
express to different degrees; they are therefore dis(Abbott 2009). It is an Arctic to Alpine, subcontinentinguished by a combination of characters, most of
tal species (Wirth 1995).
which are not exclusive to any of the species (Thell *Polychidium muscicola (Sw.) Gray – On moss on SW
& al. 2011). Specimens with rather broad and overside of large gneiss boulder. (1) 14778. – Previously
lapping marginal lobes, branched as well as simple
known only from one find in Thessalia (Abbott 2009).
rhizinae, marginal isidia in addition to laminal isidia ***Pseudephebe pubescens (L.) M. Choisy – On E side
(the latter mainly on ridges in peripheral parts but
of granite rock outcrop. (3) 14871a. – This species has
may also cover large patches of the central part of the
an Arctic-(montane)Alpine distribution (Wirth 1995).
thallus), no or occasionally little pruina, upper surface
In the Balkans it has been published from Bulgaria
± foveolate and no lobules or rarely with lobules de(Mayrhofer & al. 2005) and Macedonia (Mayrhofer
veloped from isidia in the older part of the thallus are
& al. 2013).
Willdenowia 44 – 2014
Pseudevernia furfuracea (L.) Zopf – On N side of granite
boulder. (5) 14761. – This mainly epiphytic species is
widespread in Greece (Abbott 2009).
Ramalina capitata (Ach.) Nyl. – On top and W side of
granite boulders. (2) 14768; (5) 14751. – Known
from the Aegean island Naxos, Ipiros and E Makedonia (Abbott 2009; Christensen & Alstrup 2013).
***Ramalina strepsilis (Ach.) Zahlbr. – On top of granite
rock outcrops. (3) 14866; (4) 14853. – Ramalina strep
silis is sometimes included in R. capitata (Santesson
2011) or is treated as a variety of that taxon (Nimis
& Poelt 1987). In order to bring attention to the morphological variation in the present material of the R.
capitata complex, and as the taxonomy seems not to
be finally settled, the taxa are here treated at the species level. Thus, R. strepsilis is distinguished from R.
capitata by the labriform soralia along the margin of
the lobes in addition to the capitiform soralia apically
on the lobes (R. capitata in this sense only has the
latter type of soralia). As this taxon is often treated
as being conspecific with R. capitata, the geographical distribution is not clear. Ramalina strepsilis has,
however, been reported several times from Macedonia (Mayrhofer & al. 2013).
Umbilicaria cylindrica (L.) Delise – On top of granite
rock outcrop. (3) 14865.
Umbilicaria deusta (L.) Baumg. – On N, E and W sides
of granite boulders and outcrops. (2) 14770; (3)
14869, 14872. – The one previous record for Makedonia is from the Vrondous Mts E of Thessaloniki
(Christensen & Alstrup 2013).
**Umbilicaria polyphylla (L.) Baumg. – On S side of
granite rock outcrop. (3) 14868. – Hitherto only known
from the Aegean island Samothraki (Abbott 2009).
Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale – On top
of granite outcrop and on base of E side of granite
boulder. (4)14852; (5)14760.
*Xanthoparmelia loxodes (Nyl.) O. Blanco & al. (Neo
fuscelia loxodes (Nyl.) Essl.) – On top of large granite
boulder. (1) 14791, 14792. – Known from the Aegean
islands Aegina, Kriti, Lesvos and Kos and from Peloponnisos (Abbott 2009).
Xanthoparmelia pulla (Ach.) O. Blanco & al. (Neofusce
lia pulla (Ach.) Essl.) – On granite outcrop at ground
level in patches of granite gravel and on N and almost vertical W sides of granite boulders. (5) 14750,
14753, 14756; (6) 14734.
Xanthoparmelia stenophylla (Ach.) Ahti & D. Hawksw.
– On top of large gneiss boulder. (1) 14790.
Xanthoparmelia tinctina (Maheu & A. Gillet) Hale – Below overhanging part of NW side of granite outcrop
and on weathered granite outcrop on SSE slope. (4)
14860; (6) 14732.
Epigeic species
Cetraria aculeata (Schreb.) Fr. – On the ground among
gneiss boulders and granite outcrops and on the
403
ground in patches of granite gravel. (1) 14793; (4)
14859, 14862; (5) 14737, 14742.
***Cetraria ericetorum Opiz – On the ground among
granite outcrops and on the ground in patches of
granite gravel. (4) 14856; (5) 14744. – In Europe it
is distributed from the Arctic to the temperate zone,
common in the Nordic countries and the Alps, scattered in E Europe and the Balkans, reaching Bulgaria
(Kärnefelt 1979: Fig. 42, Mayrhofer & al. 2005). In the
Balkans it is also reported from Slovenia (Mayrhofer
2006), Bosnia and Herzegovina (Bilovitz & Mayrhofer 2011), Montenegro (Kneževic & Mayrhofer 2009)
and Macedonia (Mayrhofer & al. 2012, 2013).
Cetraria islandica (L.) Ach. subsp. islandica – On the
ground among granite outcrops and on the ground in
patches of granite gravel. (4) 14861; (5) 14741.
Cladonia cervicornis (Ach.) Flot. subsp. cervicornis –
On shallow soil on granite outcrop and on the ground
in patches of granite gravel. (5) 14745, 14763. – Only
one previous record for Makedonia: Mt Cholomondas of Chalkidiki (Abbott 2009).
