Folia Cryptog. Estonica, Fasc. 32: 5–8 (1998)
A revision of Cladonia stricta
Teuvo Ahti
Department of Ecology and Systematics, P.O. Box 4, FIN-00047 University of Helsinki, Finland
Abstract: The arctic lichen Cladonia stricta (Nyl.) Nyl. is divided into three species, viz. C. stricta s. str., C. trassii Ahti, sp.
nov.., and C. uliginosa (Ahti) Ahti, comb. nov. Cladonia trassii and C. uliginosa have centrally proliferating scyphi and constantly
produce atranorin, while in C. stricta the scyphi are marginally proliferating and atranorin is inconstant. All the three species
are essentially arctic to subarctic, circumpolar, but the ranges are poorly known.
Kokkuvõte: T. Ahti. Ülevaade liigist Cladonia stricta.
Arktiline samblik Cladonia sticta (Nyl.) Nyl. jagatakse siin kolmeks liigiks: C. stricta s. str., C. trassii Ahti sp. nov. ja C. uliginosa
(Ahti) Ahti comb. nov. Liikidel C. trassii ja C. uliginosa on keskelt prolifitseeruvad karikad ning nad sisaldavad püsivalt atranoriini; liigi C. stricta karikad prolifitseeruvad servadest ning atranoriin kas esineb või puudub. Kõik kolm liiki on peamiselt
arktilise kuni subarktilise (tsirkumboreaalse) levikuga, siiski on nende areaalide ulatust seni vähe uuritud.
INTRODUCTION
Cladonia stricta (Nyl.) Nyl. was adopted by me
(Ahti, 1978) as the correct name for a widespread arctic lichen that was earlier called C.
lepidota “Nyl.”, C. gracilescens (Flörke) Vain. or
C. cerasphora Vain. The lectotype specimen of
C. stricta is from Taimyr Peninsula, Siberia. At
the same time I also somewhat provisionally
segregated C. stricta var. uliginosa Ahti for a
taxon which Vainio (e.g., 1894) had recognized
as as a distinct species under the misapplied
name C. gracilescens.
However, when outlining the distribution of
var. uliginosa I noted that the populations of C.
stricta in East Asia and western North America are in need of a more through taxonomic
analysis. I have later continued the study of C.
stricta in these regions. The final impetus for a
revision of my concept of C. stricta was provided
by Dr Taimi Piin, who pointed me out that the
type of C. stricta apparently represents a species
distinct from most of the material under this
name from Fennoscandia.
I had no possibility to make an extensive
revision of material in many herbaria, but a
new taxonomic scheme of the complex is given
below. It still does not include a treatment of
all variation noted in Canada and coastal East
Asia in this group.
Cladonia stricta (Nyl.) Nyl. (Figs. 1, 2)
Flora 52: 294. 1869. – Cladonia degenerans var.
stricta Nyl. in Middendorff, Reise Sibir. 4, Anh.
6(2): 4. 1867. Type: Russia. Krasnoyarsk Terr.:
Taimyr Peninsula, 1843, von Middendorff s.n.
(H-NYL 38841, lectotype, designated by Ahti,
1978: 11).
Cladonia degenerans f. fuscescens Nyl., Notiser Sällsk. Fauna Fl. Fenn. Förhandl. [Lich.
Lappon. Orient.] 8: 109. 1866 (preprint). Type:
Russia. Murmansk Region: island Bolshoy
Oleniy (“Olenji”), 1861, Fellman (H, lectotype,
here designated).
Cladonia cerasphora Vain., Acta Soc. Fauna
Fl. Fenn. 10: 167. 1894, nom. illeg. superfl. for
C. stricta (Ahti, 1978).
Illustration. Thomson (1984: 164).
Primary thallus evanescent, consisting of small
squamules, 1–2 mm wide. Podetia dark brown
to grey, at base strongly melanotic; 3–12 cm tall,
1.5–3 mm thick, slender, slightly branched by
dichotomy, axils closed or perforated; sterile tips
usually subsubulate, ascyphose; scyphi absent
or present, narrow (0.5–3 mm), often with 2–7
teeth or long proliferations at margins, shallow
to deep, frequently tips appearing trifurcate,
indistinctly scyphose; scyphi soon becoming
perforate, older scyphi with many perforations;
central proliferations in scyphi absent or very
rare; surface matt, clearly arachnoid, especially
towards the tips, verruculose-corticate, towards
the base checkered; podetial wall 200–300 mm,
cortex 50–70 mm, soft medulla 100–150 mm,
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Folia Cryptog. Estonica
Fig. 1. Cladonia stricta. Lectotype (H-NYL
38841). Scale = 1 cm.
stereome 50–100 mm, central canal densely
papillate. Conidiomata at tips of podetia, ovoid,
with hyaline slime, conidia 7–9 mm, falciform.
Hymenial discs infrequent, 2–5 mm wide, dark
brown, spores 10–15 × 2.5–3 mm, fusiform.
Chemistry. Fumarprotocetraric acid, often
with atranorin (e.g., in lectotype), traces of protocetraric and confumarprotocetraric acids. An
HPLC analysis cited under Cladonia phyllophora
Hoffm. by Huovinen et al. (1990: 223, no. 465).
No convirensic acid (Cph-1) has been detected
with TLC, though that substance is not uncommon in C. phyllophora.
Habitats. Observed in the field in Alaska,
the Yukon, and Sweden, and mostly seen growing in rock fields in timberline areas, either in
lower mountain tundra or in upper woodlands.
Judged from specimen labels probably mainly
in dryish upland communities rather than in
wet habitats.
Distribution. Essentially circumpolar, arctic-hemiarctic, extending to mountains in the
boreal zone. Recorded from Norway (incl. Svalbard), Sweden, Finland, Kola Peninsula, arctic
Siberia, Chukotka, Alaska, the Yukon, British
Columbia, Northwest Territories.
Representative specimens.
SVALBARD: Spitsbergen, ca. 10 km SE of Longyearbyen, 300–500 m, 1985 Kashiwadani 23537 (H,
TNS). NORWAY. Troms: Tromsö, 1906 Havås, Lich.
Exs. Norv. 528 (H), 1990 Schindler 14174 (H, KR).
SWEDEN. Jämtland: Åre, Mt. Åreskutan, 1280 m,
1975 Ahti 30303 (H). Torne Lappmark: Jukkasjärvi,
Fig. 2. Cladonia stricta. Canada, Ahti 31953
(H). Scale = 1 cm.
Riksgränsen, 1958 Bäck (H, UPS); Karesuando,
Laukkuåive, 1910 Montell (H); Kummaeno, 1910
Lynge (H). FINLAND. Lapponia enontekiensis: Enontekiö, Karesuvanto, 1867, Norrlin (H). RUSSIA. Murmansk Region: Lavna-tundra, 600 m, 1987 Dudoreva
298/5 (H, KPABG). Tyumen Region: Station 126 on
Vorkuta–Labytnangi railway, 1986, Zhurbenko 8618
(H). Krasnoyarsk Territory: Putorana, Lake Lama,
1984, Zhurbenko 848 (H); Severnaya Zemlya, SW of
Cape Barano, 1991, Safronova 913 & Andreev (H).
Magadan Region: Chukotka, Anyuy Range, 1978, Andreev (H). ALASKA. Point Barrow, 1958, Thomson et
al., Lich. Arct. 34 (US); Mt. Fairplay, 1967, Ahti 25270
(H). CANADA. British Columbia: km 108 on Haines
Highway, 1976, Otto 5897 (H). Yukon: Mile 96 Haines
Highway, 1967, Ahti 23356 (H). Northwest Territories:
District of Mackenzie, Hyndman Lake, 1966, Scotter
8045 (H), Nahanni Natl. Park, Hole-in-the-Wall Lake,
1200 m, 1977, Ahti 31953 (CANL, H, TU); District of
Keewatin, Parson’s Lake, 1959, Thomson & Larsen
5883 (US).
Cladonia stricta s. str. is the “apparently new species” mentioned under C. phyllophora Hoffm. by
Thomson & Ahti (1994: 146) and in connection
with C. alaskana A. Evans by Ahti & Zhurbenko
(1994). Perhaps most of the material of C. stricta
is found under C. phyllophora in herbaria. It
is noteworthy that C. stricta occasionally contains atranorin and such specimens have often
been referred to C. stricta while morphologically
similar specimens without atranorin have been
placed in C. phyllophora.
7
In specimens with well-developed scyphi the
numerous perforations of the scyphi constitute
a good diagnostic character.
del Fuego, Argentina (Stenroos & Ahti, 1991:
324, as C. stricta).
Representative specimens.
Cladonia trassii Ahti sp. nov.
Thallus primarius persistens aut demum evanescens, squamis 5–12 mm longis. Podetia 3–8
cm alta, elongata, irregularia, parce ramosa, ascypha vel scyphis angustis, a centro scyphorum
prolifera, glaucescentia, cortice subcontinuo,
apice pruinoso, subtomentoso, basi areolato,
medulla melanotica. Atranorinam et acidum
fumarprotocetraricum continens.
Type. Sweden. Torne Lappmark: Gällivare
(“Gellivare”), Mt. Patjanen, 550 m, 1922, Stenholm, Sandstede: Clad. Exs. 1134 (H, holo-type;
TU, isotype; as Cladonia gracilescens).
Illustration. Krog et al. (1994: 169, as Cladonia stricta).
Etymology. Named in honour of the Estonian
lichenologist Professor Hans Trass.
Primary thallus persistent to evanescent, consistings of fairly large squamules, 1–5 mm wide.
Podetia 3–8 cm tall, 1–3 mm thick, extremely
variable in shape; glaucous-grey, often browned
in part, base melanotic, with checkered surfafe;
usually slightly branched by dichotomy, axils
closed; scyphose but scyphi usually very few
and irregularly shaped, if well-developed, then
centrally proliferating; ascyphose tips bluntish;
surface areolate-corticate, rather smooth but often largely decorticate and squamulose, slightly
arachnoid and pruinose towards the tips; podetial wall 200–380 mm, cortex 20–40 mm,
medulla 80–150 mm, stereome 100–190 mm;
central canal slightly papillate. Conidiomata at
tips of podetia, globose to mammiform, pycnidial
slime hyaline, conidia 6–8 × 1 mm, falciform.
Hymenial discs infrequent, dark brown, 2–5 mm
wide, spores 12–14 × 2.5–3 mm, fusiform.
Chemistry. PD+ red, K+ yellow; contains
atranorin, fumarprotocetraric acid and traces
of protocetraric and confumarprotocetraric
acids.
Habitats. Mainly in arctic and alpine late
snow-lie communities.
Distribution. Arctic-alpine, circumpolar in
the northern hemisphere, probably somewhat
oceanic. A single locality reported from Tierra
SWEDEN. Jämtland: Åre, Mt. Snasahögarna, Silverfallet, 900 m, 1975 Ahti 30313 (H, UPS). RUSSIA.
Khabarovsk Territory: Anoy River, 1956, Rosenberg 44
(US). ALASKA. Thompson Pass, 1967, Thomson 20974
& Ahti (US). CANADA. Northwest Territories: Baffin
Island, Clyde Fjord, 1950, Hale 301 (US). Québec:
Mt. Albert, 1940, Lepage 6287 (US). GREENLAND.
Hansen: Lich. Groenl. Exs. 92 (H, US).
Cladonia trassii is actually a new name for what
Vainio (1894, 1922) called C. cerasphora and
what has recently been called C. stricta var.
stricta (e.g., Ahti, 1977, 1978; Krog et al., 1994),
though some material of C. stricta s. str. has
been included in the previous descriptions.
Cladonia uliginosa (Ahti) Ahti comb. nov.
Cladonia stricta var. uliginosa Ahti, Ann. Bot.
Fennici 15: 11: 1978. Type: Russia. Murmansk
Region (formerly Finland. Kuusamo: Salla):
Korja, “ad pedem montis Nurmitunturi”, 1937,
Laurila, Räsänen: Lich. Fenn. Exs. 475 (H, holotype; BM, H, UPS, isotypes).
Illustration. Krog et al. (1994: fig. VII, as
Cladonia stricta, in colour!).
For description see Vainio (1922), as Cladonia
gracilescens.
Chemistry. As in C. trassii, but with additional minor unknown substances (Huovinen
et al., 1990: 225).
Habitat. In periodically wet places in woodlands near timberline.
Distribution. Not uncommon in northern
boreal forest regions of Fennoscandia but its
total distribution is little known. There are few
records from North America. In Finland it is
commoner than C. trassii but in Sweden and
Norway it seems to be less common common
than that species.
Representative specimens.
NORWAY. Hordaland: Granvin, 225 m, 1918, Havås,
Lich. Norv. Occid. Exs. 101 (H). SWEDEN. Torne
Lappmark: Karesuando, Virkakursu, leg. Lång,
Crypt. Exs. Vindob. 1867 (H). FINLAND. Ostrobottnia
ultima: Simo, 1942, Räsänen, Lich. Fenn. Exs. 790
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Folia Cryptog. Estonica
(H). RUSSIA. See type. ALASKA. Alaska Peninsula,
Naknek, 1948, Lepage 22634 (US); Lake Peters, 1948,
Scholander & Flagg (US).
Cladonia uliginosa is characterized by constant
occurrence of distinct, centrally proliferating
scyphi and numerous podetial squamules. It is
very close to C. trassii and I earlier (Ahti, 1978)
treated it as a variety (under C. stricta). I have
not found distinct intermediates, but further
field investigations are required to confirm the
differences in relation to C. trassii. The latter
species usually has no or no well-developed
scyphi. Many of the North European exsiccata
distributed as C. gracilescens belong to C. trassii
rather than C. uliginosa, which has confused the
recognition of these species.
ACKNOWLEDGMENTS
This contribution is dedicated to Prof. Hans
Trass who, among many other things, has studied European and Siberian arctic macrolichens.
I am further indebted to Dr Taimi Piin (Tallinn),
Dr Mikhail P. Zhurbenko (St. Petersburg), and
Ms. Tamara Dudoreva (Kirovsk), Prof. Henry A.
Imshaug (East Lansing, MI), who have given me
material or observations on C. stricta. Dr George
W. Scotter and Prof. John W. Thomson provided
me opportunities to study this group in field in
Canada and Alaska. The study was financially
supported through a grant from the Academy
of Finland, and at the last stage through an
Andrew W. Mellon Senior Fellowship.
REFERENCES
Ahti, T. 1977. Cladonia Wigg. In Bestimmungsschlüssel europäischer Flechten. Ergänzungsheft
(eds. Poelt, J. & Vêzda, A.) 1: 45–95. J. Cramer,
Vaduz.
Ahti, T. 1978. Nomenclatural and taxonomic remarks
on European species of Cladonia. Ann. Bot. Fenn.
15: 7–14.
Ahti, T. & Zhurbenko, M. P. 1994. Distribution of
Cladonia alaskana in Asia. Graphis Scripta 6:
11–14.
Krog, H., Østhagen, H. & Tønsberg, T. 1994. Lavflora.
Norske busk- og bladlav. Edn. 2. Universitets-forlaget. 368 pp.
Huovinen, K., Ahti, T. & Stenroos, S. 1990. The composition and contents of aromatic lichen substances
in Cladonia section Cladonia and group Furcatae.
Bibl. Lichenol. 38: 209–241.
Stenroos, S. & Ahti, T. 1991 (‘1990’). The lichen family
Cladoniaceae in Tierra del Fuego: problematic or
otherwise noteworthy taxa. Ann. Bot. Fenn. 27:
317–327.
Thomson, J. W. 1984. American arctic lichens. Vol.
1. The macrolichens. Columbia University Press,
New York.
Thomson, J. W. & Ahti, T. 1994. Lichens collected
on an Alaska Highway expedition in Alaska and
Canada. Bryologist 97: 138–157.
Vainio, E. A. 1894. Monographia Cladoniarum universalis. Vol. 2. Acta Soc. Fauna Fl. Fenn. 10:
1–498.
Vainio, E. A. 1922. Lichenographia fennica. Vol. 2.
Baeomyceae et Lecideales. Acta Soc. Fauna Fl.
Fenn. 53: 1–340.
Folia Cryptog. Estonica, Fasc. 32: 9–14 (1998)
Opegrapha trassii sp. nov., a new lichenicolous fungus on
Heterodermia
Brian J. Coppins* and Sergey Y. Kondratyuk#
*Royal Botanic Garden Edinburgh, Inverleith Row, Edinburgh EH3 5LR, UK
#N. G.Kholodny Institute of Botany, 2 Tereshchenkivska St., 252601 Kiev, Ukraine
Abstract: A new lichenicolous fungus, Opegrapha trassii S. Kondratyuk & Coppins (Opegraphaceae), parasitizing several species
of Heterodermia (Physciaceae) is described and illustrated, and comments about its differences from related taxa are provided.
The new fungus is widely distributed and so far known from Sierra Leone, Uganda, Zambia, Malawi, Angola, South Africa,
India, Mauritius, Japan, New Zealand, USA (incl. Hawaii Islands and Puerto Rico) and Surinam.
Kokkuvõte: B. J. Coppins ja S. J. Kondratjuk. Opegrapha trassii sp. nov., uus lihheniseerunud seene liik
sambliku Heterodermia tallustelt.
Kirjeldatakse uus lihheniseerunud seene liik Opegrapha trassii S. Kondratyuk & Coppins (Opegraphaceae), mis parasiteerib
sambliku Heterodermia (Physciaceae) tallustel. Lisatud on ka uue liigi erinevused lähedastest taksonitest. O. trassii on laialt levinud,
uuritud eksemplaride leiukohtadena on märgitud Sierra Leone, Uganda, Sambia, Malaavi, Angoola, Lõuna-Aafrika, India,
Mauritius, Jaapan, Uus-Meremaa, USA (s.h. Havai ja Puerto-Rico) ning Surinam.
INTRODUCTION
The ascomycete genus Opegrapha comprises c.
300 species, mostly lichenized species (Hawksworth et al., 1995). However, some of the species are lichenicolous, several being previously
assigned to the genus Leciographa A. Massal.
In their world key to lichenicolous fungi, Clauzade et al. (1989) recognized about 20 species of
lichenicolous Opegrapha, but since then a further 24 species have been newly described, validated or introduced (or re-introduced) into the
genus: O. stereocaulicola Alstrup & D. Hawksw.
on Stereocaulon (Alstrup & Hawksworth, 1990),
O. physciaria (Nyl.) D. Hawksw. & Coppins on
Xanthoria parietina (Atienza, 1992; Coppins et
al., 1992); O. sphaerophoricola Isbrand & Alstrup
on Sphaerophorus (Isbrand & Alstrup, 1992); O.
geographicola (Arnold) Hafellner on Rhizocarpon
geographicum, O. rotunda Hafellner on Physconia and O. zwackhii (A. Massal. ex Zwackh)
Källsten on Phlyctis argena (Hafellner, 1994);
O. brevissima Kalb & Hafellner on Haematomma
hilare (Kalb et al., 1995), O. leuckertii S. Kondratyuk & D.J. Galloway on Pseudocyphellaria
and Sticta (Kondratyuk & Galloway, 1995), O.
physcidiae Kalb & Elix on Physcidia sp. (Kalb &
Elix, 1995), O. rouxiana Nav.-Ros. & Hladun on
Polyblastia (Navarro-Rosinés & Hladun, 1995),
O. cryptotheciae Matzer on Cryptothelia, O.
ectolechiaceaum Matzer & R. Sant. on foliicolous
Ectolechiaceae, O. epiporina Matzer on Porina, O.
kalbii Matzer on Byssoloma, O. mazosiae Matzer
on Mazosia, O. phyllobathelii Matzer & R. Sant.
on Phyllobathelium, O. phylloporinae Müll. Arg.
and O. sipmannii Matzer on Porina epiphylla gr.,
O. porinicola Matzer on Phyllophiale alba and
Porina epiphylla, O. strigula Matzer & R. Sant.
and O. uniseptata Matzer on foliicolous Strigula
(Matzer, 1996); O. phlyctidicola (Vouaux) Etayo
on Phlyctis agelaea (Etayo, 1996); O. encephalographoidea Diederich & Aptroot on Pyrenula,
and O. pleactocarpoidea Diederich on Phaeographis (Aptroot et al., 1997). Furthermore, we are
aware of several additional, undescribed species,
one of which is widely occurring on various species of Heterodermia (Physciaceae).
Recent, more critical morphological studies
of lichenicolous Opegrapha have shown each
species to have a restricted host range, most
being either species- or genus-specific, or more
rarely family-specific (e.g. O. ectolechiacearum
on foliicolous Ectolechiaceae and O. rupestris
Pers. on crustose, foveolate Verrucariaceae on
limestones), and no species has yet been reliably
reported from hosts of different families. No
Opegrapha or ‘Leciographa’ has hitherto been
reported on Heterodermia, and the fungus on
10
Folia Cryptog. Estonica
this genus clearly differs from other Opegrapha
species on hosts belonging to the Physciaceae.