Cladonia foliacea (Huds.) Willd. – On the ground among
granite outcrops and on the ground in patches of granite gravel. (4) 14855; (5) 14747; (6) 14733.
Cladonia furcata (Huds.) Schrad. – On the ground among
granite outcrops and on the ground in patches of granite gravel. (4) with Cetraria aculeata 14862; (5) 14743.
Cladonia rangiformis Hoffm. – On the ground among
granite outcrops. (4) 14858.
Diploschistes scruposus (Schreb.) Norman var. scrupo
sus – On the ground in patches of granite gravel. (5)
14748. – Only two previous records for Makedonia,
both from Halkidiki (Abbott 2009).
Leptogium pulvinatum (Hoffm.) Otálora var. pulvinatum
– On litter. (6) 14731. – For Makedonia this taxon
is only reported from two localities on Mt Olymbos
(Szatala 1959).
Peltigera rufescens (Weiss) Humb. – On the ground among
outcrops of gneiss and granite and on the ground in
patches of granite gravel. (1) 14794; (4) 14857; (5)
14738.
Discussion
At Mt Verno, the principal sampling area, including localities 2 – 6, the vegetation zone of the deciduous oaks, the
Quercetalia pubescentis, range from 700 – 1000 m. From
850 – 1700 m the dominant tree species is Fagus sylvatica
of the Fagetalia zone (Fotiadis & al. 2005). In both zones
the woodlands have been cleared over large expanses giving room for montane grasslands used for the grazing of
livestock (Fig. 3). The lichens were sampled on granite
and gneiss outcrops and boulders in these grasslands (loc.
2, 3, 4, 5, 7, Fig. 2). On Mt Varnous (loc. 1) and on Mt
Verbista (loc. 7) the zonation is similar. At loc. 1 and 6 the
lichens were collected on gneiss outcrops and boulders in
404
Christensen: Lichens from granite and gneiss in mountains of Makedonia, Greece
Juniperus communis thickets. The environment of the localities is open, exposed to wind and sun (the grasslands,
Fig. 2), or semi-open (the Juniperus thickets). Juniperus
communis thickets are indicative of former grazing areas
– at least it is the case in N Europe outside fell-heaths and
other natural vegetation types. Thus, with the exception of
the outcrops and boulders, the biotopes are anthropogenic
in origin (Fig. 3).
Christensen & Svane (2009) reported a number of
species with northern affinities from mountains of N
Greece. Later Christensen & Alstrup (2013) published
species lists from different rock types from a number of
sites in the mountains of N Greece. They found that crystalline rocks at high altitude harbour a number of species
with boreal to Arctic and Alpine distribution. Allowing
for the few study sites in the present study, this seems
to confirm a connection between crystalline rocks, high
altitude and a host of species with a northern distribution
type. The outcrops and boulders of granite and gneiss
included in the present study were found at altitudes between 750 m and 1800 m. The three Umbilicaria species
(U. cylindrica, U. deusta and U. polyphylla) occurred
only above 1750 m, while Lasallia pustulata was found
as low as 750 m. Cetraria ericetorum and C. islandica
were not found below 1250 m, while C. aculeata was
found as low as 950 m (it occurs as low as sea level –
Linda in Arcadia 2013). Bryoria chalybeiformis was
found above 1250 m. Cornicularia normoerica occurred
above 1400 m. Pseudephebe pubescens was found above
1750 m. Ramalina capitata was found above 1400 m and
R. strepsilis above 1250 m. Brodoa oroarctica occurred
above 1750 m. Lecanora swartzii occurred at 1200 m.
The distribution of the above-mentioned species
along the altitude gradient represented by the seven
(eight) localities of the present study seems to mirror
the geographical distribution of the species: The species
found in the highest levels are those with the most pronounced bias toward north in their distribution patterns.
Of the two localities above 1750 m the Umbilicaria
species occurred on one (U. cylindrica, U. polyphylla) or
both (U. deusta) suggesting that the three species probably are very common on acidic rocks at this altitude.
Lasallia pustulata was found at all altitudes in four of the
seven localities and seems, therefore, to be rather common. Cornicularia normoerica was found at all four localities above 1400 m. It must be very common on acidic
rocks at this altitude. The three Cetraria species are very
sensitive to the high grazing intensity of these montane
grasslands. They were found as fragmented cushions
or as detached lobes patchily in the grass swards. The
species were, therefore, not an obvious element in the
grassland and could easily have been overseen in some of
the localities, as time did not allow for intensive searching. Bryoria chalybeiformis was found at two of the five
localities above 1250 m (or three of the six, if loc. 8 is
included). It is, therefore, likely to be rather common
on acidic rocks at this altitude. The ornithocoprophilous
character of Ramalina capitata and R. strepsilis makes it
difficult to evaluate their occurrences. Together they occur at four of the seven localities. The presence of Brodoa
oroarctica at the two very highest localities underlines its
Arctic-Alpine affinity. It may be relatively rare. The same
applies to Pseudephebe pubescens, which was found at
one of these two localities (above 1750 m).
The findings of this small study, with emphasis on
macrolichens from a small number of sites, may indicate that the northern element of the Greek lichen flora is
much more abundant than suggested from previous publications. Macrolichens of acidic crystalline rocks of the
high mountains are certainly under-collected in Greece.
This applies even more to crustose lichens.
Acknowledgements
Thanks are due to Helmut Mayrhofer (GZU) and another,
anonymous reviewer for constructive suggestions on an
earlier draft of this paper. Nicholas Turland kindly prepared the map.
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