Accordingly, the collections on Heterodermia
are here referred to the species newly described
below.
Type: [SOUTH AFRICA]: B. G. Hope, on Heterodermia sp., date and collector unmentioned
[Herb. Archibald Menzies (1754–1842)] (BM –
holotype; E – isotype [South Africa: Cape of Good
Hope, on H. speciosa, 1791, A. Menzies]).
Opegrapha trassii S. Kondratyuk & Coppins sp. nov.
Etymology: Opegrapha trassii is named after the
famous Estonian lichenologist Prof. Hans Trass
in recognition of his important contribution to
our knowledge on the flora of Estonia and other
regions of Eurasia, as well as on the family Physciaceae, especially the genus Heterodermia.
Fungus lichenicola in thallo specierum Heterodermiae; cortex hospitis brunneolescens.
Ascomata lirelliformia, aggregata, simplicia
vel interdum breviter furcata, epruinosa, 200–
300(–420) µm longa, 125–160(–200) µm lata,
disco rimiformi vel demum hianti. Excipulum
integrum, fuscoatrum, admodum K-. Hymenium
(40–)45–63.5(–72.5) µm altum, I+ caeruleum.
Asci clavati cum annulo apicali amyloideo
minuto, (30.5–)32.5–40(–47) × (11–)12–14.5(–16)
µm, (2–)4–6(–8)-spori. Ascosporae 3(–4–5)-septatae, hyalinae vel demum granulis brunneascentes, 12.5–17.5(–19) × (3.6–)4.5–5(–5.5) µm.
Pycnidia pauca, ±immersa, c. 40–50 µm diam.,
conidiis bacilliformibus 3.8–5 × 0.5–0.7 µm.
Lichenicolous fungus, parasitic on thalli (rarely
on thalline margin of host apothecia) on various Heterodermia species. Ascomata arising,
±superficially (i.e. not developing below, and
bursting through, the host cortex), in clusters up to 3.5 mm across and containing up
to 15(–38) ascomata; surrounding host tissue
discoloured brown; clusters characteristically
elliptical, with the long axis of the clusters and
most individual ascomata parallel with that
of the lobes of the host. Individual ascomata
Fig. 1 a–b. Opegrapha trassii. a: habitus of ascomata on the host thalli (× 72); b: part of ascomata
(× 48).
11
Fig. 2 a–b. Opegrapha trassii. a: asci, b: ascospores. Scale 7 µm.
black, epruinose, 200–300(–420) µm long,
125–160(–200) µm wide, and, in sections,
(127–)140–150 µm tall; simple or occasionally
shortly 1–2-furcate, straight or slightly curved,
disc slit-like or eventually gaping. Exciple well
developed, entire, 18–27(–32.5) µm wide laterally,
(21.5–)30–43(–53) µm wide below, dark brown to
blackish brown, sometimes with reddish tinge
K- or K+ fuscous brown (loosing red tinge) or
slightly olivaceous (never bright green or redbrown). Hymenium hyaline, (40–)45–63.5(–72.5)
µm tall, I+ blue, K/I+ blue; epihymenium absent,
with the excipulum remaining incurved over the
top of the hymenium or becoming evident in
widely gaping ascomata and then hyaline to
pale brown, 9.0–21.5 µm tall. Hypohymenium
dilute brownish, K- or K+ slightly olivaceous,
(12.5–)16–23.5 µm tall. Paraphyses branched,
1–1.5(–2) µm wide (in KOH). Asci broadly
clavate, (30.5–)32.5–40(–47) × (11–)12–14.5(–16)
µm, with K/I+ blue ring around the top of the
ocular chamber, (2–)4–6(–8)-spored. Ascospores
hyaline, 3(–4–5)septate, upper 2 (3 or 4) cells
somewhat more swollen than the lower cells,
12.5–17.5(–19) × (3.6–)4.5–5(–5.5) µm, apices
obtuse, sometimes with a thin perispore <1
µm thick; old spores released from asci sometimes pale brown due the external deposition of
minute brown granules. Pycnidia sometimes
present around the outside of the ascomatal
clusters, immersed in the host tissue, dark
brown to black, c. 40–60 µm diam.; wall tissue
dark brown, K-; conidia (microconidia) bacilliform, 3.8–5 × 0.5–0.7 µm (Fig. 1, 2).
Opegrapha trassii is very variable with respect
to the numbers of spores in the ascus; this can
be 2, 4, 6 or 8, and all four types have been seen
in sections of a single ascoma. However, asci
with 4 or 6 spores are those most commonly
encountered.
Only two species of Opegrapha have been previously described from hosts in the Physciaceae,
and both differ from O. trassii in having rounded
ascomata and larger ascospores: O. rinodinae
Vêzda on Phaeorrhiza nimbosa (ascospores 22–
26 × 4–6 µm) and O. rotunda Hafellner on Physconia (ascospores 18–22 × 5–6 µm) (Hafellner,
1994). Despite its specific epithet, O. physciaria
(Nyl.) D. Hawksw. & Coppins grows on Xanthoria parietina (Teloschistaceae) and has 8-spored
asci that lack an apical K/I+ blue ring. Two collections in E have an unidentified Opegrapha
on Pyxine cocoes (Physciaceae) from Botswana
12
Folia Cryptog. Estonica
Fig. 3 Distribution of Opegrapha trassii.
(Long 12449, Long & Rae 881). They differ from
O. trassii in that the clusters of ascomata arise
below and eventually burst through the cortex
of the host, the asci are 8-spored, and the
ascospores are somewhat broader, with many
reaching a width of 5.5–6 µm.
What seems to be an additional Opegrapha species on Heterodermia, has recently been found. A
small specimen, collected in Papua New Guinea
and sent to the first author by Dr P. Diederich,
differs from O. trassii in having more robust
ascomata with a soon expanding disc, to 460 µm
wide. In sections the ascomata are much taller
(to 280 µm) with a taller hymenium (80–85 µm)
and much taller combined hypothecium and
lower exciple (to 200 µm; mostly less than 60
µm in O. trassii). The asci are 4-spored, but the
ascospores are consistently 5-septate at maturity (no mature 3-septate ascospores being
detected), and longer (17.5–21.5 × 4.5–6,5 µm).
It seems unlikely that this collection (Diederich
10317) represents an extreme morphotype of O.
trassii, and further collections will surely prove
it to be a distinct species.
Additional specimens (all on Heterodermia spp.)
(Fig. 3). INDIA: Madras: Nilgias district, 3000 ft, on
H. leucomela, November 1886 J.S.Gamble 18438 (BM);
Madras, on H. leucomela, 1775 (collector unknown)
[Herb. Ind. Oc. Hook fil. & Thomson] (BM).
MAURITIUS: on H. obscurata, no date, Dr Ayres (E).
JAPAN: Shikoku, Pref. Ehime, Odamachi, Ochira ca.
560 m alt., on rock, on H. obscurata, 18 Nov. 1972 M.
Inoue 2227 (BM). SIERRA LEONE: Loma Mountains,
Camp 2, on tree at edge of forest gallery at about
1400 m alt., on Anaptychia boryi [=H. leucomela subsp.
boryi], 15 March 1971 P. W. Richards R7253L (BM).
UGANDA: Toro, Burahya, Street in Fort Portal, Lat.
0º40’N, Long. 30º16’E, alt. 1500 m, SR-95-75, trunk
of tree in avenue, on H. leucomela, Juny 1970 T.D.V.
Swinscow 2U 17/18-2 (BM; 1 km N of Fort Portal.
Toro. Common on tree bole in avenue, on H. leucomela, 25.07.1971 A.Pentecost RE29 (BM). ZAMBIA [N.
Rhodesia], Namwala, Kafue National Park, Lubalansuki Hill, Bald granite Hill, on H. pseudo-speciosa, 7
May 1964 Mitchell (BM). MALAWI: Mlanje, S.Province,
D.C.’s grounds, alt. 3200 ft, on H. magel-lanica, April
1969 M. Jellicoe (BM). ANGOLA: Welwitsch, Iter Angolense, No. 34. Hab. frequens in summis jugis de
Serra de Etella et circa Mumpulla ad 4500 ped. elev.,
truncis Iarchonanthi spec. adnascens. Distr. Huilla
(3800–5500 ped. elev.). Inter 14 et 16o Lat austr.,
on Anaptychia neoleuco-melaena f. squarrosa [=H.
leucomela subsp. boryi], Jun. 1860 L. Welwitsch (BM
– isotype of A. neo-leucomelaena f. squarrosa). SOUTH
AFRICA: Cape Province, E. prov, on a small dead bush,
on H. leuco-mela, 15 March 1898 R.E.Raud (BM); Cape
of Good Hope, on H. lepidota, date and collector unmentioned (BM, from Herb. A.Menzies. Recd. 1886).
Natal: Umgae Mountains, SW Plant, on H. sp. (‘Physcia
leucomelas v. subcomosa [ Syn. p. 415]’), 1853 (Collector uncertain) (BM). USA: Louisiana: East Feliciana
Parish, 5.6 km SE of Clinton, Idlewild Research Station, 30o48’N, 90o45’W, on H. albicans, 18 April 1984,
S.C.Tucker 26346 (E); Puerto-Rico,
13
Coamo, on H. sp., 13 December 1885 P. Sintenis (BM
ex K sub Parmotrema tinctorum); Hawaii, Sandwich
Islands, on H. sp. [without date and collector] [Herb.
A. Menzies. Recd. 1886] (BM). SURINAM: Paramaribo,
Palmgarden, 0 m alt., on H. albicans on Roystonea
regia, 25–29 January 1985, A. Aptroot 14842a (Herb.
Aptroot). NEW ZEALAND: loc. uncertain, on H. speciosa [date unknown], Knight [Herb. Leighton] (BM).
North Island, Mantime rocks, Russell, on Anaptychia
obscurata, 9.3.1966 Wade (BM); North Island, coast
rocks, Mount. Maunganui, on H. obs-curata, 24–26
May 1966, Wade (BM).
ACKNOWLEDGEMENTS
We are grateful to the curators and keepers of
the cited herbaria for assistance with loan of
specimens, Dr A. Aptroot for sending his collection of O. trassii from Surinam, and to Dr P.
Diederich for sending us material of the probable second Opegrapha species on Heterodermia.
K.S.Y. is also thankful to Ms A.M.Coppins and to
the first author for extremely warm hospitality
and generous help with accommodation during his stay in Edinburgh. He acknowledges the
invaluable help given him by staff at E, during
his stay there in 1996, and is grateful to the
Royal Society for financial support too.
REFERENCES
Alstrup, V. & Hawksworth, D. L. 1990. The lichenicolous fungi of Greenland. Medd. Grønland, Bioscience 31: 1–90.
Aptroot, A., Diederich, P., Sérusiaux, E. & Sip-man, H.
1997. Lichens and lichenicolous fungi from New
Guinea. Bibl. Lichenol. 64: 1–220.
Atienza, V. 1992. Peridiothelia oleae (Körber) D.
Hawksw. and Opegrapha physciaria (Nyl.) D.
Hawskw. et Coppins, two poorly known west
mediterranean fungal taxa. Anales Jard. Bot.
Madrid. 50: 159–162.
Clauzade, G., Diederich, P. & Roux, C. 1989. Nelikenigintaj fungoj likenlogaj. Bulletin Soc. Linn. Provence,
Numéro spécial 1: 1–142.
Coppins, B. J., James, P. W. & Hawksworth, D. L.
1992. New species and combination in “The lichen
Flora of Great Britain and Ireland”. Lichenologist
24: 351–370.
Etayo, J. 1996. Aportación a la flora liquénica de las
Islas Canarias. I. Hongos liguenícolas de gomera.
Bull. Soc. Linn. Provence 47: 93–110.
Hafellner, J. 1994. Beitrage zu einem Prodromus der
lichenicolen Pilze Osterreichs und angrenzender
Gebiete. I. Einige neue oder seltene Arten. Herzogia 10: 1–28.
Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler,
D. N. 1995. Ainsworth & Bisby’s Dictionary of the
Fungi. 8th edn. Wallingford: CAB International.
Isbrand, S. & Alstrup, V. 1992. Opegrapha sphaerophoricola sp. nov. Bryologist 95: 233–234.
Kalb, K. & Elix, J. 1995. The lichen genus Physcidia.
Biblioth. Lichenol. 57: 265–296.
Kalb, K. Hafellner, J. & Staiger, B. 1995. Haematomma-Studien. II. Lichenicole Pilze auf arten der
Flechtengattung Haematomma. Biblioth. Lichenol.
59: 199–222.
Kondratyuk, S. & Galloway, D. J. 1994. Liche-nicolous
fungi and chemical patterns in Pseudo-cyphellaria. Bibl. Lichenol. 57: 327–345.
Matzer, M. 1996. Lichenicolous ascomycetes with
fissitunicate asci on foliicolous lichens. Mycol.
Papers 171: i–x, 1–202.
Navarro-Rosinés, P. & Hladun, N. L. 1995. Opegrapha
rouxiana sp. nov., nelikeniginta fungo likenloga
ce enpetraj Polyblastia. Bull. Soc. linn. Provence
46: 25–90.
14
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 15–20 (1998)
A revision of the Lecanora subfusca group in Estonia
Inga Jüriado
Institute of Botany and Ecology , University of Tartu, 38 Lai St., EE2400 Tartu, Estonia
Abstract: 12 species of the Lecanora subfusca group are known in Estonia, 10 of them are present in the contemporary
flora, one species is considered extinct, and one doubtful. Lecanora circumborealis is new for the country, four species are very
rare (L. caesiosora, L. epibryon, L. glabrata, L. impudens) and seven species are common in Estonia. A key to the species of the
L. subfusca group in Estonia is presented.
Kokkuvõte: I. Jüriado. Ülevaade Lecanora subfusca rühmast Eestis.
Eestist on teada 12 samblikuliiki, mis kuuluvad Lecanora subfusca rühma, neist 10 on kindlasti esindatud Eesti kaasaegses flooras,
üks liik on tõenäoliselt hävinud ja üks – kaheldav. Lecanora circumborealis on uus liik Eesti lihhenoflooras; neli liiki on väga
haruldased (L. caesiosora, L. epibryon, L. glabrata, L. impudens) ja seitse liiki on üsna või väga tavalised Eestis. Käesolevas töös
on esitatud nende liikide määramistabel ja üldine levik Eestis. Tutvustatud on ka liikide määramiseks vajalikke anatoomilisi
tunnuseid.
INTRODUCTION
The name “Lecanora subfusca” has been used by
many authors as a collective name for a group
of epiphytic and epilithic species, many of which
are widely distributed in the Northern Hemisphere. The combination Lecanora subfusca (L.)
Ach. was proposed by Acharius in 1810, who
divided the species into eight varieties, seven of
which were later raised to the specific level. Today the name L. subfusca is out of use according
to the proposal of Vitikainen & Brodo (1985) to
include it in the list of rejected names.
The characteristic features of this group
are:
* ellipsoid to broadly ellipsoid ascospores (approximately 10–20 × 6–9 mm),
* apothecial discs are generally reddish
brown,
* apothecial margin (amphithecium) contains
calcium oxalate crystals,
* thallus and apothecial margin react K + yellow
(atranorin),
* thallus is crustose, ± grey (Brodo, 1984; Eigler,
1969).
The aim of this study was to revise the L. subfusca group in Estonia. A list of species, data on
their distribution and some ecological notes are
presented. This study is part of a larger project
to compile a new checklist of Estonian lichens.
As morphological features in the L. subfusca
group are very variable and overlapping, it is
necessary to study apothecial anatomy while
identifying the species. Therefore some aspects
of apothecial anatomy (epihymenium and amphithecium types) in this group are briefly discussed following Brodo (1984).
MATERIALS AND METHODS
The study was based on collections in the
herbarium materials of the Institute of Botany
and Ecology at the University of Tartu (TU); the
specimens collected in Estonia and kept in the
Universities of Helsinki (H) and Riga (RIG) were
also considered. Altogether about 1100 specimens were studied. Additional material for
comparison was received from Helsinki (H). Besides the herbarium materials, publications by
Bruttan (1870), Mereschkowski (1913), Räsänen
(1931), Trass (1967, 1970) were taken into account when composing the list of species.
Cross-sections of apothecia were made by
hand and examined by light microscope (MBI
- 3). Spot tests were made with 10 % potassium hydroxide (K) and paraphenylenediamine
in ethanol (Pd); K and 50 % nitric acid (HNO3)
were used to determine the solubility of epihymenial granules, HNO3 was used also for hymenial colour tests. The secondary substances
were identified using thin-layer chromatography
(TLC) described by Culberson & Kristinsson
(1970) and Culberson (1972) only in a few occasions – when inevitable for the identification
of species (in 31 specimens).
16
Folia Cryptog. Estonica
The herbarium data were sorted using the
computer program BRAHMS and the program
DMAP was used to compose the distribution
map in Estonia.
The figures 1 and 2 (from Brodo, 1988,
drawn by Susan Laurie-Bourque) are reproduced with permission from the Canadian
Museum of Nature, Ottawa, Canada.
The following key books and papers were
used to identify the specimens and to compose
the key: Brodo (1984), Brodo & Vitikainen
(1984), Brodo et al. (1994), Foucard (1990), Poelt
& Vêzda (1981), Purvis et al. (1992), Tønsberg
(1992), Wirth (1995).
RESULTS AND DISCUSSION
Important apothecial characters in the Lecanora subfusca group
The anatomy of the L. subfusca group has been
studied in detail by different authors (Brodo,
1984; Eigler, 1969; Magnusson, 1932; Poelt,
1952). Several types of apothecia have been
described according to the structure of the epihymenium and amphithecium. Hereafter only
some of those epihymenium types – occuring in
Estonian species - will be discussed.
The epihymenium is the upper layer of the
hymenium that is differentiated by pigmentation
(within or between the tips of the paraphyses)
and granulation (above or between the tips of
the paraphyses) (Brodo, 1984).
Epihymenium types (according to Brodo,
1984):
1. chlarotera-type – epihymenium coarsely
granular, granules are on the surface of the
hymenium, soluble in KOH and HNO3; not
pigmented or pigmented reddish brown to
olivaceous brown (pigmentation disappears
in KOH, in dark apothecia epihymenium
HNO3 + reddish) (Fig. 1);
2. pulicaris-type – epihymenium finely granular (granules are much smaller than in
chlarotera-type epihymenium), located on
the surface of the hymenium and between
the tips of paraphyses (up to 20 mm in
depth), soluble in KOH but insoluble in
HNO 3 ; pigmented brown to olivaceous
brown (pigmentation disappears in KOH,
in dark apothecia epihymenium HNO3 +
reddish) (Fig. 1);
Fig. 1. Epihymenial granules: (a) chloroteratype, (b) pulicaris-type in L. hybocarpa. Scale:
each unit = 10 µm.
3. glabrata-type – epihymenium not granular
or inspersed, pigmented clear reddish brown
(pigmentation persists in KOH).
The amphithecium has been considered to be
all those tissues external to the hypothecium
and hymenium, and includes also the cortex
(Brodo, 1984).
Three amphithecium types can be distinguished in the group (according to Brodo,
1984):
1. pulicaris-type – very large, irregular crystals in clumps in amphithecial medulla
(Fig. 2);
2. campestris-type – small, irregular crystals
entirely or partially filling amphithecial medulla, not entering the cortex;
3. allophana-type – small, irregular crystals
extending from the medulla into the cortex
(Fig. 2).
In some cases the amphithecial crystals may be
rather few or even absent, but generally they are
abundant. The pulicaris-type is usually easily
recognized while the differences between the
campestris- and allophana-types are less obvious. Combining the epihymenium and amphithecium types, the species can be divided into
groups (Table 1).
17
Fig. 2. Amphithecium types:(a) pulicaris-type in L. rugosella, (b) allophana-type in L. glabrata.
Scale: each unit = 100 µm.
Table 1. The epihymenium and amphithecium types in L. subfusca group in Estonia
Epihymenium glabrata-t.
Epihym. chlarotera-t.
Epihym. pulicaris-t.
Amphit. with small
crystals (allophana- or
campestris-t. [*])
Amphit. with large
crystals (pulicaris-t.)
Amphit. with large
crystals (pulicaris-t.)
Amphit. with large
crystals (pulicaris-t.)
L. allophana
L. campestris *
L. epibryon
L. glabrata
L. impudens
L. argentata
L. caesiosora
L. cenisia
L . chlarotera
L. rugosella
L. circumborealis
L. pulicaris
List of species of the Lecanora subfusca
group in Estonia
Ten species are present in the contemporary
flora, one species (marked with asterisk *) is
considered extinct, and one (marked with ?) is
doubtful. Only those synonyms in the Lecanora
subfusca group are listed that are mentioned
in the checklist of Estonian lichens by Trass
(1970).
LECANORA ALLOPHANA Nyl. – very widely spread all
over the country (more than 200 localities),
common in parks in the countryside and
also in towns. On deciduous trees (Acer,
Fraxinus, Populus, Tilia), rarely on wood.
L. ALLOPHANA f. SOREDIATA Nyl. – rare in Estonia,
four specimens from the western, northern
and southern parts of Estonia, on Populus
tremula (TLC: atranorin, terpenoids).
L. ARGENTATA (Ach.) Malme (syn. L. subfuscata H.
Magn., L. subrugosa Nyl.) – very common
in wooded meadows and deciduous forests
in western regions, frequent also in other
parts of Estonia. On deciduous trees (Acer,
Fraxinus, Populus, Quercus, Sorbus, Tilia),
rarely on wood.
L. CAESIOSORA Poelt – very rare, only one locality
in western Estonia: Hiiumaa Co., Saarnaki
Is., on stone fence, 16 Aug. 1974, leg. E.
Sander, det. I. Brodo (TLC 1120-16 by Brodo
1993: atranorin, chloroatranorin, roccellic
acid) (TU) (Jüriado, 1997).
L. CAMPESTRIS (Schaer.) Hue – quite frequent in
western and northern Estonia, only one
specimen has been collected at an inland
location (from southern Estonia). On calcareous and siliceous rocks, also on manmade substrata.
18
Folia Cryptog. Estonica
L. CENISIA Ach. [syn. L. atrynea (Ach.) Nyl.] – quite
common on the coast and islands in western and northern Estonia, sparse inland. On
siliceous rocks, sometimes also on wood.
L. CHLAROTERA Nyl. – very common all over the
country (more than 100 localities), in forests and parks in the countryside and in
towns. On deciduous trees, also on wood
(14 specimens have been studied by TLC:
atranorin, gangaleoidin, ± norgangaleoidin,
± roccellic acid).
L. CIRCUMBOREALIS Brodo & Vitik. – new for Estonia, rare – five localities distributed in
different parts of Estonia (Fig. 3).
Specimens examined. 1. Harjumaa Co., near Vasalemma, on Betula pubescens, 15 June 1947, leg. E.
Parmasto, verified by O. Vitikainen (TLC: atranorin,
roccellic acid) (TU). 2. Harjumaa Co., near Maardu, on
Betula sp, 23 May 1942, leg. J. Seim (TLC: atranorin)
(TU). 3. Läänemaa Co., Vormsi Is., Diby, on wood,
July 1987, leg. R. Allmäe (TLC: atranorin) (TU). 4.
Valgamaa Co., Kääriku, on Betula pendula, 27 March
1966, leg. H. Trass (TLC: atranorin, roccellic acid) (TU).
5. Viljandimaa Co., Vihi, on wood, 30 July 1996, leg.
I. Jüriado (TLC: atranorin) (TU).
available and, therefore, it is not possible to
verify the taxon. According to the literature
(Santesson, 1993), the species occurs
mainly on Fagus, which does not occur in
Estonia. Today we still include the species
in the checklist of Estonian lichens but
consider it doubtful.
L. IMPUDENS Degel. – very rare, two localities
only: Saaremaa Co., Kuressaare, on Fraxinus excelsior, 28 June 1980, leg. & det. T.
Rand-lane, verified by G. Thor 1991 (TLC:
atranorin) (TU) (Randlane, 1986); Tartumaa
Co., Luunja, in humid forest on Tilia cordata,
1 Jan. 1949, leg. H. Trass (TLC: atranorin)
(TU).
L. PULICARIS (Pers.) Ach. [syn. L. chlarona (Ach.)
Nyl., L. coilocarpa (Ach.) Nyl., L. pinastri
(Schaer.) H. Magn.] – widespread in all
parts of Estonia (more than 200 localities).
On coniferous trees (very often on Pinus sylvestris), frequently on wood, on deciduous
trees (Alnus incana, Betula, etc.).
L. RUGOSELLA Zahlbr. – very common all over the
country (more than 100 localities), in forests
and parks, in the countryside and in towns.
On deciduous trees, also on wood.
Comments on the species’ list of the Lecanora subfusca group in Estonia
Fig. 3. Distribution of Lecanora circumborealis
in Estonia.
*L.
(Ach.) Ach. – very rare, only one locality from the last century: Saaremaa Co.,
Muhu Is. (Moon) (TU), “an der Erde über
Mossen” (Bruttan, 1870).
?L. GLABRATA (Ach.) Malme – very rare, one locality; mentioned by Mereschkowski (1913)
(Harjumaa Co., Tallinn), the specimen is not
EPIBRYON
Lecanora circumborealis is morphologically
and anatomically very close to L. pulicaris and
therefore, it is necessary to use the complex of
characteristics to distinguish the two taxa (p.
19); for more information see Brodo (1984) and
Brodo & Vitikainen (1984).
L. impudens is morphologically rather similar to L. allophana f. sorediata but, according to
Tønsberg (1992), their terpenoid patterns differ.
TLC was carried out on four local specimens
and showed that two specimens of L. im-pudens
contained only atranorin, while two specimens
of L. allophana f. sorediata contained some terpenoids in addition to atranorin.
Lecanora rugosella has been treated as a
form or synonym of L. chlarotera (Clauzade &
Roux, 1985; Poelt, 1952; Wirth, 1995) or also as
a distinctive species (Brodo, 1984; Purvis et al.
1992; Santesson, 1993; Vitikainen et al. 1997).
They both have the same epihymenium and
amphithecium types (Table 1), and other
19
anatomical and chemical features are also
similar. These species have been separated according to their morphology: L. rugosella has
coarsely verrucose thallus and more thicker
and verrucose apothecial margin than L. chlarotera. In Estonian herbarium material some
specimens of L. chlarotera are slightly sorediose, and there are quite a number of specimens
with soralia-like heaps among both L. chlarotera
and L. rugosella. Further investigation of those
species is needed.
Those specimens, which earlier were determined as L. subrugosa, are now considered
extreme morphological variants of L. argentata
caused by the substrate; no anatomical differences between those species could be observed,
futhermore, morphological intermediates exist
(Lumbsch & Feige, 1996). Morphological intermediate specimens are also present among
Estonian herbarium materials, and the distinguishing between these two species was often
impossible.
Key to the species of the Lecanora subfusca group in Estonia
A Saxicolous
1 Thallus sorediate, soralia pale yellow to yellowish white, usually rounded, slightly convex to
hemisphaerical, 1–2 mm in diameter, thallus grey, whitish or yellowish white. Apothecia infrequent ....................................................................................................
L. caesiosora
– Thallus not sorediate, thallus greyish white to grey, apothecia frequent .......................... 2
2 Thallus whitish grey to grey, apothecial discs red-brown to almost black, epruinose, margins
smooth and even. Epihymenium glabrata-type, amphithecium with small crystals ...........
L. campestris
– Thallus greyish white, apothecial discs varying in color - yellowish brown, greyish brown to
almost black, usually lightly pruinose, margins smooth to verrucose. Epihymenium chlaroteratype, amphithecium with large crystals .............................................................. L. cenisia
B Muscicolous
Thallus thick, verrucose, apothecia constricted at the base, crowded, discs red-brown. On moss
and dead vegetation ........................................................................................ L. epibryon
C Corticolous or lignicolous
1 Thallus sorediate ........................................................................................................... 2
– Thallus not sorediate ..................................................................................................... 3
2 Soralia mostly rounded, flat to convex, discrete or coalescing into larger, continuous patches.
Apothecia rare , <1 mm in diameter ............................................................... L. impudens
– Soralia flat to hemiglobose, usually discrete. Apothecia abundant, larger than 1 mm ............
........................................................................................
L. allophana f. sorediata
3 Amphithecium with small crystals (allophana-type) ........................................................ 4
– Amphithecium with large crystals (pulicaris-type) ........................................................... 5
4 Apothecia large, 1–3 mm diameter, constricted at the base, margin prominent, commonly
flexuose. Spores 12–21 × 7–11 mm ................................................................ L. allophana
– Apothecia small, 0.3–0.6 mm diameter, closely adnate, margin even with disc, finally excluded.
Spores 9–13 × 6–8 mm ........................................................................
L. glabrata
5 Epihymenium granular (pulicaris- or chlarotera-type) ..................................................... 6
– Epihymenium without granules (glabrata-type) ............................................................. 10
6 Epihymenium pulicaris-type ..........................................................................................
7
– Epihymenium chlarotera-type ........................................................................................ 8
7 Apothecial disks pale reddish brown to dark brown, thallus and/or only apothecial margin
Pd+ red (sometimes Pd-). Apothecial cortex 18–25 mm laterally, expanded to (25) 30–45 (66)
mm at the base, spores 11–15 × 7.5–9.5 mm, walls 0.8–1.0 mm ...................... L. pulicaris
– Apothecial discs dark red-brown to black, thallus and apothecial margin Pd+ yellow or Pd-.
Apothecial cortex 22–38 mm laterally, expanded to 35–65 (90) mm at the base, spores 13–17.5
× 8–12 mm, walls 1.0–1.2 mm ..................................................
L. circumborealis
20
Folia Cryptog. Estonica
8 Apothecial discs usually dark brown to brown-black, often lightly pruinose, apothecial margins
becoming blackish, adjacent to disc. On lignum .........................................
L. cenisia
– Apothecial discs pale dull brown, orange- or red-brown, pruinose or not, apothecial margins
remain unblackened. On bark, sometimes on lignum ..................................................... 9
9 Thallus coarsely verrucose, apothecia constricted at the base, raised, margin very thick and
verrucose, discs often lightly pruinose ............................................................ L. rugosella
– Thallus smooth to verrucose, apothecia sessile, margin thin or thick, smooth to verrucose,
discs usually epruinose ................................................................................
L. chlarotera
10 Thallus thin or thick, rough to verrucose, apothecia broadly attached or constricted at the
base, apothecial margin thin or thick, ± smooth, crenulate to verrucose .......... L.argentata
ACKNOWLEDGEMENTS
The author is very grateful to her supervisors T.
Randlane and A. Saag, sincere thanks also to
O. Vitikainen and A. Piterans for their help and
the opportunities to visit the herbaria in Helsinki and Riga. Dr. I. Brodo is thanked for kindly
permitting to reprint the figures of anatomical
characters. The stydy was undertaken as part
of the project “Composition and analysis of Estonian lichen flora” financed by the Estonian
Science Foundation (Grant No. 1297).
REFERENCES
Brodo, I. M. 1984. The North American species of the
Lecanora subfusca group. Beih. Nova Hedwigia
79: 63–185.
Brodo, I. M. 1988. Lichens of the Ottawa region. Second
Edition. Ottawa Field Naturalist’s Club Special
Publication 3, Ottawa. 115 pp.
Brodo, I. M., Owe-Larsson, B. & Lumbsch, H. T. 1994.
The sorediate, saxicolous species of the Lecanora
subfusca group. Nord. J. Bot. 14: 451–461.
Brodo, I. M. & Vitikainen, O. 1984. The typification
of Lecanora subfusca (L.) Ach., its varieties, and
some of its related taxa published before 1850.
Mycotaxon 21: 281–298.
Bruttan, A. 1870. Lichen Est-, Liv- und Kurlands.
Arch. Naturk. Liv-, Ehst- u. Kurl. Zweite Serie 7:
163–326.
Clauzade, G. & Roux, C. 1985. Likenoj de okcidenta
Europo. Bull. Soc. Bot. Centre-Ouest Nouv. s. Num.
Spec. 7: 1–893.
Culberson, C. F. 1972. Improved conditions and new
data for the identification of lichen products by a
standardized thin-layer chromatographic method.
J. Chromatogr. 72: 113–125.
Culberson, C. F. & Kristinsson, H. 1970. A standardized method for the identification of lichen
products. J. Chromatogr. 46: 85–93.
Eigler, G. 1969. Studien zur Gliedrung der Flechtengattung Lecanora. Diss. Bot. 4: 1–195.
Foucard, T. 1990. Svensk skorplavs flora. Lund. 306 pp.
Jüriado, I. 1997. Epilithic species of the lichen genera
Lecanora, Protoparmelia and Tephromela in Estonia. Folia Cryptog. Estonica 31: 26–29.
Lumbsch, H. T. & Feige, G. B. 1996. Comments on
the exsiccat “Lecanoroid Lichens” III. Mycotaxon
58: 259–267.
Magnusson, A. H. 1932. Beiträge zur Systematic der
Flechtengruppe Lecanora subfusca. Meddel. Göteborgs Bot. Trädg. 7: 65–87.
Mereschkowski, K. 1913. The checklist of lichens in the
Baltic provinces (in Russian). Kazan.
Poelt, J. 1952. Die Lecanora subfusca-Gruppe in Süddeutschland. Ber. Bayer. Bot. Ges. 29: 58–69.
Poelt, J. & Vêzda, A. 1981. Bestimmungsschlüssel europäischer Flechten. Erg. II. Bibl. Lichenol. 16.
Purvis, O. W., Coppins, B. J., Hawksworth, D. L.,
James, P. W. & Moore, D. M. (eds.) 1992. The
lichen flora of Great Britain and Ireland. Natural
History Publications, London. 710 pp.
Randlane, T. 1986. New species to the Estonian lichen-flora (in Russian). Folia Cryptog. Estonica
21: 8–10.
Räsänen, V. 1931. Die Flechten Estlands. Helsinki,
162 pp.
Santesson, R. 1993. The lichens and lichenicolous
fungi of Sweden and Norway. SBT -förlaget,
Lund. 240 pp.
Tønsberg, T. 1992. The sorediate and isidiate, corticolous, crustose lichens in Norway. Sommerfeltia
14: 1–331.
Trass, H. 1967. Analysis of the lichen-flora of Estonia
(in Russian). Doctoral dissertation. Tartu State
University.
Trass, H. 1970. The elements and development of the
lichen-flora of Estonia (in Russian). Pap. Bot. 9:
5–233.
Vitikainen, O., Ahti, T., Kuusinen, M., Lommi, S. &
Ulvinen, T. 1997. Checklist of lichens and allied
fungi of Finland. Norrlinia 6: 1–123.
Vitikainen, O. & Brodo, I. M. 1985. Proposal to reject
Lichen subfuscus L. (Lichenized fungi). Mycotaxon
34: 533–534.
Wirth, V. 1995. Flechtenflora. Ulmer, Stuttgart. 661 pp.
Folia Cryptog. Estonica, Fasc. 32: 21–25 (1998)
A catalogue of Heterodermia (Physciaceae)
Syo Kurokawa
1101 Kurose 47, Toyoma 939–8213, Japan
Abstract: All taxa of Heterodermia known at present are listed with special reference to the priority of combinations, especially
when the same combination was proposed in two or more times. New combinations under Heterodermia are proposed for
12 species.
Kokkuvõte: S. Kurokawa. Heterodermia (Physciaceae) kataloog.
Loetletud on kõik perekonna Heterodermia praegu teada olevad liigid viidetega prioriteetsetele kombinatsioonidele. Uute
kombinatsioonidena esitatakse 12 Heterodermia liiki.
INTRODUCTION
ENUMERATION OF TAXA
Since the genus Heterodermia Trevis. was resurrected by Poelt (1965), various authors have
proposed new combinations under the genus.
They are Culberson (1966), Skorepa (1972),
Follmann & Rédon (1972), Awasthi (1973),
Follmann (1974, 1983), K. P. Singh & S. R.
Singh (1976), Swinscow & Krog (1976), Hale
(Vêzda, 1976), Dey (1976), Wetmore (1976), K.
P. Singh (1979), A. Singh (1980), Weber (1981),
Hawksworth (Shaw, 1984), Elix (1985), Wei &
Jiang (1986), Weber (Egan, 1987), Wei (1991)
and Trass (1992). However, there are some
confusions in author citation. For instance, the
combination Heterodermia boryi was proposed
by three different authors in 1976. Although H.
boryi (Fée) Kr. P. Singh & S. R. Singh is earliest
as shown below and keeps the priority, H. boryi
(Fée) Hale was employed by Park (1990). Even
though Trevisan (1861) did not propose such
a combination, Aptroot (1987) used Heterodermia galactophylla (Tuck.) Trevis. In the present
paper, all taxa now considered to belong to Heterodermia will be listed, with special reference
to the priority of combination under the genus.
A number of new species of the genus described
by various authors in recent years are also included. In addition, new combinations under
Heterodermia are proposed for 12 species, which
have not yet been transferred to the genus.
HETERODERMIA ALBICANS (Pers.) Swinscow & Krog,
Lichenologist 8: 113. 1976.
Heterodermia speciosa var. domingensis (Ach.)
Trevis., Atti Soc. Ital. Sci. Nat. Milano 11:
614. 1868.
Heterodermia domingensis (Ach.) Trevis.,
Nuovo Giorn. Bot. Ital. 1: 114. 1869.
HETERODERMIA ALBIDIFLAVA (Kurok.) D. D. Awasthi,
Geophytology 3: 113. 1973.
H ETERODER MIA ALBOPRUINOSA (Kurok.) Kurok.,
comb. nov.
Anaptychia albopruinosa Kurok., Beih. Nova
Hedwigia 6: 32. 1962.
HETERODERMIA ALLARDII (Kurok.) Trass, Folia Cryptog. Estonica 29: 6. 1992.
HETERODERMIA ANGUSTILOBA (Müll. Arg.) D. D. Awasthi, Geophytology 3: 113. 1973.
HETERODERMIA ANTILLARUM (Vain.) Swinscow & Krog,
Lichenologist 8: 114. 1976.
HETERODERMIA APPALACHENSIS (Kurok.) W. L. Culb.,
Bryologist 69: 479. 1966.
HETERODERMIA APPENDICULATA (Kurok.) Swinscow &
Krog, Lichenologist 8: 114. 1976.
HETERODERMIA ARSENEI (Kurok.) Kurok., comb.
nov.
Anaptychia arsenei Kurok., Beih. Nova Hedwigia 6: 89. 1962.
HETERODERMIA AWASTHII (Kurok.) D. D. Awasthi,
Geophytology 3: 113. 1973.
HETERODERMIA BARBIFERA (Nyl.) Kr. P. Singh, Bull.
Bot. Survey India 21: 221. 1979.
Heterodermia barbifera (Nyl.) W. A. Weber,
Mycotaxon 13: 101. 1981.
Heterodermia barbifera (Nyl.) J. C. Wei, Enum.
Lich. China 106. 1991.
The present paper is dedicated to Prof. Dr
H. Trass who has been contributing to the
taxonomy of lichens, especially to that of the
Physciaceae.
22
Folia Cryptog. Estonica
HETERODERMIA BORYI (Fée) Kr. P. Singh & S. R.
Singh, Geophytology 6: 33. 1976 (July 31).
Heterodermia boryi (Fée) Swinscow & Krog,
Lichenologist 8: 124. 1976 (October).
Heterodermia boryi (Fée) Hale in Vêzda, Lich.
Selecti Exs., Fasc. 57, no. 1421. 1976 (October).
Heterodermia neoleucomelaena (Kurok.) Follmann & Rédon, Willdenowia 6: 446. 1972.
Heterodermia neoleucomelaena f. circinalis
(Zahlbr.) Follmann & Rédon, Willdenowia
6 : 446. 1972. – Heterodermia circinalis
(Zahlbr.) W. A. Weber, Mycotaxon 13 : 101.
1981. – Heterodermia boryi var. squarrosa f.
circinalis (Zahlbr.) J. C. Wei in Wei & Jiang,
Lich. Xizang 110. 1986.
Heterodermia neoleucomelaena (Kurok.) D. D.
Awasthi, Geophytology 3: 114. 1973.
Heterodermia neoleucomelaena f. sorediosa
(Jatta) D. D. Awasthi, Geophytology 3: 114.
1973. – Heterodermia boryi var. squarrosa
f. sorediosa (Jatta) J. C. Wei., Enum. Lich.
China 107. 1991.
Heterodermia neoleucomelaena f. soredobullata (D. D. Awasthi) D. D. Awasthi, Geophytology 3: 114. 1973.
Heterodermia neoleucomelaena f. squarrosa
(Vain.) D. D. Awasthi, Geophytology 3: 114.
1973. – Heterodermia boryi var. squarrosa
(Vain.) J. C. Wei in Wei & Jiang, Lich. Xizang
110. 1986.
HETERODERMIA CASSARETTIANA (A. Massal.) Trevis.,
Atti Soc. Ital. Sci. Nat. Milano 11: 624.
1868.
HETERODERMIA CHILENSIS (Kurok.) Swinscow & Krog,
Lichenologist 8: 115. 1976.
HETERODERMIA CHONDROIDEA W. A. Weber & D. D.
Awasthi, Bryologist 74: 181. 1971.
HETERODERMIA COMOSA (Eschw.) Follmann & Rédon, Willdenowia 6: 446. 1972.
Heterodermia comosa (Eschw.) D. D. Awasthi,
Geophytology 3: 113. 1973.
Heterodermia comosa (Eschw.) Follmann &
L. I. Ferrago in Follmann, Philippia 5: 118.
1983.
HETERODERMIA CONGOENSIS (Kurok.) Swinscow &
Krog, Lichenologist 8: 116. 1976.
HETERODERMIA CORALLOPHORA (Taylor) Skorepa,
Bryologist 75: 490. 1972.
Heterodermia hypoleuca var. dactylina * isidiophora Trevis., Atti Soc. Ital. Sci. Nat. Milano
11: 615. 1868.
HETERODERMIA CORONATA (Kurok.) D. D. Awasthi,
Geophytology 3: 113. 1973.
HETERODERMIA CROCEA R. C. Harris, Some Florida
Lichens 78. 1990.
HETERODERMIA CUBENSIS (Kurok.) Trass, Folia Cryptog. Estonica 29: 10. 1992.
HETERODERMIA CYATHIFORMIS (Kurok.) Kurok., comb.
nov.
Anaptychia cyathiformis Kurok., J. Hattori
Bot. Lab. 37: 602. 1973.
HETERODERMIA DACTYLIZA (Nyl.) Swinscow & Krog,
Lichenlogist 8: 117. 1976.
Heterodermia hypoleuca var. dactyliza (“dactylina”) (Nyl.) Trevis., Atti Soc. Ital. Sci. Nat.
Milano 11: 615 1868.
HETERODERMIA DENDRITICA (Pers.) Poelt, Nova Hedwigia 9: 31. 1965.
HETERODERMIA DIADEMATA (Taylor) D. D. Awasthi,
Geophytology 3: 113. 1973.
Heterodermia dispensa (Nyl.) Trévis., Atti Soc.
Ital. Sci. Nat. Milano 11: 615. 1868.
Heterodermia speciosa var. cinerascens (Nyl.)
Trevis., Atti Soc. Ital. Sci. Nat. Milano 11:
615. 1868.
Heterodermia diademata f. brachyloba (Müll.
Arg.) D. D. Awasthi, Geophytology 3: 113.
1973.
HETERODERMIA DISSECTA (Kurok.) D. D. Awasthi,
Geophytology 3: 113. 1973.
Heterodermia dissecta var. koyana (Kurok.) D.
D. Awasthi, Geophytology 3: 113. 1973.
Heterodermia dissecta var. koyana (Kurok.) J.
C. Wei, Enum. Lich. China 109. 1991.
HETERODERMIA ECHINATA (Taylor) W. L. Culb., Bryologist 69: 481. 1966.
HETERODERMIA ERINACEA (Ach.) W. A. Weber in Egan,
Bryologist 90: 163. 1987.
HETERODERMIA FAURIEI (Kurok.) Kurok., comb. nov.
Anaptychia fauriei Kurok., Beih. Nova Hedwigia 6: 83. 1962.
HETERODERMIA FIRMULA (Nyl.) Trevis., Atti Soc. Ital.
Sci. Nat. Milano 11: 615. 1868.
HETERODERMIA FLABELLATA (Fée) D. D. Awasthi,
Geophytology 3: 113. 1973.
Heterodermia flabellata var. rottbollii (Vain.) J.
C. Wei, Enum. Lich. China 109. 1991.
HETERODERMIA FLAVOSQUAMOSA Aptroot & Sipman in
Sipman, Willdenowia 19: 546. 1990.
H ETERODERMIA FOLLMANNII Sipman in Daniels,
Schulz & Peine, Flechten Follmann 333.
1995.
HETERODERMIA FRAGILISSIMA (Kurok.) J. C. Wei & Y.
M. Jiang, Lich. Xizang 111. 1986.
23
Heterodermia fragilissima (Kurok.) J. C. Wei,
Enum. Lich. China 109. 1991.
Heterodermia fragilissima (Kurok.) Trass, Folia
Cryptog. Estonica 29: 12. 1992.
HETERODERMIA GALATOPHYLLA (Tuck.) W. L. Culb.,
Bryologist 69: 482. 1966.
HETERODERMIA GRANULIFERA (Ach.) W. L. Culb., Bryologist 69: 482. 1966.
Heterodermia hypoleuca * isidiophora Trevis.,
Atti Soc. Ital. Sci. Nat. Milano 11: 615.
1868.
HETERODERMIA HIMALAYAENSIS (D. D. Awasthi) D. D.
Awasthi, Geophytology 3: 113. 1973.
HETERODERMIA HYPOCAESIA (Yasuda) D. D. Awasthi,
Geophytology 3: 113. 1973.
HETERODERMIA HYPOCHRAEA (Vain.) Swinscow &
Krog, Lichenologist 8: 119. 1976.
HETERODERMIA HYPOLEUCA (Ach.) Trevis., Atti Soc.
Ital. Sci. Nat. Milano 11: 615. 1868.
HETERODERMIA INCANA (Stirt.) D. D. Awasthi, Geophytology 3: 114. 1973.
HETERODERMIA INDICA (H. Magn.) D. D. Awasthi,
Geophytology 3: 114. 1973.
HETERODERMIA INTERMEDIA Trass, Folia Cryptog.
Estonica 29: 23. 1992.
HETERODERMIA ISIDIOPHORA (Nyl.) D. D. Awasthi,
Geophytology 3: 114. 1973.
Heterodermia speciosa ** isidiophora (Nyl.)
Trevis., Atti Soc. Ital. Sci. Nat. Milano 11:
614. 1868.
Heterodermia speciosa var. domingensis * isidiophora (Nyl.) Trevis., Atti Soc. Ital. Sci. Nat.
Milano 11: 614 1868.
HETERODERMIA ISIDIOSA (Kurok.) Kurok., comb.
nov.
Anaptychia rugulosa var. isidiosa Kurok., Beih.
Nova Hedwigia 6: 41. 1962.
HETERODERMIA JAPONICA (M. Satô) Swinscow & Krog,
Lichenologist 81: 122. 1976.
HETERODERMIA KUROKAWAE Trass, Folia Cryptog.
Estonica 29: 14. 1992.
HETERODERMIA LAMELLIGERA (Taylor) Follmann &
Rédon, Willdenowia 6: 446. 1972.
Heterodermia lamelligera (Taylor) Trass, Folia
Cryptog. Estonica 29: 15. 1992.
HETERODERMIA LEPIDOTA Swinscow & Krog, Lichenologist 8: 122. 1976.
HETERODERMIA LEUCOMELA (L.) Poelt, Nova Hedwigia
9: 31. 1965.
Heterodermia leucomela f. albociliata (Nyl.) D.
D. Awasthi, Geophytology 3: 114. 1973.
Heterodermia verruculifera (Kurok.) W. A. Weber, Mycotaxon 13 : 102. 1981.
HETERODERMIA LORIFORMIS (Kurok.) Swinscow &
Krog, Lichenologist 8: 129. 1976.
H ETERODER MIA LUTESCENS (Kurok.) Follmann,
Philippia 2: 73. 1974.
HETERODERMIA MAGELLANICA (Zahlbr.) Swinscow &
Krog, Lichenologist 8: 130. 1976.
HETERODERMIA MICROPHYLLA (Kurok.) Skorepa, Bryologist 75: 490. 1972.
Heterodermia microphylla (Kurok.) Swinscow
& Krog, Lichenologist 8: 132. 1976.
Heterodermia microphylla f. granulosa (Kurok.)
J. C. Wei, Enum. Lich. China 111. 1991.
HETERODERMIA MULTICILIATA (Kurok.) Follmann &
Rédon, Willdenowia 6: 446. 1972.
Heterodermia multiciliata (Kurok.) Trass, Folia
Cryptog. Estonica 29: 16. 1992.
HETERODERMIA NAMAQUANA Brusse, Bothalia 22:
183. 1992.
HETERODERMIA OBESA (Pers.) Trevis., Atti Soc. Ital.
Sci. Nat. Milano 11: 614. 1868.
Heterodermia obesa var. caesiocrocata (Nyl.)
Trevis., Atti Soc. Ital. Sci. Nat. Milano 11:
614. 1868.
Heterodermia obesa (Pers. ) Trass, Folia Cryptog. Estonica 29: 16. 1992.
HETERODERMIA OBSCURATA (Nyl.) Trevis., Nuovo
Giorn. Bot. Ital. 1: 114. 1869.
Heterodermia speciosa var. obscurata (Nyl.)
Trevis., Atti Soc. Ital. Sci. Nat. Milano 11:
614. 1868.
HETERODERMIA PACIFICA (Kurok.) Kurok., comb.
nov.
Anaptychia pacifica Kurok., J. Hattori Bot.
Lab. 37: 592. 1973.
HETERODERMIA PALPEBRATA (Taylor) Trass, Folia
Cryptog. Estonica 29: 17. 1992.
HETERODERMIA PANDURATA (Kurok.) J. C. Wei, Enum.
Lich. China 112. 1991.
Heterodermia pandurata (Kurok.) Trass, Folia
Cryptog. Estonica 29: 17. 1992.
HETERODERMIA PAPUANA Aptroot & Sipman, Willdenowia 20: 241. 1991.
HETERODERMIA PELLUCIDA (D. D. Awasthi) D. D.
Awasthi, Geophytology 3: 114. 1973.
H ETERODER MIA PERUVIANA (Kashiw. & Kurok.)
Kurok., comb. nov.
Anaptychia peruviana Kashiw. & Kurok., Bull.
Nat. Sci. Museum, Tokyo, ser. B, 16: 154.
1990.
HETERODERMIA PINNATA Sipman in Daniels, Schulz
& Peine, Flechten Follmann 330. 1995.
24
Folia Cryptog. Estonica
HETERODERMIA PODOCARPA (Bél.) D. D. Awasthi,
Geophytology 3: 114. 1973.
HETERODERMIA POLYRHIZA (Kurok.) Kurok., comb.
nov.
Anaptychia polyrhiza Kurok., Beih. Nova Hedwigia 6: 32. 1962.
HETERODERMIA PROPAGULIFERA (Vain.) J. P. Dey in
Parker & Roane, Distr. Hist. Biota S. Appalachians 4: 403. 1976.
Heterodermia dendritica var. propagulifera
(Vain.) Poelt, Nova Hedwigia 9: 31. 1965.
Heterodermia dendritica var. propagulifera
(Vain.) Ajay Singh, Lichenol. Ind. Subcont.
1966–1977. 47. 1980 (nom. illeg., basionym
not cited).
HETERODERMIA PSEUDOSPECIOSA (Kurok.) W. L. Culb.,
Bryologist 69: 484. 1966.
HETERODERMIA PUNCTIFERA (Kurok.) D. D. Awasthi,
Geophytology 3: 114. 1973.
HETERODERMIA RUBESCENS (Räsänen) D. D. Awasthi, Geophytology 3: 114. 1973.
HETERODERMIA RUGULOSA (Kurok.) Wetmore, Bryologist 79: 304. 1976.
Heterodermia rugulosa (Kurok.) Trass, Folia
Cryptog. Estonica 29: 19. 1992.
HETERODERMIA SITCHENSIS Goward & W. J. Noble in
Goward, Bryologist 87: 366. 1984.
HETERODERMIA SPECIOSA (Wulf.) Trevis., Atti Soc.
Ital. Sci. Nat. Milano 11: 614. 1868.
HETERODERMIA SPINIGERA (Kurok.) Kurok., comb.
nov.
Anaptychia spinigera Kurok., Beih. Nova Hedwigia 6: 66. 1962.
HETERODERMIA SPINULOSA (Kurok.) J. C. Wei, Enum.
Lich. China 113. 1991.
Heterodermia spinulosa (Kurok.) Trass, Folia
Cryptog. Estonica 29: 20. 1992.
HETERODERMIA SQUAMULOSA (Degel.) W. L. Culb.,
Bryologist 69: 484. 1966.
HETERODERMIA STELLATA (Vain.) W. A. Weber, Mycotaxon 13: 102. 1981.
HETERODERMIA SUBASCENDENS (Asahina) Trass, Folia
Cryptog. Estonica 29: 20. 1992.
Heterodermia subascendens (Asahina) W. L.
Culb. in Wei, Enum. Lich. China 113. 1991.
(nom. invalid., basionym not cited).
HETERODERMIA SUBCOMOSA (Nyl.) Elix, Lich. Austr.
Exs., no. 80. 1985.
Heterodermia subcomosa (Nyl.) Trass, Folia
Cryptog. Estonica 29: 20. 1992.
HETERODERMIA SUBISIDIOSA (Kurok.) Kurok., comb.
nov.
Anaptychia magellanica var. pectinata f. subisidiosa Kurok., Beih. Nova Hedwigia 6: 68.
1962.
HETERODERMIA SZECHUANENSIS (J. D. Zhao, L. W.
Hsu & Z. M. Sun) J. C. Wei, Enum. Lich.
China 113. 1991.
Heterodermia szechuanesis f. albo-marginata
(J. D. Zhao, L. W. Hsu & Z. M. Sun) J. C.
Wei, Enum. Lich. China 113. 1991.
HETERODERMIA TOGASHII (Kurok.) D. D. Awasthi,
Geophytology 3: 114. 1973.
HETERODERMIA TRANSLUCENS (Kurok.) D. Hawksw.
in Shaw, Microorgan. PNG Res. Bull. 33:
252. 1984.
HETERODERMIA TREMULANS (Müll. Arg.) W. L. Culb.,
Bryologist 69: 485. 1966.
HETERODERMIA TRICHOPHORA (Kurok.) Trass, Folia
Cryptog. Estonica 29: 21. 1992.
HETERODERMIA TRICHOPHOROIDES (Kurok.) Kurok.,
comb. nov.
Anaptychia trichophoroides Kurok., Beih. Nova
Hedwigia 6: 101. 1962.
HETERODERMIA TROPICA (Kurok.) Kurok., comb.
nov.
Anaptychia tropica Kurok., Beih. Nova Hedwigia 6: 36. 1962.
HETERODERMIA UNDULATA (J. D. Zhao, L. W. Hsu
& Z. M. Sun) J. C. Wei, Enum. Lich. China
113. 1991.
HETERODERMIA USAMBARENSIS (Kurok.) Swinscow &
Krog, Lichenologist 8: 137. 1976.
HETERODERMIA VULGARIS (Vain.) Follmann & Rédon,
Willdenowia 6: 447. 1972.
HETERODERMIA YUNNANENSIS (J. D. Zhao, L. W. Hsu
& Z. M. Sun) J. C. Wei, Enum. Lich. China
113. 1991.
ACKNOWLEDGEMENTS
My sincere appreciation is expressed to Dr. D. D.
Awasthi in India, Dr. J. A. Elix of the Australian
National University, Dr. R. C. Harris of the New
York Botanical Garden, Dr. D. L. Hawksworth of
the International Mycological Institute, Kew, Dr.
K. P. Singh of the Botanical Survey in India and
Dr. D. K. Upreti of the National Botanical Research Institute, Lucknow for their cooperations
in providing informations about the literature.
25
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Skorepa, A. C. 1972. A catalogue of the lichens reported from Tennessee. Bryologist 75: 481–500.
Swinscow, T. D. V. & Krog, H. 1976. The genera
Anaptychia and Heterodermia in East Africa.
Lichenologist 8: 103–138.
Trass, H. 1992. Synopsis of the lichen genus Heterodermia (Ascomycotina, Physciaceae sive Pyxidaceae). Folia Cryptog. Estonica 29: 2–24.
Trevisan, V. G. 1861. Über Atestia, eine neue Gattung der Ramalineen aus Mittel-America. Flora
44: 49–53.
Vêzda, A. 1976. Lichenes Selecti Exiccati, Fasc. 57,
Nos 1401–1425. Brno.
Weber, W. A. 1981. Lichenes Exiccati distributed by
the University Colorado Museum, Boulder, Fascicle 1–15, Nos. 1–600, 1961–1074. Mycotaxon
13: 85–104.
Wei, J. C. 1991. An Enumeration of Lichens in China.
278 pp. Intern. Academic Pr., Bejing.
Wei, J. C. & Y. M. Jiang 1986. Lichens Xizang. 130
pp. Science Pr., Bejing.
Wetmore, C. M. 1976. Macrolichens of Big Bend National Park, Texas. Bryologist 70: 296–313.
26
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 27–35 (1998)
Problems related to the marine lobate and subfruticose species of
Caloplaca
E. Ingvar Kärnefelt
The Botanical Museum, Lund University, Östra Vallgatan 18, S-223 61 Lund, Sweden
Abstract: 25 lobate and subfruticose species of Caloplaca occurring mainly on coastal rocks are discussed. This special
group of lichens is relatively well-known from western Europe and North America. However, for other parts of the world
with suitable environmental conditions such as in Australia, southern Africa and South America, there are still large gaps of
information on this basically ecologically delimited group of lichens.
Kokkuvõte: E. I. Kärnefelt. Mariinsete hõlmiste ja poolpõõsasjate kuldsamblike (Caloplaca) liikidega
seotud probleemid.
Käsitletakse 25 hõlmadega või poolpõõsasja tallusega kuldsambliku (Caloplaca) liiki, mis levivad peamiselt rannakaljudel. Need
samblikud on Lääne-Euroopas ja Põhja-Ameerikas üsna hästi uuritud. Teistest piirkondadest – Austraaliast, Lõuna-Aafrikast
ja Lõuna-Ameerikast – on selle ökoloogiliselt hästi piiritletud samblike rühma kohta veel vähe teada.
INTRODUCTION
The supergenus Caloplaca (Fr.) Th. Fr. comprises
a large diversity of presumably near 1100 species, and can partly be divided in more or less
distinctive species groups (Kärnefelt, 1989; Poelt & Hinteregger, 1993; Wetmore & Kärnefelt,
1998). It is under discussion whether some of
these groups could represent genera or not
(Kärnefelt, 1989; Poelt & Pelleter, 1984). This
is also a matter of principles and different generic concepts (Kärnefelt, 1991; Nimis, 1997).
Here, I guess we will always have to deal with
personal views related to what value one could
assign to important characters, in addition to
general knowledge of the diversity and variation
in a certain group.
In the relatively new Estonian macrolichen
flora by Trass & Randlane (1994) several placodioid groups were included. One of these groups
was Caloplaca sect. Gasparrinia comprising 11
species, of which C. marina, C. microthallina,
C. scopularis, C. thallincola and C. verruculifera were treated, all well-known from western
Europe occurring in the spray zone on coastal
rocks (Nordin, 1972; Purvis et al., 1992). Coastal
rocks in mainly temperate and subtropical regions are fairly richly colonized by lichens of
the Teloschistaceae, particularly of the genus
Caloplaca but also Teloschistes and Xanthoria.
Presumably the combination of several factors,
involving nutrient rich sites, exposure and
competition, have made this group of lichens
successful in coastal situations throughout
the world.
Is Caloplaca section Gasparrinia worth being recognized taxonomically at generic level,
or is it just one of these names which has been
mentioned in floras from time to time? An answer like ”usually the Gasparriniae can be easily recognized” shows some weak points. Seen
from the character (1) ”lobed or with distinctive
marginal lobes”, there are species which will
be difficult to place within a possible genus.
The other important characters defining a possible genus Gasparrinia are (2) ”anthraquinone
pigmented lobes” and (3) ”cortical layers with
more or less distinctive paraplectenchymatous
hyphae” (Wetmore & Kärnefelt, 1998). However, there are species within the grey or black
species group which are both lobed and lack
anthraquinones. Most of the yellow and lobed
species have distinctive paraplectenchymatous
cortical layers, but there are also species in this
group with different cortical structures.
The aim of this paper, in connection with
other projects in the genus Caloplaca (Wetmore
& Kärnefelt, 1998), is the present an overview,
worldwide, of these species.
28
Folia Cryptog. Estonica
DISCUSSION
The lobate species of Caloplaca occurring on
coastal rocks have been fairly well investigated
in western Europe (James, 1962; Nordin,
1972; Purvis et al., 1992) and recently also in
North America (Arup, 1995b). In other parts
of the World, however, the knowledge of this
group is much less well documented. The most
spectacular maritime belt dominated by Teloschistaceae is generally found on the western
parts of the continents, mainly in temperate but
also in subtropical regions, where the species
encounter the large ocean waters.
List of marine lobate and subfruticose species discussed
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
C.
BONAE-SPEI
Almb. & Poelt
Weber
CORALLOIDES (Tuck.) Hult.
CRIBROSA (Hue) Zahlbr.
EUDOXA (Müll. Arg.) Zahlbr.
FRAGILLIMA Wern.
GOMERANA J. Steiner
IGNEA Arup
IMPOLITA Arup
ISIDIOCLADA Zahlbr.
ISIDIOSA (Vain.) Zahlbr.
LITTOREA Tav.
LUCENS (Nyl.) Zahlbr.
MARINA (Wedd.) Zahlbr.
MAURITANICA Wern.
MICROTHALLINA (Wedd.) Zahlbr.
ORA Poelt & Nimis
ORTHOCLADA Zahlbr.
REGALIS (Vain.) Zahlbr.
ROSEI Hasse
SCOPULARIS (Nyl.) Lett.
SUBLOBULATA (Nyl.) Zahlbr.
THALLINCOLA (Wedd.) Du Rietz
THAMNODES Poelt
VERRUCULIFERA (Vain.) Zahlbr.
BRATTEI
The Arctic and adjacent regions
Coastal rocks, particularily in the arctic and
adjacent regions, are rarely colonized by a rich
lichen cover. This is probably associated to the
harsh environmental conditions during the long
winter season. In King Karls Land in eastern
Svalbard, where I spent the summer of 1980, I
did not observe any marine Teloschistaceae at
all, apart from scattered Xanthoria elegans .
Extensive field work has been carried out
especially in Greenland within the Arctic region.
However, Hansen et al. (1987) only recorded C.
scopularis and C. verruculifera from coastal
rocks.
Western Europe
The most frequent species along the sea coast
in western Europe is probably C. marina, an
exclusively littoral species, well adapted to its
habitat and with rather limited morphological
variation. Caloplaca marina varies, however, in
the development of the marginal lobes and it
has been discussed whether it should be treated
among the lobate species or not (Arup, 1992;
Wetmore & Kärnefelt, 1998). Caloplaca littorea,
known from the British Isles and the southern
part of the European coast line, characterized
by small isidia and with less well developed
marginal lobes, is presumably closely related
with C. marina (Kärnefelt, 1990b; Purvis et al.,
1992). Nordin (1972) treated C. microthallina in
the Gasparrinae. This species, characterized by
a rather small thallus composed of squamulose
lobules, is probably more closely related to the
C. squamosa group (Wetmore & Kärnefelt,
1998).
Three other species with marginal lobes
are rather widespread along the European sea
coast, i.e. C. scopularis, C. thallincola and C.
verruculifera. Caloplaca scopularis is the smallest of these species, characterized by lobes with
normal paraplectenchymatous layers and polarilocular spores. Caloplaca thallincola is usually
much larger, with distinctly effigurated lobes.
The citriform shape of the ascospores, however,
suggests that this species is more closely related
to two mainly inland species, C. aurantia and C.
flavescens. Caloplaca verruculifera is unique in
having verruciform or globular isidia covering the
lobes (Kärnefelt, 1990b). The cortex also differs
in the presence of more conglutinated or prosoplectenchymatous hyphae. Presumably, C. verruculifera is most closely related to C. granulosa,
which can also occur at much higher altitudes
(Nimis, 1993; Poelt & Romauch, 1977). Seen
particularly on the cortical structure and general habitat ecology C. verruculifera could also
29
show affinities with C. gomerana from the
Mediterranean region, and furthermore with C.
lucens and C. orthoclada from South America.
The Mediterranean region and northern
Africa
This region seems in general to be poor in lobate species occurring on coastal rocks. In the
first place this might be related to the warmer
climatic conditions and to warmer water.
Among the species discussed so far, C.
thallincola occurs extremely rarely from coastal
rocks in the Mediterranean region (Nimis & Poelt, 1987; Nimis, 1993). This beautiful species
is otherwise more widespread in the southern
parts of western Europe and in the British
Isles, where it prefers calcareous rocks. The
most closely related species, C. aurantia and C.
flavescens, are both relatively widespread in the
Mediterranean region also in coastal situations
(Clauzade & Roux, 1985; Ozenda & Clauzade,
1970).
Among the species known from coastal rocks
in the Mediterranean region there are entities
within the C. marina group, which, as mentioned
before, not really belong to the lobate species.
Caloplaca ora was separated from C. marina on
the generally smaller thallus with thinner lobes,
spores, and paraphyses (Nimis & Poelt, 1987).
It is obviously widespread in the Mediterranean
region. Caloplaca littorea was listed by Nimis
from Sardinia, where it is, however, very rare
(1993).
Another beautiful species, C. gomerana or
until recently referred to as C. gloriae, known
from the eastern Iberian Peninsula and from
north-western Africa and Macaronesia (Hafellner, 1995), is not a strictly littoral species,
occurring mainly near the coast up to 250 m
altitudes. Caloplaca gomerana is unique in its
laminal pseudocyphellae and a special type of
cortical layers (Llimona & Werner, 1975). Similar
pseudocyphellae are occasionally developed in
C. verruculifera, to which this species is probably
related (Arup, 1994).
Eastern North America
The marine species of Caloplaca occurring on
sea shore rocks in North America were treated
by Arup (1995b). Five species, of which at
least two have distinct marginal lobes, i.e. C.
scopularis and C. verruculifera were recorded
for eastern North America (Arup, 1994). Many
plant groups, and biota in general, distributed
in western Europe, occur in corresponding habitats in eastern North America, which is linked
both by pre-Pleistocene history and reinvasion of
biota after Pleistocene glaciations (Hultén, 1963;
Kärnefelt, 1979). Of the two species mentioned,
however, C. verruculifera also occurs on the
North American west coast.
Arup (1994) also treated the squamulose C.
microthallina from both the eastern and western
coasts.
Western North America
Arup (1992, 1995a) treated several different species of Caloplaca occurring on coastal rocks in
western North America. Among these, however,
only three belong to the lobate group. One of
them is C. brattiae, a strictly littoral species occurring in the lowest belt along the Californian
coast line (Arup, 1995a). It is presumably closest to C. scopularis. These two species actually
behave as vicarious taxa, having only minor differences in spore size, quantity of apothecia and
structure of lobe tips. Caloplaca scopularis only
occurs on sea shore rocks on the north American
east coast and C. brattiae on the west coast.
Caloplaca brattiae differs from C. verruculifera,
which occurs further north, in the structure of
the cortical layer.
Two other species, C. ignea and C. impolita,
confined to the coast line, have distinctive
marginal lobes (Arup, 1995a). Caloplaca ignea,
characterized by bright orange to reddish lobes
and the central portions covered with apothecia,
is not a strictly littoral species and may occur
as high up as to 500 m. This species is probably most closely related the C. saxicola group.
Caloplaca impolita, which has much broader and
flatter marginal lobes, may also occur at slightly
higher altitudes and not only on the shore. Both
species are mainly distributed along the southwestern coast from central California to Baja
California (Arup, 1995a; Wetmore & Kärnefelt,
1998).
Arup (1992) also reported C. marina from
the North American west coast. Caloplaca rosei,
which occurs within the same range as C. marina, has less well developed marginal lobes and
a more continuously areolated thallus.
30
Folia Cryptog. Estonica
The most unusual species occurring on
coastal rocks in western North America, however, is C. coralloides (Arup, 1995a; Poelt &
Pelleter, 1984; Wetmore & Kärnefelt, 1998).
This is a strictly littoral species, known from
Oregon to Baja California and characterized
mainly by a subfruticose thallus consisting of
yellowish coralloid lobes (Poelt & Pelleter, 1984).
A slightly different species, C. thamnodes, with
broader and more dark orange, coralloid lobes
and scattered pseudocyphellae, occurs further
south in the northern coast of Baja California,
where both species may occur in the same habitat on rather exposed coastal rocks. Otherwise
these species do not differ much in anatomical
features, both being characterized by rather
massive prosoplectenchymatous cortical layers and by the typical subfruticose shape of
the thallus. It is remarkable that a similar pair
of species, C. bonae-spei and C. eudoxa, has
developed under similar environmental conditions along the Atlantic coast in Southern Africa
(Kärnefelt, 1991).
Eastern Asia
The lichen flora in the eastern asiatic region
has not been well investigated, perhaps with
the exception of Japan. There are also relatively
few duplicate collections of maritime species of
Caloplaca distributed in European and North
American herbaria.
There seem to be rather few lobate marine
species in Caloplaca from this region. Caloplaca
scopularis seems to be one of the most common
species, at least reported from Japan (Yoshimura, 1987). This species dominates the coastal
rocks I studied in Petrov Island in Primorye,
Russia in 1991. Presumably, C. brattiae could
also occur in this region, which is much more
probable than the occurrence of C. scopularis
judging from their biogeographical patterns.
There are many examples of Amphi-Beringian
plants which have wide distributions in western North America, over the Aleutains, down to
Kamtchatka and the Far East region (Kärnefelt,
1979). Caloplaca scopularis does not match this
pattern at all, belonging to an eastern North
American and Amphi-Atlantic element.
Caloplaca marina probably occurs rarely in
the far eastern region seen from my own collections from Japan and Primorye.
Australasia and New Zealand
There are several, strictly littoral species of
Caloplaca in Australia. Some of them are still
undescribed and the whole genus is currently
being treated for the Flora of Australia project.
Among the littoral species, however, only a few
have marginal lobes, the most conspicuous
being C. sublobulata. This species, which originally was described from southernmost South
America, has been slightly misunderstood due
to its broad morphological variation (Kärnefelt,
1988; Santesson, 1944). During my fieldwork in
Australia in October–November 1997, I observed
large populations of C. sublobulata both from
coastal rocks in Western Australia and from
Victoria in southeasternmost Australia (Fig. 1).
I have seen material of this distinctive species
also from New Zealand. Among the other littoral
species, I also recorded a few small populations
of C. marina on coastal rocks in Victoria.
Caloplaca cribrosa occasionally occurs on
coastal rocks in Tasmania and New Zealand
(Poelt & Pelleter, 1984). This species is characterized by rather coarse lobes with distinct pseudocyphellae which occasionally are also slightly
raised like in the other subfruticose species (Fig.
2). It is strange that this species does not seem
to occur on the Australian mainland such as
in the SW region or along the southern coast.
Furthermore, Caloplaca cribrosa does not form
a biogeographical pair similar with the related
species occurring in SW North America and in
SW Africa. Caloplaca cribrosa is more likely to
be associated with C. regalis in the Western
Hemisphere. The anatomical characters in the
cortical layers are otherwise the same in this
species and in the American and African ones
known (Fig. 3).
Africa
Apart from various local floristic works, Africa is
still a lichenologically very much underinvesitgated continent. I have seen almost no records of
littoral species of Caloplaca from the large tropical regions north of the southern African states.
In South Africa C. sublobulata has been recorded
from coastal rocks along with a closely related
isidiate species C. isidiosa (Kärnefelt, 1990a, b).
Caloplaca marina also occurs scattered in the
same region.
31
Fig. 1. Caloplaca sublobulata . a. Habitus, b. details of lobes, c. apothecia, d. ascospores, e. cross
section of thallus portion with apothecium, f. paraphyses, g. ascus, h. cross section of lobe, i.
longitudinal section of lobe (MEL 1031729) (drawing by Rex Filson).
32
Folia Cryptog. Estonica
Fig. 2. Caloplaca cribrosa. a. Habitus, b. details of lobe (MEL 1000439), c. apothecium, e. cross
secion of apothecium (MEL 1012612), d. apothecium with pseudocyphellate margin, f. cross section (MEL 1000439) (drawing by Rex Filson).
33
approach, or at least coastal. If any, this group
of seven species, characterized by a combination
of anatomical and morphological characters and
habitat selection, should be separated at generic
level (Figs 2, 3).
The Antarctica and adjacent regions
Fig. 3. Caloplaca cribrosa. a. Cross section, b.
longitudinal section (MEL 1000439) (drawing
by Rex Filson).
Two of the unique subfruticose species occur within the western part of southern Africa
on coastal rocks, C. bonae-spei from the southernmost part, and the closely related C. eudoxa
further north from central Namibia to southern
Angola (Kärnefelt, 1991; Poelt & Pelleter, 1984).
Caloplaca bonae-spei is a strictly littoral species,
often occurring within populations of C. isidiosa and C. sublobulata, while C. eudoxa can
occur also in the inland. Much further north in
Mauritania, a third species C. mauritanica has
been recognized within this remarkable group
of lichens. I have not seen any material from the
only known locality, situated more inland and
not really littoral. But I would guess that C. mauritanica is extremely close to, or even identical
with, C. eudoxa. The whole subfruticose group
preferably seems to be littoral in its ecological
Søchting & Øvstedal (1991) made a survey of
the genus in the Western Antarctic region. In
spite of the limited open land available and of
the many scientific expeditions, the lichen flora
of this region is still very poorly known. Fourteen
Caloplaca species were recorded altogether, but
only a few of them develop distinctive marginal
lobes and occur on coastal rocks, i.e. C. isidioclada, C. lucens, C. sublobulata and C. regalis.
Of these species C. isidioclada, C. lucens and
C. sublobulata have all more or less distinctive
marginal lobes, while C. regalis belongs to the
group of subfruticose species. Caloplaca isidioclada is mainly characterized by lobes partly
being covered by finger-like to coralloid isidia
and cortical layers composed of paraplectenchymatous cells (Søchting & Øvstedal, 1992). Its
relation to C. isidiosa (Vain.) Zahlbr. is unclear,
and type material has to be examined.
The identity of C. lucens is still somewhat
unclear. The material referred to by Søchting
& Øvstedal (1992) is saxicolous, while the type
from Kerguelen Island was based on lignicolous
material (Santesson, 1944). This species, furthermore, is probably not very closely related
to the other marginally lobed species, apart
from C. gomerana, C. verruculifera and the
inland species C. trachyphylla (Hansen et al.,
1987; Wetmore & Kärnefelt, 1998). Caloplaca
lucens and the other species have more or less
rough convex lobes with isidia or papillae, tiny
or distinct pseudocypheallae and prosoplectenchymatous cortical layers.
Caloplaca regalis belongs to the group of
subfruticose taxa (Poelt & Pelleter, 1984). It
differs slightly from the pairs C. bonae-spei/C.
eudoxa and C. coralloides/C. thamnodes in the
presence of more horizontally directed marginal
lobes and thin paraplectenchymatous external
cortical layers. Furthermore C. regalis does not
seem to be strictly littoral, but it is more general
coastal, occurring higher up to several hundred
meters above sea level. Caloplaca ambitiosa,
34
Folia Cryptog. Estonica
which was described from the subantarctic islands, has shown to fall within the variation of
C. regalis (Søchting & Øvstedal, 1992).
South America
South America is a large continent with a very
long coastal line and, similarily to Africa and
Australia, with a relatively little known lichen
flora. Apart from a few special treatments, e.g.
by Malme (1926) on the Teloschistaceae, only
larger coastal species of Caloplaca have been
mentioned (Grassi, 1952; Santesson, 1944).
Among the species already discussed C. isidiosa is known from scattered localities in Brasil
and in the Galapagos Islands (Kärnefelt, 1988,
1990a, b).
Other littoral lobate species known are C.
sublobulata, mainly from southernmost South
America (Kärnefelt, 1988; Santesson, 1944).
From Chile I have seen material of a species
named C. orthoclada, which in some ways reminds of C. lucens with distinctly effigurated
marginal lobes. The surface of the lobes, however, is not rough as in C. lucens, but the cortical layer composed of prosoplectenchymatous
layers is, of the same structure. Type materials
must be more carefully examined in these taxa
in order to settle possible affinities.
Among the subfruticose species, C. regalis is
known from coastal situations in southernmost
South America including the Falkland Islands.
Further north along the Chilean coast near Valparaiso, another species, C. fragillima, has been
recognized, growing on coastal rocks (Poelt &
Pelleter, 1984). Caloplaca fragillima, known only
from the type locality, is presumably very closely
related to C. coralloides or even identical with
this species. Much more field work should be
carried out along the great Pacific coastline, and
also along the Atlantic coast in South America
which probably will reveal very interesting material for phytogeographical discussions and
theories on evolution and affinities especially
among Southern Hemisphere entities.
ACKNOWLEDGMENTS
I wish to thank Drs Lars Fröberg, Pier Luigi
Nimis and Arne Thell for valuable comments
and suggestions for improvement of the text. I
also wish to thank Dr Cliff Wetmore, St Paul for
company during the field work in Australia and
Magn. Bergvalls Stiftelse for financial support.
Dr Rex Filson, Boral, N.S.W., is greatly thanked
for providing me with his research material on
the genus from Australia including the illustrations.
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and Evolution 148: 51–88.
Poelt, J. & Romauch, E. 1977. Die Lagerstrukturen
placoidaler Küsten- und Inlandflechten. In Beiträge zur Biologie der niederen Pflanzen (eds Frey,
W., Hurka, H. & Oberwinkler, F.), pp. 141–153.
Jena, Gustav Fischer-Verlag.
Purvis, O., Coppins, B., Hawksworth, D., James, P. &
Moore, D. 1992. The Lichen Flora of Great Britain
and Ireland. London. ix + 710 pp.
Santesson, R. 1944. Contributions to the lichen flora of
South America. Arkiv för Botanik 31 A, 7: 1–28.
Søchting, U. & Øvstedal, D. O. 1992. Contributions
to the Caloplaca flora of the western Antarctic
region. Nordic Journal of Botany 12, 1: 121–134.
Trass, H. & Randlane, T. (eds) 1994. Eesti suursamblikud. Tartu. 399 pp.
Wetmore, C. & Kärnefelt, E. I. 1998. The lobate and
subfruticose species of Caloplaca in North America and Central America. Bryologist (in press).
Yoshimura, I. 1987. Lichen flora of Japan in color.
Higashi-ku, Osaka. 349 pp.
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Folia Cryptog. Estonica, Fasc. 32: 37–42 (1998)
Bioindication of air quality by lichens in a small town Viljandi in
southern Estonia
Siiri Liiv & Enel Sander
Tallinn Botanic Garden, 52 Kloostrimetsa Rd., EE0019 Tallinn, Estonia
Abstract: Air quality was estimated in Viljandi in 1987, 1991 and 1997 by the method of mapping of the distribution of
indicator lichens growing on small-leaved lime (Tilia cordata) and Norway maple (Acer platanoides). The data obtained in these
years show constant deterioration of air quality in Viljandi. Basing on the results of 1997, we distinguished four lichen zones:
I, the zone of lichens tolerant to strong acidic air pollution (acidification of the substrate); II, the zone of lichens tolerant
to moderate acidic air pollution; III, the zone of lichens tolerant to alkaline dust pollution (eutrophication of the substrate)
and IV, the zone of lichens sensitive to acidic air pollution.
Kokkuvõte: S. Liiv ja E. Sander. Õhu seisundi lihhenoindikatsiooniline hindamine Viljandis.
Viljandis on õhu seisundit hinnatud 1987., 1991. ja 1997. a., kasutades harilikul pärnal (Tilia cordata) ja harilikul vahtral (Acer
platanoides) kasvavate indikaatorsamblike leviku kaardistamise meetodit. Nende kolme aasta andmete põhjal võib täheldada õhu
kvaliteedi jätkuvat halvenemist Viljandis. 1997. a. tulemuste põhjal eraldati 4 tsooni: I – õhu tugevat happelist saastet taluvate
samblike tsoon; II – õhu mõõdukat happelist saastet taluvate samblike tsoon; III – tolmusaastet (substraadi eutrofeerumist)
taluvate samblike tsoon; IV – õhu happelise saaste suhtes tundlike samblike tsoon.
INTRODUCTION
In recent years relatively numerous investigations into the accumulation of elements in
lichens as a characteristic of air quality have
been conducted. However, beside this method
of the bioindicational assessment of air
quality, also the classical method of mapping
of the distribution of indicator species is used,
often combined with the index method (Wirth,
1987; Hosiaisluoma, 1994; Hobohm, 1994;
Kirschbaum, 1995; van Dobben, 1996; Liiv
& Sander, 1996). The method of mapping of
the distribution of indicator lichens serves as
the quickest and simplest way for establishing
the problematic areas of air pollution, but also
for fixing the improvement in air quality in
some region on the basis of the distribution of
indicator species (Hobohm, 1994; van Dobben,
1996; Ammann, 1997).
In Estonia, the use of lichens as bioindicators
for the assessment of air quality was initiated in
the late 60s at the then Tartu State University
with the study of lichen groupings growing on
trees in the town of Tartu (Trass, 1968 a, b).
In Viljandi, irregular assessments of air quality
have been performed since 1972, using different
methods of lichen analysis. In this year field
works were started to collect material for a
graduation thesis under the supervision of
Prof. Hans Trass (Liiv, 1973). Lichen studies
were repeated in Viljandi in 1976 and 1983
(Liiv, 1978, 1987, 1989 a, b; Eensaar, Liiv,
1983). Analyses of epiphytic lichen groupings
in Viljandi were carried out also in 1987 and
1991. These were mostly aimed at solving
the methodical problems of using lichens as
bioindicators. The present paper gives a brief
comparison of the results of the lichen analyses
performed in Viljandi in these years with the
data we obtained during mapping of indicator
lichens in 1997.
MATERIAL AND METHODS
Viljandi is a small town (22 000 inhabitants) in
Southern Estonia (Fig. 1). Viljandi is a holiday
town by the picturesque Lake Viljandi with Castle
Park and old wooden houses. As a county centre
Viljandi traditionally accommodates enterprises
involved in food production. Enterprises involved
in production of building materials are mainly
situated in Männimäe industrial district,
southwestern part of Viljandi.
Assessing air quality, we mapped the
distribution of indicator lichens and evaluated
the state of lichens.
38
Folia Cryptog. Estonica
In 1997 we conducted 252 analyses of lichen
groupings on 126 trees in Viljandi – 68 on smallleaved lime (Tilia cordata Mill.) and 58 on Norway
maple (Acer platanoides L.) bark of which has
relatively similar properties to that of lime. The
results of the study of lichen groupings in 1987
and 1991 were used for comparison. In 1987
we performed 94 analyses on 45 limes and 2
maples; in 1991 128 analyses were made on 57
maples and 7 limes.
Mapping the distribution of indicator lichens
on lime and maple, we used the plan of Viljandi
on the scale of 1:10 000, where the town is
divided into 500 × 500 m quadrats. In 1997
lichen groupings were analysed in 30 quadrats.
All lichens in two, mostly N and S expositions
at a height of 1.3 m on a tree were identified
and their cover in per cent was estimated using
a 20 × 20 cm grid, divided into hundred units
of 2 × 2 cm. Additionally, we identified lichens
on the trunk at a height of 0.5–2 m. Selecting
the substrate, we followed the standardization
requirements set for this method: we chose
faultless, upright, solitary trees with the average
circumference of 130 cm.
Indicator lichens are selected and grouped
according to differences in their relative toxicitytolerance, by comparing the frequency of lichens
growing on lime and data on the coexistence of
species in areas with comparatively different
pollution levels in Estonia (Liiv, 1989 a, b; 1992).
This selection of indicator species coincides
largely with the scale of indicator species
elaborated in other European countries by
different methods and used for the bioindication
of air quality (Skye, 1979; Hultengren et al.,
1991; Hosiaisluoma, 1994; Hobohm, 1994;
Wirth, 1995).
By the compilation of the lichen-zonal map
we based on the data of the distribution of the
following indicator lichens: Hypocenomyce
scalaris (Ach.) M. Choisy, Lepraria incana s.lat.,
Scoliciosporum chlorococcum (Graewe ex Stenh.)
Vêzda, Hypogymnia physodes (L.) Nyl. tolerant
to strong acidic air pollution (acidification
of the substrate); Pseudevernia furfuracea (L.)
Zopf., Platismatia glauca (L.) W.L. Culb. & C.F.
Culb., Tuckermannopsis chlorophylla (Willd.)
Hale, Vulpicida pinastri (Scop.) J.-E. Mattsson
& M.J. Lai tolerant to moderate acidic air
pollution; Phaeophyscia nigricans (Flörke)
Moberg, P. orbicularis (Neck.) Moberg, Physconia
enteroxantha (Nyl.) Poelt, Xanthoria candelaria
(L.) Th. Fr., X. parietina (L.) Th. Fr. tolerant to
strong alkaline dust pollution (eutrophication
of the substrate) and acidic air pollution;
Anaptychia ciliaris (L.) Körb., Melanelia
subargentifera (Nyl.) Essl., Physconia distorta
(With.) J.R. Laundon, Ramalina fastigiata (Pers.)
Ach., R. fraxinea (L.) Ach., R. pollinaria (Westr.)
Ach. tolerant to alkaline dust pollution but
sensitive to acidic air pollution.
RESULTS AND DISCUSSION
In 1997 we identified 58 lichen species on
lime and maple; the data on the distribution
of 46 species are presented in a manuscript
report (Liiv & Sander, 1997). In 1987, when
we studied lichen flora mostly on lime, we
identified 45 species in Viljandi, and in 1991 we
determined 55 species predominantly on maple.
Differences in the number of epiphytic lichens
in Viljandi are mainly caused by the tree species
under observation – the abundance of lichens
growing on the maple with the subneutral bark
is higher than on the lime with the subacidic
bark. This was also confirmed by the results of
the investigations of 1993–95 when we analysed
lichen groupings on 693 lime and 777 maple
trees growing outside the towns in parks of
estates and country cemeteries.
Basing on the distribution of indicator
lichens, we distinguished four zones in Viljandi
(Fig. 1).
Zone I, the zone of lichens tolerant to
strong acidic air pollution, is located in the
centre of Viljandi. The area of the zone is about
0.4 km2.
This zone is characterised by frequent
occurrence of the indicators of strong acidic
pollution H. scalaris, L. incana, S. chlorococcum
and H. physodes on trees with the subacidic
and subneutral bark. H. physodes grows on lime
and maple also outside the towns, in parks of
estates and country cemeteries (Table 1). The
other three extremely toxicity-tolerant lichens
H. scalaris, L. incana and S. chlorococcum have
not been recorded on lime and maple outside
the town areas.
Poleophobous species sensitive to acidic
pollution A. ciliaris, M. subargentifera, P.
distorta, R. fastigiata, R. fraxinea and R.
pollinaria, occurring commonly on lime and
39
Fig. 1. Lichen zones in Viljandi. I – zone of lichens tolerant to strong acidic air pollutions; II – zone
of lichens tolerant to moderate acidic air pollution; III – zone of lichens tolerant to strong dust
pollution; IV – zone of lichens sensitive to acidic air pollution.
maple in relatively unpolluted air outside the
towns, are completely lacking in zone I (Table 1,
2). Indicators of dust pollution are represented
in zone I by P. orbicularis, P. enteroxantha,
X. candelaria and X. parientina, but also by
other comparatively toxicity-tolerant lichens as
Buellia punctata (Hoffm.) A. Massal., Melanelia
exasperatula (Nyl.) Essl., Lecanora carpinea (L.)
Vain., L. chlarotera Nyl. and L. hagenii (Ach.) Ach.
growing on eutrophicated substrate. In all, we
identified 28 lichen species on lime and maple
in this zone. However, lichens showing tolerance
to acidic air pollution, e.g. Evernia prunastri and
Ramalina farinacea or indicators of moderate
acidic pollution P. furfuracea and T. chlorophylla,
but also more tolerant Parmelia sulcata and H.
physodes, are extremely tiny, strongly deformed
and hidden in bark cracks, being on the verge of
dying or already dead. The conditions are highly
suitable for H. scalaris, S. chlorococcum and L.
incana only. The mean general cover of lichens
on trees of the quadrats analysed is only 4% in
this zone.
40
Folia Cryptog. Estonica
Table 1. Frequency (%) of indicator lichens on lime and maple in Viljandi in 1987, 1991 and 1997,
and in Estonian parks outside the towns in 1993–95
Zone II, the zone of lichens tolerant
to moderate acidic air pollution, comprises
almost the entire northern part of Viljandi (Fig.
1). The area of the zone is about 6 km2 and it
takes up most of Viljandi’s territory.
This zone is characterised by the distribution
of P. furfuracea, P. glauca, T. chlorophylla and V.
pinastri, tolerant to moderate acidic pollution,
also H. physodes, very tolerant to acidic
pollution and P. orbicularis, P. enteroxantha, X.
candelaria and X. parietina which are tolerant to
alkaline dust and acidic air pollution, but also
by poleophobous A. ciliaris, M. subargentifera, P.
distorta, P. perisidiosa, R. fastigiata, R. fraxinea
and R. pollinaria. Tolerant to strong acidic
pollution H. scalaris and S. chlorococcum are
missing in this zone. In all, 52 lichen species have
been identified in zone II. Such a large number
of lichens should be put down to the cemetery
at Riia Road, but also to Viljandi Lossipark with
great biomass. In this area poleophobous lichens
grow side by side with indicators of moderate
acidic pollution. Thus zone II represents a socalled transitional zone for lichens, showing the
occurrence of poleophobous species as well as
indicators of acidic air pollution. H. physodes
has been identified even on 89% of trees (Table
2). Many dead lichens are found on trees as
well.
As a rule, the cover of lichens on trees is
higher in relatively unpolluted air. The mean
general cover of lichens is 20% in zone II, but the
highest value of 26% is recorded in zone IV.
Zone III, the zone of lichens tolerant
to strong dust pollution, spreads mostly in
two regions (Fig. 1). A 0.2 km2 area occurs
as an island in the zone of moderate acidic
pollution. The other dust pollution area of 2.2
km2 embraces most of the Männimäe industrial
and residential district.
On 89% of the trees examined in the socalled dust zone grow P. nigricans, P. orbicularis,
P. enteroxantha, X. candelaria and X. parietina,
tolerant to strong dust pollution and moderate
acidic pollution (Table 2). Indicators of acidic
pollution are represented in this zone by H.
physodes and L. incana on single trees, but H.
scalaris and S. chlorococcum, tolerant to strong
acidic pollution, and P. furfuracea, P. glauca,
41
Table 2. Frequency (%) of groups of indicator lichen zones of Viljandi in 1997
T. chlorophylla and V. pinastri, which are tolerant
to moderate acidic pollution, are lacking.
The total number of lichen species is 32 in
the dust zone, whereas on 44% of the trees grow
also poleophobous, tolerant to dust pollution
A. ciliaris, M. subargentifera, P. distorta, P.
perisidiosa, R. fastigiata and R. fraxinea, which,
however, are sensitive to acidic pollution.
Zone IV, the zone of lichens sensitive to
acidic air pollution, comprises the marginal
areas of the town (Fig. 1). In the zone of relatively
unpolluted air of Viljandi 30 lichens have been
identified. The main indicators of this zone
A. ciliaris, M. subargentifera, P. distorta, R.
fastigiata, R. fraxinea and R. pollinaria, which
tolerate weak dust pollution but are sensitive
to acidic pollution, grow on 78% of the trees
investigated for lichen groupings. The lichens
indicative of dust pollution, which occur
relatively frequently on lime and maple also
outside the town areas, grow in this zone even
on all trees (Table 1, 2).
The lichen flora on lime and maple of Viljandi
provokes a lot of interest as most of the town
falls in the zone of moderate acidic pollution,
where lichens of the trees with the subacidic
and subneutral bark “compete” with the lichens
characterising the trees with the acidic bark.
This accounts for a relatively large number
of lichens in zone II in comparison with the
zones of lichens tolerant to dust pollution (III)
or sensitive to acidic air pollution (IV). Lichens
are relatively numerous also in the most polluted
zone of Viljandi, as we considered as growing
every at least recognisable or damaged lichen
remain. A very expressive characteristic is the
comparatively low cover of lichens which makes
4% in this zone. Moreover, most of this value
represents the cover of crustaceous lichens P.
argena and L. incana tolerant to pollution, or
of H. physodes which is also very tolerant to
acidic pollution.
Considering the results of mapping of
indicator lichens in 1987 and 1991, we note
that the number of thalli of dead lichens as well
as of damaged and mutilated lichens was greater
in 1997. The location of the zones coincided in
all three years. As compared to the years of
1987 and 1991, the frequency of some lichens
42
Folia Cryptog. Estonica
indicative of acidic air pollution has increased
(Table 1). The distribution of H. scalaris, the
indicator of strong acidic pollution, has widened.
In 1987 this species was not identified in
Viljandi, in 1991 it grew on lime in the Park
of Coloured Fountains. The frequency of H.
physodes, L. incana and P. furfuracea has more
increased on lime. The reason of lower increase
on maple may be not only the differences of bark
of different tree species but also a shorter time
interval between maple-analyses (1991/1997)
compared with lime-analyses (1987/1997).
Among indicators of moderate acidic
pollution of air V. pinastri was not identified
on lime and maple in 1987 and 1991, but now
it occurs in three town quadrats. Analyses of
lichen groupings, performed mostly on maple
in 1991, did not show the occurrence of P.
furfuracea, which we identified on lime in two
town quadrats in 1987 and in six quadrats in
1997. The distribution of toxicity-tolerant H.
physodes has expanded considerably as well.
These facts indicate the growth of acidic air
pollution in Viljandi. Although for some years
lichens stay inert in response to changes in
air properties, we may affirm that air quality
is constantly deteriorating in Viljandi, as the
state of lichens has worsened practically in the
whole town. More sensitive lichens, of which
only fragments have preserved, are dying out.
The intact exterior and great increment of
lichens indicative of acidic pollution, however,
give evidence of moderate and not strong acidic
pollution in the most part of the town. Still,
young thalli of poleophobous lichens were
recorded nowhere in Viljandi. On the contrary,
their cover has decreased almost everywhere,
the state of lichens has deteriorated also on the
outskirts.
ACKNOWLEDGMENTS
Financial support to carry out this study has
been received from the Environmental Fund of
Estonia. The authors are grateful also to Agu
Eensaar and Anne Noor.
REFERENCES
Ammann, K. 1997. Flechten als Bioindikatoren. Uni
Press, 94: 18–25.
Eensaar, A. & Liiv, S. 1983. Viljandi õhu seisund. In
Väikelinnade ja maa-asulate keskkonnakaitse
ning maastikuhooldus. Ettek. teesid. Tallinn
– Viljandi. Lk. 104–107.
Hobohm, C. 1994. Baumflechten und Luftbelastungen
in Lüneburg und Umgebung – eine neue Methode
der Bioindikation auf der Basis von Zeigerwerten.
Beiträge zur Naturkunde Niedersachsens 47, 2:
49–61.
Hosiaisluoma, V. 1994. Pääkaupunkiseudun puistopuiden jäkälistö ilmanlaadun kuvaajana.
Pääkaupunkiseudun julkaisusarja C, 2: 1–52.
Hultengren, S., Martinsson & P-O., Stenström, J.
1991. Lavar och luftföroreningar. Känslighetsklassning och indexberäkning av epifytiska lavar.
Naturvårdsverket RAPPORT 3967. 113 pp.
Kirschbaum, U. 1995. Immissionsbezogene Flechtenkartierung von Hessen nach der neuen Richtlinie VDI 3799, Blatt 1. Staub – Reinhaltung der
Luft 55: 305–309.
Liiv, S. 1973. Linnade õhu saastatuse astme bioindikatsioonist. Diplomitöö. Tartu. 84 lk. Käsikiri
Tartu Ülikoolis.
Liiv, S. 1978. Lichenoindicational mapping of air pollution in Tartu and Viljandi (in Russian). In Studies on problems of ecology and rational exploitation
of natural resources, pp. 5–8. Tartu.
Liiv, S. 1987. Lichenoindication of air pollution in
towns in southern Estonia (in Russian). Dissertation. Tallinn. 230 pp.
Liiv, S. 1989 a. Kas samblikud tulevad tagasi? Eesti
Loodus 9: 554–560.
Liiv, S. 1989 b. Kas samblikud tulevad tagasi? Eesti
Loodus 10: 648–654.
Liiv, S. 1992. Eestin ilmanlaatu jäkäläkartoituksella
mitattuna. Ilmansuojelu-uutiset 1: 8–12.
Liiv, S. & Sander, E. 1996. Distribution of epiphytic
lichens indicating air pollution in Estonia. In:
Progress and Problems in Lichenology in the Nineties. Abstracts of the third symposium IAL 3. Salzburg, Austria 1–7 September 1996. P. 231.
Liiv, S. & Sander, E. 1997. Viljandi linna õhu seisundi
hindamine lihhenoindikatsiooni meetodil. Tallinn,
29 lk. Käsikiri Tallinna Botaanikaaias.
Skye, E. 1979. Lichens as biological indicators of air
pollution. Ann. Rev. Phytopathol. 17: 325–341.
Trass, H. 1968 a. Samblikud – õhu saastatuse indikaatorid. Eesti Loodus 2: 80–83.
Trass, H. 1968 b. Indeks samblikurühmituste kasutamiseks õhu saastatuse määramisel. Eesti
Loodus 10: 628.
Van Dobben, H. F. 1996. Decline and recovery of
epiphytic lichens in an agricultural area in The
Netherlands (1900–1988). Nova Hedwigia 62,
3–4: 477–485.
Wirth, V. 1987. Die Flechten Baden-Württembergs.
Eugen Ulmer Verlag. Stuttgart. 528 pp.
Wirth, V. 1995. Flechtenflora. Bestimmung und ökologische Kennzeichnung der Flechten Südwestdeutschlands und angrenzender Gebiete. Eugen Ulmer
Verlag. Stuttgart. 661 pp.
Folia Cryptog. Estonica, Fasc. 32: 43–46 (1998)
List of Estonian calicioid lichens and fungi
Piret Lõhmus
Institute of Botany and Ecology, University of Tartu, 38 Lai St., EE2400 Tartu, Estonia
Abstract: In the beginning of the 1970s 34 species of calicioid lichens and fungi were reported from Estonia. However,
after critical revision of herbarium material only 23 species, according to the present taxonomy, can be confirmed. In the
last two decades 25 new to Estonia species have been found and thus, altogether 48 species are verified in Estonia today. In
addition, two previously reported species are treated as doubtful because of the lack of vouchers. A map of the frequency
of the species in different parts of Estonia is presented, and species that may occur in Estonia are discussed.
Kokkuvõte: P. Lõhmus. Eesti kalitsioidsete samblike ja seente nimestik.
1970. aastate alguseks oli Eestist teada 34 kalitsioidse sambliku ja seene liiki (Trass, 1967, 1970; Sõmermaa, 1970). Pärast
herbaarmaterjalide kriitilist läbivaatust, arvestades muutusi süstemaatikas ning lisades hilisemad andmed, võib Eestist tõestatuks
lugeda 48 kalitsioidse sambliku ja seene liiki. Kahe liigi (Chaenotheca cinerea, Sclerophora peronella) esinemine Eestis jääb kaheldavaks,
kuna varasem herbaarmaterjal puudub ja uusi leide seni pole. Võrreldes Trassi (1967, 1970) ning Sõmermaa (1970) esitatud
nimestikega on lisandunud 25 liiki. Esmakordselt avaldatakse järgmised liigid: Calicium denigratum, C. parvum, Chaenotheca
gracillima, C. laevigata, C. subroscida, C. xyloxena, Chaenothecopsis consociata, C. debilis, C. epithallina, C. nana, C. pusiola, C. savonica,
C. viridireagens, Cyphelium sessile, Phaeocalicium populneum. Umbes 40% levikuruutudest (10’E–W × 6’N–S) on kogutud vähemalt
üks kalitsioidse sambliku või seene liik. Edasistel uuringutel võib praegustele lisanduda veel vähemalt kümme liiki.
INTRODUCTION
Trass (1967, 1970) and Sõmermaa (1970) reported a total of 34 species of calicioid lichens
and fungi from Estonia. During the last decades
new species have been added and some previously published species were redetermined. The
aims of this paper are (1) to check critically the
species in Trass (1967, 1970) and Sõmermaa
(1970), (2) to compile an updated list, (3) to
discuss the state of knowledge about Estonian
calicioid lichens and fungi, and (4) to present
the species that may occur in Estonia.
MATERIAL AND METHODS
The study is based mainly on the material of
calicioid lichens and fungi kept in the herbaria
of the University of Tartu (TU), the Botanical
Garden of Tallinn (TBA), the Institute of Ecology in Tallinn (IE), the University of Helsinki
(H) and the University of Riga (RIG). Altogether
over 1700 specimens were examined, 740 of
them were collected by the author within the
last three years.
The morphology and anatomy of the lichens were studied with a stereo lens and light
microscope, and the colour tests of apothecia
were made with 10% KOH, 50% nitric acid and
Lugol’s solution. For identification the following
publications were used: Foucard (1990), Purvis
et al. (1992), Tibell (1975, 1976, 1978, 1980).
Additional comparsion material was received
from H. The nomenclature follows Santesson
(1993), the abbreviations of authors’ names are
given according to Kirk & Ansell (1992).
The herbarium data were sorted with the
computer program BRAHMS. To illustrate
the distribution of collected material, a map
was compiled with the DMAP 6.4 program for
Windows. As a record unit a square 10'(E–W) ×
6'(N–S) was used.
Using the species lists of the 15 best studied
squares (with more than 10 species in each;
representing a total of 37 species) the rate of
discovery of new species is shown. For this purpose the lists were taken at random and for each
list the number of new species (not recorded in
previous lists) was determined.
RESULTS
After critical revision of herbarium material
only 21 species of calicioid lichens and fungi
presented in Trass (1967, 1970) and Sõmermaa (1970) can be confirmed according to the
44
Folia Cryptog. Estonica
present taxonomy. Furthermore, the occurrence
of four species (Chaenotheca cinerea, C. chlorella,
C. hispidula, Sclerophora peronella) could not be
confirmed, because voucher material was not
found. However, Chaenotheca chlorella and C.
hispidula have been collected later again. Two
additional species are not known from Estonia
with certainty. The following species were excluded from the lists in Trass (1967, 1970) and
Sõmermaa (1970) because of previous uncorrect
identification: Calicium corynellum was redetermined (®) as Microcalicium arenarium (det. V.
Räsänen; RIG), C. lenticulare ® C. abietinum and
C. glaucellum (det. P. Lõhmus; TU), all specimens (except three) of Chaenotheca chlorella
® Chaenotheca phaeocephala (det. P. Lõhmus;
TU), Chaenothecopsis faginea ® C. pusilla (det.
P. Lõhmus; TU), C. viridireagens ® C. epithallina
(det. A. Titov; TU), Phaeocalicium populneum ®
Mycocalicium subtile and Chaenothecopsis consociata (det. P. Lõhmus, A. Titov; TU). Some of
these taxa were later found as new to the Estonian lichen flora.
List of calicioid species in Estonia
A checklist of Estonian calicioid lichens and
fungi is compiled. New taxa for Estonia (compared with Trass 1967, 1970; Sõmermaa 1970,
taking into account redeterminations) are given
in bold; doubtful species are marked with ?. As
synonyms only these names are added which
have been used in Trass (1970) and Sõmermaa
(1970). In the right column the total number of
records in Estonia in 1998 is presented.
CALICIUM Pers.
ABIETINUM Pers.
49
ADSPERSUM Pers.
5
DENIGRATUM (Vain.) Tibell
1
GLAUCELLUM ACH.
60
PARVUM Tibell
13
QUERCINUM Pers.
18
SALICINUM Pers.
28
TRABINELLUM (Ach.) Ach.
22
VIRIDE Pers.
120
CHAENOTHECA (Th. Fr.) Th. Fr.
BRACHYPODA (Ach.) Tibell = Coniocybe sulphurea
(Retz.) Nyl.
19
BRUNNEOLA (Ach.) Müll. Arg.
16
CHLORELLA (Ach.) Müll. Arg. = C. carthusiae
(Harm.) Lett.
11
(Turner ex Ach.) Th. Fr. 118
(Pers.) Tibell = Calicium schaereri
DNot., C. schaereri (DNot.) Zahlbr.
1
FERRUGINEA (Turner & Borrer) Mig. = C. melanophaea (Ach.) Zwackh.
123
FURFURACEA (L.) Tibell = Coniocybe furfuracea
(L.) Ach.
67
GRACILLIMA (Vain.) Tibell = Calicium gracilis
(Nadv.) Oxn.
2
HISPIDULA (Ach.) Zahlbr.
4
LAEVIGATA Nádv.
3
PHAEOCEPHALA (Turner) Th. Fr.
37
STEMONEA (Ach.) Müll. Arg.
21
SUBROSCIDA (Eitner) Zahlbr.
10
TRICHIALIS (Ach.) Th. Fr.
93
XYLOXENA Nádv.
39
CHAENOTHECOPSIS Vain.
CONSOCIATA (Nádv.) A.F.W. Schmidt
8
DEBILIS (Turner & Borrer ex Sm.) Tibell
1
EPITHALLINA Tibell
3
NANA Tibell
4
PUSILLA (Ach.) A.F.W. Schmidt = Calicium alboatrum Flörke, Calicium pusillum Flörke
Calicium. subpusillum Vain.
35
PUSIOLA (Ach.) Vain.
1
RUBESCENS Vain.
2
SAVONICA (Räsänen) Tibell
4
SUBPAROICA (Nyl.) Tibell
1
VAINIOANA (Nádv.)Tibell
1
VIRIDIREAGENS (Nádv.) A.F.W. Schmidt = Calicium
viridireagens Nádv.
1
CYPHELIUM Ach.
INQUINANS (Sm.) Trevis.
7
LUCIDUM (Th. Fr.) Th. Fr.
1
SESSILE (Pers.) Trevis.
1
TIGILLARE (Ach.) Ach.
3
MICROCALICIUM Vain. emend. Tibell
ARENARIUM (Hampe ex A. Massal.) Tibell = Calicium arenarium Hampe
4
DISSEMINATUM (Ach.) Vain.
5
MYCOCALICIUM Vain.
SUBTILE (Pers.) Szatala = Calicium subtile
Pers.
86
PHAEOCALICIUM A.F.W. Schmidt
POPULNEUM (Brond. ex Duby) A.F.W. Schmidt =
Calicium populneum De Brond.
1
SCLEROPHORA Chevall.
CONIOPHAEA (Norman) Mattson & Middelb. 2
FARINACEA (Chevall.) Chevall.
4
NIVEA (Hoffm.) Tibell = Coniocybe pallida (Pers.)
Fr.
10
? PERONELLA (Ach.) Tibell = Coniocybe hyalinella
Nyl.
1
CHRYSOCEPHALA
?
CINEREA
45
SPHAEROPHORUS Pers.
GLOBOSUS (Huds.) Vain.
STENOCYBE (Nyl.) Körb.
PULLATULA (Ach.) Stein
THELOMMA A. Massal.
OCELLATUM (Körb.) Tibell
2
27
3
Thus there are 48 verified and two doubtful
species of calicioid lichens and fungi in Estonia
today. Most common species (over 50 localities)
are Calicium glaucellum, C. viride, Chaenotheca
chrysocephala, C. ferruginea, C. furfuracea, C.
trichialis, Mycocalicium subtile. Twelve species
– Calicium denigratum, Chaenotheca gracillima,
Chaenothecopsis debilis, C. pusiola, C. subparoica, C. vainioana, C. viridireagens, Cyphelium
lucidum, C. sessile, Phaeocalicium populneum,
Sclerophora coniophaea and Sphaerophorus
globosus – have been found only once or twice.
In the present list 25 species are new to
Estonia, compared to Trass (1967, 1970) and
Sõmermaa (1970). Eleven of these have already
been published by Randlane (1978, 1986;
Chaenotheca phaeocephala, Cyphelium inquinans), Ekman et al. (1991; Calicium glaucellum,
Chenothecopsis subparoica, C. vainioana, Microcalicium disseminatum, Sclerophora coniophaea,
S. farinacea, Thelomma ocellatum), Nilson et al.
(1997; Cyphelium lucidum) and Thor et al. (1998;
Cyphelium sessile). A.Titov, having checked
some TU herbarium material of calicioid lichens
and fungi in 1984, identified Chaenotheca subroscida, C. xyloxena, Chaenothecopsis consociata,
C. epithallina and C. pusiola for the first time.
My revisions of all herbarium material and additional fieldwork in 1995–1997 added Calicium
denigratum (leg. H. Ting, 1963), C. parvum (leg.
A.-L. Sõmermaa, 1967), Chaenotheca gracillima
(leg. P. Lõhmus, 1997), C. laevigata (leg.A.-L.
Sõmermaa, 1967), Chaenothecopsis debilis (leg.
I. Jüriado, 1996), C. nana (leg. P. Lõhmus, 1997),
C. savonica (leg. P. Lõhmus, 1996), C. viridireagens (leg. P. Lõhmus, 1997) and Phaeocalicium
populneum (leg. M. Sarv, 1995) to the list.
DISCUSSION
A map of the geographical frequency of all
calicioid lichens and fungi species in Estonia
is presented where the squares are numbered
according to the abundance of species in the
squares (Fig. 1). Altogether in ca. 40% of squares
at least one species has been collected, this indicates that thorough investigations have been
Fig. 1. The known distribution of calicioid species of lichens and fungi and the abundance of known
species in squares (1 = less than 6; 2 = 6–10; 3 = 11–15; 4 = 16–20; 5 = more than 20 species).
46
Folia Cryptog. Estonica
lane, L. Tibell and A. Lõhmus for criticism on
the manuscript. The study was undertaken as
part of the project “Composition and analysis of
Estonian lichen flora” financed by the Estonian
Science Foundation (Grant No. 1297).
REFERENCES
Fig. 2. The rate of discovery of new species in
the 15 best studied (with more than 10 species)
squares with a total of 37 species. For the analysis the squares were taken in random order.
done only in a few regions. The best studied
areas are Alam-Pedja Nature Reserve (22 species
are known in 260 km²), Endla Nature Reserve
(18 species in 81 km²) and Poruni primeval
forest with its surroundings (16 species in 0.5
km²). Preferred habitats for calicioid lichens
and fungi (mire complexes, old deciduous forests and abandoned homesteads) are typical for
these areas. The knowledge of calicioid flora in
central Estonia and mainland western Estonia
is still very poor and needs more attention in
the future research.
An analysis of the rate of discovery of new
species has been carried out in 15 best studied
squares (Fig. 2). After the first decline the rate
of adding new species has remained stable in
Estonia, i. e. a significant part of calicioid lichens and fungi seems to have been not found
yet. In southern Finland 63 species (Vitikainen
et al., 1997; biogeographical provinces No. 1–7)
and in Latvia 23 species (Piterans, 1982, pers.
comm.) are known. Considering the distribution
and habitats of the species in these lists, I guess
that at least Calicium adaequatum, C. lenticulare, Chaenothecopsis hospitans, C. viridialba,
Cybebe gracilenta, Cyphelium notarisii and
Sphinctrina anglica might be found in Estonia.
Probably about ten species are as yet not discovered in Estonia.
ACKNOWLEDGEMENTS
I am grateful to the curators of herbaria H ,
IE, RIG and TBA. My warm thanks are due to
A. Saag for his advices about the computer
programs BRAHMS and DMAP, and to T. Rand-
Ekman, S., Fröberg, L., Kärnefelt, J., Sundin, R., Thor,
G. 1991. New or interesting lichens from Estonia.
Folia Cryptog. Estonica 28: 5–25.
Foucard, T. 1990. Svensk skorplavs flora. Lund. 306
pp.
Kirk, P. M. & Ansell, A. E. 1992. Authors of fungal
names. C.A.B. International, Wallingford. 95 pp.
Nilson, E., Kannukene, L., Truus, L., Ratas, U., Puurmann, E., Tobias, M. 1997. Biological diversity. In
Small islands of Estonia (eds. Ratas, U. & Nilson,
E.), pp. 149–153. Tallinn.
Piterans, A. 1982. Lichens of Latvia (in Russian). Zinante, Riga. 352 pp.
Purvis, O. W., Coppins, B. J., Hawksworth, D. L.,
James, P. W. & Moore, D. M. (eds.) 1992. The
lichen flora of Great Britain and Ireland. Natural
History Publications, London. 710 pp.
Randlane, T. 1978. New lichen species to the Estonian
lichen-flora from the Viidumäe Nature Reserve
(Island Saaremaa). Folia Cryptog. Estonica. 11:
7–8.
Randlane, T. 1986. New species to the Estonia lichenflora. Folia Cryptog. Estonica. 21: 8–10
Santesson, R. 1993. The lichens and lichenicolous
fungi of Sweden and Norway. SBT -förläget,
Lund. 240 pp.
Sõmermaa, A. 1970. Epiphytic lichens found in main
forest types of Estonia. Pap. Bot. 9: 265–297.
Thor, G. & Nordin, A. 1998. 16 lichens new to Estonia.
Folia Cryptog. Estonica. 32: 123–125.
Tibell, L. 1975. The Caliciales of Boreal North America.
Symb. Bot. Upsal. XXI: 2.
Tibell, L. 1976. Calicium denigratum (Vain.) L. Tibell,
comb. nov. Notes on Caliciales 3. Bot. Notiser
129: 131–136.
Tibell, L. 1978.The genus Microcalicium. Bot. Notiser
131: 229–246.
Tibell, L. 1980. The lichen genus Chaenotheca in the
northern hemisphere. Symb. Bot. Upsal. XXIII:
1.
Trass, H. 1967. Analysis of the lichen-flora of Estonia
(in Russian). Doctoral dissertation. Tartu State
University.
Trass, H. 1970. The elements and development of the
lichen-flora of Estonia (in Russian). Pap. Bot. 9:
5–233.
Vitikainen, O., Ahti, T., Kuusinen, M., Lommi, S. &
Ulvinen, T. 1997. Checklist of lichens and allied
fungi of Finland. Norrlinia 6: 1–123.
Folia Cryptog. Estonica, Fasc. 32: 47–55 (1998)
Epiphytic macrolichens in Estonian forests
Ljudmilla Martin & Jüri Martin
International Center for Environmental Biology at Tallinn University of Educational Sciences,
PO Box 676, Tallinn EE 0026, Estonia
Abstract: Estonian forests were studied in 116 random sample plots to estimate epiphytic macrolichen distribution fre-
quency. 57 lichen species were recorded 1164 times on 2328 trees. Lichen species were divided into seven frequency classes.
Frequency class 1 (1–15 records) includes 42 species which belong to 5 geographic elements. Multiregional species Hypogymnia
physodes and Parmelia sulcata demonstrate the highest frequency. Several rare and floristically interesting macrolichen species
belong to frequency class 1: Alectoria sarmentosa, Bryoria furcellata (new species to Estonia), Hypocenomyce friesii, Lobaria pulmonaria,
Menegazzia terebrata, Physcia semipinnata, Usnea fulvoreagens, U. scabrata.
Kokkuvõte: L. Martin ja J. Martin. Epifüütsed suursamblikud Eesti metsades.
Eesti metsades uuriti epifüütsete suursamblike sagedust 116 juhuslikult valitud punktis. 57 samblikuliiki registreeriti 1164
korral 2328 puul. Samblikuliigid jagunesid seitsmesse sagedusklassi. Suurima liikide arvuga (42) oli esimene sagedusklass
(1–15 leidu). Need liigid kuuluvad viide geograafilisse elementi. Suurima sagedusega olid multiregionaalsed samblikuliigid
Hypogymnia physodes ja Parmelia sulcata. Esimesse sagedusklassi kuuluvad haruldased ja floristiliselt huvitavaid liigid Alectoria
sarmentosa, Bryoria furcellata (uus liik Eesti lihhenoflooras), Hypocenomyce friesii, Lobaria pulmonaria, Menegazzia terebrata, Physcia
semipinnata, Usnea fulvoreagens, U. scabrata.
INTRODUCTION
Estonian lichen flora is relatively well studied
and includes about 800 species (Trass, 1970).
The last review and identification key “Macrolichens of Estonia” (Trass & Randlane, 1994 )
includes records of 332 species of macrolichens.
Among them, 42 species have been recorded
before 1950 and are considered as extinct from
the Estonian lichen flora. In this review, 131
species are registered as epiphytic macrolichens
(excluding the genus Cladonia), and 20 of them
have not been recorded after 1950.
A. Sõmermaa (1972) published a monographic study on the ecology of epiphytic lichens
in the main Estonian forest types. In her study
the distribution of 26 epiphytic macrolichen species (excluding the genus Cladonia) in different
forests was described.
The latest review of forest lichens in Estonia
was presented by H. Trass (1996) in the study
of Estonian natural (old-growth) forests flora.
He presented data on the distribution and frequency of 95 lichen species that are considered
as indicators of virgin forests. This list includes
47 epiphytic macrolichen species.
The main goal of the present study is to
make an inventory of epiphytic macrolichen
flora in the Estonian forests and to evaluate
and quantify the frequency of epiphytic macrolichens. In 1994–1997 lichen data were collected under the co-operative Baltic–US Forest
Ecological Monitoring Project and Estonian State
Environmental Monitoring Program.
This is the first paper in a series of publications to analyze different aspects of lichen
species distribution in relation to structure
and composition of Estonian forests.
MATERIAL AND METHODS
The general methodology and the forest sample plot design are described in ”Forest Health
Monitoring Field Methods Guide (International)”
(1994, 1995, 1996) and in “Forest Ecological
Monitoring in the Baltic Countries” (1996). According to this methodology forest stand was
considered as a forested area of more than 0.4
ha and sample trees were >12.5 cm in diameter.
Sample plots in the forested areas were selected
by a randomized sampling grid (Overton et al.,
1990 ). The locations of sample plot centers
were estimated using satellite and topographic
maps. In the field the location of sample plot
was identified by a GPS unit.
48
Table 1. Macrolichen species recorded for Estonian forests
Folia Cryptog. Estonica
49
50
Folia Cryptog. Estonica
Fig. 1. Forest epiphytic macrolichens sample plots.
The lichen sample area (“lichen plot”) was a
circular area with 34.7 m radius (0.378 ha)
around the forest sample plot center. On each
lichen plot the occurrence of the lichen species
on different tree species (at least 10 trunks of
each tree species) was noted. Macrolichens were
recorded at a height of more than 0.5 m from the
tree base up to 2.5 m and separately for twigs of
Picea abies. In addition, six sampled forest sites
along the air pollution gradient crossing Estonia from northeast to south-west (Martin et al.,
1994) were included in the database (Table 1). In
this study 2328 tree trunks were observed. The
lichen species were recorded on fourteen tree
species including Coryllus avellana, Malus sp.
and Juniperus communis. The most frequent tree
species in this sample was Pinus sylvestris (71
plots) and the least frequent Tilia cordata (1 plot).
Location of the 116 sampled forest sites is shown
in Fig. 1. During the field work, more than 1000
lichen samples were collected for identification
in the laboratory. The nomenclature presented
by R. Santesson (1993) was followed.
Similarity of epiphytic lichen cover species
composition registered on more frequent phorophytes was calculated by Sørensen’s similarity
coefficient.
The classification of geographical elements
proposed by H. Trass (1970) was used for macrolichen flora characterization.
The collected specimens were deposited in
the Herbarium of the International Center for
Environmental Biology, Tallinn (ICEB).
RESULTS
In the studied forests 57 macrolichen species
were identified and total number of macrolichen records selected by substrate types (phorophytes) was 1164 (Table 1). The highest similarity in lichen vegetation was found between
Pinus sylvestris trunks and Picea abies twigs
(Table 2). The lowest similarity value was between Picea abies twigs and Populus tremula.
Using the data presented in Table 1 the
lichen species frequency classes’ intervals were
calculated using the following formula:
where c is frequency class interval, xN – maximal variant (99, number of records for Parmelia sulcata), x1 – minimal variant (x1 = 1) and N
– number of observations (number of species,
N = 57). The calculated class interval was 15
51
Table 2. Similarity of lichen vegetation species composition (Sørensen’s similarity coefficient) on
different sampled phorophytes
Picea. abies, twigs
Pinus sylvestris
Picea abies, trunks
Betula spp.
Alnus incana
Alnus glutinosa
Populus tremula
Picea abies,
twigs
Pinus
sylvestris
1
0.800
0.760
0.710
0.512
0.378
0.213
1
0.627
0.698
0.500
0.368
0.250
(14.98). Frequency class 1 was further divided
into two subclasses: 1a (1 record) and 1b (2–15
records).
Using the data presented in Table 1 and
the calculated frequency class intervals a lichen
species frequency distribution curve was created
(Fig. 2). The recorded epiphytic lichen species
were divided into 7 classes.
Several rare and floristically interesting lichen species are included in frequency class 1.
Some of the new findings of these species are
described below.
Picea
abies,
trunks
Betula
spp.
Alnus
incana
Alnus
glutinosa
Populus
tremula
1
0.621
0.513
0.364
0.233
1
0.510
0.356
0.400
1
0.538
0.500
1
0.333
1
Lichen species in frequency class 1a ( 10
species, 18%) are as follows. Alectoria sarmentosa*, a very rare holarctic epiphyte, in Estonia
previously recorded once in 1930. In the current study this lichen species was found on
Picea abies twigs (North-East Estonia, Alutaguse, Kivinõmme, HEX 104386, 59°08’48”N,
27°30’18”E) in spruce-pine-birch mixed stand,
age >120 years (Fig.1.).
Lobaria pulmonaria*, a nemoral species,
recorded locally over the whole territory of Estonia on deciduous trees, mostly in broad-leaved
Fig. 2. Frequency distribution of the epiphytic macrolichen species in the studied forests.
* Asterisk indicates lichen species, which are included in the draft proposal for the Red Data Book
of Estonia.
52
Folia Cryptog. Estonica
mixed spruce forests. In this study, L. pulmonaria was recorded once on Populus tremula
(North-East Estonia, Oonurme, HEX 106135,
59°07’21”N, 26°57’46”E) in spruce-pine-birchaspen mixed stand, age >120 years.
Usnea fulvoreagens, a boreal lichen, has
been found earlier in Estonia in less than 20
localities. In this study U. fulvoreagens was
found on Picea abies (Vetla, HEX 111451,
59°12’23”N, 25°29’19”E) in spruce-pine stand,
age >120 years.
Usnea scabrata*, a boreal lichen, has been
recorded earlier in Estonia for more than 20
localities. In present study U. scabrata was recorded on Alnus glutinosa (Hageri–Adila, HEX
115043, 59°09’04”N, 24°35’15”E).
Frequency class 1b (32 species, 56%)
includes also some rare lichen species in Estonia. First of all, Bryoria furcellata – new for
the lichen flora of Estonia species (Randlane
et al., 1997) and up to now has been recorded
7 times for 6 localities in Estonian forests
(Aegna: 59°34’31”N, 24°45’49”E; 59°34’52”N,
24°45’25”E; Naissaar, 59°34”54”N, 24°31’10”E;
Nõva, 59°13’53”N, 23°38’21”E ; Tallinn, Kloostrimetsa, 59°28’12”N, 25°02’16”E; HEX 113220,
59°32’08”N, 25°36’16”E): 5 records on Pinus sylvestris, 1 on Betula pubescens and 1 on Picea
abies twigs. This species was always found in
similar habitats in open forests on sandy soils
in the coastal zone ( Fig. 1 ).
Hypocenomyce friesii, a boreal lichen, and
in Estonia recorded earlier for three localities.
In the current sample this lichen was also recorded three times for Pinus sylvestris (Aegna,
59°34’52”N, 24°45’25”E; 59°34’49”N, 24°
45’46”E; Naissaar, 59° 34”54”N, 24° 31’10”E).
Menegazzia terebrata*, a rare submontaneous lichen, was found on the same forest plot
as A. sarmentosa on Betula pubescens, Alnus
glutinosa, and Salix caprea (Fig.1). Several
interesting and relatively rare microlichen species were also collected here, i.e. Dimerella pineti
(Ach.) Vêzda, Lecanactis abietina (Ach.) Körb.,
Mycoblastus sanguinarius (L.) Norman, etc. M.
terebrata is considered as endangered in Europe
(Trass, 1997). In Estonia this lichen was earlier
known from 6 localities.
Physcia semipinnata*, a nemoral epiphytic
lichen, usually found on trunks of broad-leaved
trees (less than 10 records in Estonia). In this
study P. semipinnata was found on three forest
plots in South-Eastern Estonia (Fig.1): twice
on Populus tremula ( HEX 99245, 58°00’18”N,
27°08’37”E; HEX 100966, 58°08’51”N, 26° 46’
00”E) and once on Salix caprea (HEX 102700,
57°57’03”N, 26°05’43”E).
In addition to the species listed above, in frequency class 1 were included: (1) lichen species
which usually grow in old forests on conifers and
Betula spp. – Bryoria capillaris , B. fuscescens
, B. nadvornikiana , B. subcana , Hypogymnia
farinacea , Melanelia olivacea, Parmeliopsis
hyperopta, Tuckermannopsis sepincola, Usnea
filipendula; (2) species usually inhabiting on
deciduous trees such as Anaptychia ciliaris,
Melanelia exasperata, M. exasperatula, M.
fuliginosa, M. subargentifera , M. subaurifera,
Phaeophyscia ciliata, P. nigricans , P. orbicularis,
Physcia adscendens, P. aipolia, P. stellaris, P.
tenella , Physconia distorta, P. enteroxantha,
P. perisidiosa, Ramalina baltica, R. fastigiata,
R. fraxinea, R. pollinaria , Xanthoria fulva, X.
parietina, X. polycarpa.
In some cases listed species were found
on not typical for them substrata (for example,
Physcia tenella on Pinus sylvestris, on twigs and
trunk of Picea abies, Xanthoria parietina on P.
sylvestris, X. fulva on P. abies).
Frequency class 2 (4 species, 7%) includes
lichen species that are locally common in different habitats but relatively rare in the sampled forests: Evernia prunastri, Hypocenomyce
scalaris , Tuckermannopsis chlorophylla, Usnea
subfloridana.
Frequency class 3 (3 species, 5%) includes
lichen species common in old coniferous forests
Hypogymnia tubulosa, Imshaugia aleurites and
Ramalina farinacea found on twigs of spruces
as well on trunks of deciduous trees.
Frequency class 4 (1 species, 2%) has only
one relatively frequent species for Picea abies,
Pinus sylvestris and Betula spp. – Usnea hirta.
Frequency class 5 (3 species, 5%) includes
group of species that are typical for forests:
Platismatia glauca, Pseudevernia furfuracea,
Vulpicida pinastri.
Frequency class 6 (2 species, 4%) includes a
frequent for several phorophytes lichen Parmeliopsis ambigua, and several species of Cladonia
usually inhabiting on tree base but sometimes
climbing up on trunk.
Frequency class 7 (2 species, 4%) includes
multiregional and very frequent in forests Hypogymnia physodes and Parmelia sulcata.
53
Fig. 3. Distribution of the epiphytic macrolichen species belonging to different geographic elements
in the studied forests.
The sampled forests epiphytic macrolichen
lichen flora (Fig. 3, column 1) consists of 39%
boreal species, 30% nemoral species, 18% multiregional species, 9% holarctic species, and 4%
submontaneous species.
Lichen species in the frequency class 1
belong to the 5 geographic elements (Fig. 3,
column 2). In the higher frequency classes (2
to 7) species of the submontaneous geographic
element were absent and the percentage of boreal and multiregional species was higher (Fig.
3, column 3).
DISCUSSION
Lichens are considered as excellent biomonitors
of change in air quality, climate change, and
change in the structure of the forest community.
Indicative value of lichen communities variables,
such as presence/absence, frequency, coverage,
etc. has been recognized by many researchers
(Alstrup, 1996; Goward, 1992; Kuusinen, 1994,
1996; McCune, 1993; Smith, et al., 1993; Trass,
1996).
Decreasing area and continuity and increasing fragmentation of forests due to forest
management have led to the situation where
many forest lichen species are in danger. For example, about half of the 138 endangered lichen
species in Finland belong to the forest lichens (
Kuusinen, 1996).
H. Trass (1997) presented the list of lichen
species and their frequencies in 5 grade scale
found in natural forests in Estonia. Most of macrolichens included in this list were recorded in
present study but with lower frequency values
(mostly in our frequency classes 1 and 2). It
means that so-called indicator or typical species
of the old or natural forests become rare in other
forested areas. For example, such species are:
Bryoria capillaris, B. fuscescens, Hypogymnia
farinacea, Lobaria pulmonaria, Melanelia fuliginosa, M. olivacea, Menegazzia terebrata, Parmeliopsis hyperopta, Tuckermannopsis sepincola, Usnea fulvoreagens, U. scabrata. From 21 common
species in these two studies only Platismatia
glauca and Vulpicida pinastri had comparable
values of the frequency.
Although the lichen species distribution and
vegetation of the boreal forests has been studied
quite extensively in Northern Europe, there is
limited number of published data with which
the present results can be compared.
P.-A. Esseen (1981) found 18 macrolichens
on Pinus sylvestris and 24 on Picea abies in
Central Sweden (middle and southern boreal
subzones).
54
Folia Cryptog. Estonica
A. Sõmermaa (1972) surveyed the epiphytic
lichen vegetation of several forest types in Estonia (hemiboreal zone, Ahti et al., 1968), where
the number of epiphytic macrolichen species
(except Cladonia) on P. sylvestris was 23, P.
abies 19, on Betula pendula and B. pubescens
17, and 14 on Alnus glutinosa.
M. Hyvärinen et al. (1992) described 32
macrolichen species on P. sylvestris including
14 Cladina and Cladonia species, and 23 on P.
abies including 8 Cladonia species.
Differences between the total numbers of
epiphytic macrolichen species recorded by A.
Sõmermaa, P.-A. Esseen and M. Hyvärinen et
al. and in present study are relatively small,
approximately 7 lichen species per tree species
in average.
At the same time the comparison of the
species percentage frequencies established in
present study and presented by M. Hyvärinen
et al. (1992) for Muhos (Oulu region, the middle
boreal zone) showed considerable differences.
From 14 common species only three found on
pines (Hypogymnia physodes, Parmeliopsis
ambigua, Platismatia glauca) and two found
on spruces (H. physodes, P. sulcata) had close
frequencies. In central part of Finland frequency
values for most of the macrolichen species were
higher than estimated in present study.
These differences could be explained partly
by differnces in study site selection. M. Hyvärinen et al. (1992) as well A. Sõmermaa (1972)
studied typical forests sites. In present study all
sites were selected randomly and several forest
plots were placed in transition areas.
The epiphytic lichen vegetation is known to differ
greatly between branches and trunks of conifers
(Pike et al., 1975, 1977). The trunks of P. abies
receive less light, which could be the main reason for the greater abundance of macrolichens
on their branches than on the trunks. We found
in literature just one publication concerning lichen distribution on P. abies branches (Hilmo,
1994). She recorded 16 macrolichen species on
P. abies branches in central Norway. Comparison
is difficult because of Bryoria and Usnea species
were not identified and she studied mean cover
of the lichen species on branches. In present
study 27 macrolichen species were found on
P. abies twigs. It is interesting to point out that
the similarity between macrolichen species on
the P. abies twigs and P. sylvestris trunks was
higher than between P. abies twigs and trunks
(Sørensen’s similarity coefficients respectively
0.8 and 0.76 ).
Most of the recorded lichen species ( 42
species or 74 %) had relatively low frequency
(class 1). The most frequent lichen species in
sampled forests belonged to the nemoral, boreal
and multiregional geographic elements, which
may be due to their wide amplitude to habitat
conditions.
ACKNOWLEDGMENTS
The authors are grateful to colleagues H. Pärn
and M. Temina for their cooperation during the
field work. Also we owe thanks to T.Randlane,
University of Tartu , for consultations in identification of some Usnea taxa and Oleg Blum, Central Botanical Garden of the Ukrainian National
Academy of Sciences, for fruitful discussion of
Ramalina species. The financial support has
been received from the Bowling Green State
University, US EPA, US DA Forest Service, and
from the Estonian Ministry of the Environment
to carry out Forest Monitoring Program.
REFERENCES
Ahti, T., Hämet-Ahti, L. & Jalas, J. 1968. Vegetation
zones and their sections in northwestern Europe.
Ann. Bot. Fenn. 5: 169–211.
Alstrup, V. 1996. Natural forests and endangered
lichens in Denmark. In IAL 3: Progress and
Problems in Lichenology in the Nineties, p. 43.
Salzburg.
Esseen, P.-A. 1981. Host specifity and ecology of
epiphytic macrolichens in some central Swedish
spruce forests. Wahlenbergia 7: 73–80.
Forest Ecological Monitoring in the Baltic Countries.
1994. (Martin, J., ed.). 1996. Tallinn. 106 pp.
Forest Health Monitoring Field Methods Guide (International). US DA Forest Service, 1994, 1995,
1996.
Goward, T. 1992. Epiphytic lichens going down with
trees. In Proceedings of the Symposium “Community Action for Endangered Species”, pp. 153–158.
Vancouver, B.C., Canada.
Hilmo, O. 1994. Distribution and succession of
epiphytic lichens on Picea abies branches in a
boreal forest, central Norway. Lichenologist 26
(2): 149–169.
55
Hyvärinen, M., Halonen, P.& Kauppi, M. 1992. Influence of stand age and structure on the epiphytic
lichen vegetation in the middle-boreal forests of
Finland. Lichenologist 24: 165–180
Kuusinen, M. 1994. Epiphytic lichens flora and
diversity on Populus tremula in old-growth and
managed forests of southern and middle boreal
Finland. Ann. Bot. Fenn. 31: 245–260.
Kuusinen, M. 1996. Effect of forestry on epiphytic
lichen diversity in boreal forests of Finland. In
IAL 3: Progress and Problems in Lichenology in the
Nineties, p. 48. Salzburg.
Martin, J., Aaspõllu, J., Jaenes, A., Martin, L., Mehtijeva, N., Piin, T., Pärn, H., Tekko S., Vilde,
R. & Virolainen, V. 1994. Atmospheric pollution
deposition gradient studies in Estonia. In Climate
and Atmospheric Deposition Studies in Forests (eds
Solon, J., Roo-Zielinska, E. & Bytnerowicz, A.),
pp. 117–136.
McCune, B. 1993. Gradients in epiphyte biomass in
three Pseudotsuga-Tsuga forests of different age
in Western Oregon and Washington. Bryologist
96: 405–411.
Overton, W. S., White, D. & Steven, D. L. 1990. Design
report for EMAP, Environmental Monitoring and Assessment Program, I.
Pike, L. H., Denison, W. C., Tracy, D., Sherwood, M.
& Rhoades, F. 1975. Floristic survey of epiphytic
lichens and bryophytes growing on living, oldgrowth conifers in western Oregon. Bryologist
78: 391–404.
Pike, L. H., Rydell, R.A. & Denison, W. C. 1977. A
400-year-old Douglas fir tree and its epiphytes:
biomass, surface area and their distribution. Can.
J. Forest Res. 7: 680–699.
Randlane, T., Jüriado, I., Martin, J., Martin, L., Nilson, E. & Temina, M. 1997. Lichens of Naissaar
Island (Gulf of Finland, Estonia). Folia Cryptog.
Estonica 31: 20–25.
Santesson, R.1993. The lichens and lichenicolous fungi
of Sweden and Norway. Lund. 240 pp.
Smith, C., Geiser, L., Gough, L., McCune, B., Ryan,
B. & Showman, R. 1993. Species and communities. Chapter 4. In Lichens as Bioindicators
of Air Quality. US DA Forest Service Gen. Tech.
Rep. RM-224.
Sõmermaa, A. 1972. Ecology of epiphytic lichens in
main Estonian forest types. Scripta Mycol. 4:
1–117.
Trass, H. 1970. The elements and development of the
lichen flora of Estonia (in Russian). Pap. on Bot.
9: 5–233.
Trass, H.1996. Eesti põlislooduse taimekoosluste
bioindikatsioon: teoreetilised alused, meetodid
ja koosluste seisundi analüüs. Tartu Ülikooli
Botaanika ja ökoloogia instituut, Eesti Teadusfond. (Manuscript, 47 pp.).
Trass, H. 1997. Lichen mapping in Europe: Letharia
vulpina, Menegazzia terebrata. Proc. Estonian
Acad. Sci. Biol. Ecol. 46: 4, 195–213.
Trass, H. & Randlane, T. (eds.) 1994. Eesti suursamblikud. Tartu. 399 pp.
56
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 57–62 (1998)
Lichens of the islets of Kolga bay (Gulf of Finland, Estonia). I.
Distribution and frequency of epiphytic lichen species
Eva Nilson1 and Taimi Piin2
Institute of Ecology, Tallinn University of Educational Sciences, 2 Kevade St., EE0001 Tallinn, Estonia
2
Tallinn Botanic Garden, 52 Kloostrimetsa Rd., EE0019 Tallinn, Estonia
1
Abstract: Distribution and frequency of the most common epiphytic lichen species on five islets of Kolga bay (Gulf of
Finland, Estonia) are described based on floristic data. The most wide-spread and frequent lichen species are those indicating
eutrophication of bark substrate. Commonly overlooked crustose species are shown to be quite frequent, whereas some more
easily detectable species common for pine in the mainland part of Estonia are rarely met on the islets studied. The similarity
of epiphytic lichen flora of the islet pairs is briefly discussed, as compared to the data on vascular plants and bryophytes.
Kokkuvõte: E. Nilson ja T. Piin. Kolga lahe väikesaarte samblikud. I. Epifüütsete samblikuliikide levik ja
sagedus.
Floristiliste andmete alusel kirjeldatakse epifüütsete samblike levikut ja sagedust Kolga lahe väikesaartel. Kõige sagedamini
esinevad neil saartel epifüütsed liigid, mis viitavad puukoore eutrofeerumisele. Mitmed kooriksamblike liigid, mis kogumisel
võivad jääda märkamata ja mida seetõttu on peetud harvaesinevaiks, osutusid materjali läbitöötamisel üsna sagedasteks,
seevastu mõned Eesti mandriosas tavalised männil kasvavad samblikuliigid on Kolga lahe väikesaartel suhteliselt haruldased.
Lühidalt on võrreldud saarte soon- ja sammaltaimede ning epifüütsete samblike floora sarnasust saarepaaride vahel.
INTRODUCTION
The lichen flora of the islands at the northern
coast of Estonia was very poorly known until
recently. During the Soviet time the islands
belonged to the closed frontier zone and were
nearly inaccessible for scientists. Up to now,
the lichen species list is published for Naissaar
(Randlane et al., 1997).
Kolga bay at the northern coast of Estonia
has the greatest concentration of small islands
(islets) in this area (Figure 1). The islets are
situated between 59°30’–59°40’N and 25°10’–
25°30’E. They belong to the landscape region of
North-Estonian Coastal Lowland (Varep, 1964)
and to the corresponding geobotanical region of
Northern Maritime Estonia – Estonia maritima
borealis (Lippmaa, 1935). Administratively the
islets belong to the municipalities of Jõelähtme
and Kuusalu, Harjumaa County.
The islets lie on Vendian and Cambrian sedimentary rocks covered with Quaternary deposits
of various composition (Karukäpp & Malkov,
1993). The prevalent landforms are plains, escarpments, terraces and dunes. Northern and
western coasts of the islets are characterised
by presence of escarpments and coarse-grained
Fig. 1. Location of the islets of Kolga bay.
58
Folia Cryptog. Estonica
sediments (cobbles, boulders), while southern
and eastern coasts are normally flat and sandy
(Lepland, 1995). Parent material of soils (marine
and aeolic sands, coastal deposits consisting
much crystalline pebble) is poor in calcareous
matter. Soils are mainly podzols, sod-podzolic
soils, gley-podzols and saline littoral soils. Species diversity of flora and vegetation types of
the islets depend on age, area, elevation a.s.l.
and landscape diversity of the islets, as well as
on their management history and the influence
of seabird colonies (Ratas et al., 1995). Smaller
islets (Allu, Põhja-Malusi, Vahekari and Umblu)
carry heavily bird-influenced, fragmentary and
sparse herbaceous vegetation. Various types of
woody vegetation that make possible the occurrence of epiphytic lichens are met on five bigger
islets (Lõuna-Malusi, Rohusi, Koipse, Pedassaar
and Rammu).
A grove of young aspen trees (Populus tremula L.), some older trees of mountain ash (Sorbus
aucuparia L.) and Scots pine (Pinus sylvestris
L.), as well as a number of juniper (Juniperus
communis L.) and elder (Sambucus racemosa L.)
shrubs are met on Lõuna-Malusi. One third of
the area of Rohusi island is covered with pine
forest with the undergrowth of mountain ash
and elder. Old trees of mountain ash, aspen,
black alder (Alnus glutinosa L.) and goat willow
(Salix caprea L.) are met on forest edges near
the seashore. Woody vegetation is represented
mostly by juniper scrub on Koipse. A grove of
deciduous trees on this island consists of aspen and birch (Betula pubescens Ehrh.), while
conifers are represented by some single trees of
Scots pine and Norwegian spruce [Picea abies
(L.) Karst.]. The Calluna – Empetrum heath in
the southern part of the islet has been planted
with pine seedlings in 1987–88. Pedassaar, the
second largest islet of the group examined, has
nearly 98% of its territory covered with dry pine
forests with the undergrowth of mountain ash
and juniper. Black alder, aspen, mountain ash
and goat willow are met on forest edges. The
largest islet, Rammu, carries two small pine
groves of natural origin, surrounded by developing juniper scrub – one around a little lakelet
in the central part and another on a long sandy
spit extending to southeast. Deciduous trees are
very scarce on this islet. The Empetrum heath
in the eastern part of the islet is afforested with
pine seedlings.
For the islets of Kolga bay the following lichen species are mentioned in earlier literature:
Cladina arbuscula (Wallr.) Hale & W. L. Culb.,
Cladina rangiferina (L.) Nyl., Cladonia fimbriata
(L.) Fr. (Rebassoo, 1987); Cetraria islandica (L.)
Ach., C. nivalis (L.) Ach., Tremolecia atrata (Ach.)
Hertel, Xanthoparmelia mougeotii (Schaer. ex D.
Dietr.) Hale (Nilson & Piin, 1993); Cyphelium
lucidum (Th. Fr.) Th. Fr. and Peltigera scabrosa
Th. Fr. (Nilson et al., 1997).
In the 1990s the Kolga Bay Islets’ Nature
Reserve (consisting of ten islets) was established
to protect breeding bird populations and rare
or interesting plant communities. For conservation of any group of organisms basic information on distribution diversity and dynamics
is needed. Representative data on the frequency
of epiphytic lichens is of key importance when
these organisms are used as bioindicators of air
pollution or when Red Lists are to be established
Table 1. Area, elevation a.s.l. (Loopmann, 1996) and species number of vascular plants (Ploompuu,
1995), bryophytes (Ratas et al., 1995) and epiphytic lichens of the islets of Kolga bay
Islet
Rammu
Pedassaar
Koipse
Rohusi
Lõuna-Malusi
Põhja-Malusi
Allu
Umblu
Vahekari
Area, ha
Max elevation, m
Vascular plants
Bryophytes
Epiphytic lichens
102.6
90.5
34.3
12.5
7.0
3.1
1.6
1.6
0.6
4
13
7
6
6
3
2
3
2
229
161
189
175
126
72
57
50
36
68
58
65
44
4
2
0
0
0
29
79
68
74
35
0
0
0
0
Table 2. Distribution and frequency of the most common epiphytic lichen species on the islets of Kolga bay on different phorophytes
Lichen species
N
Phorophyte*
Ra
Pe
Ko
Ro
LM
Pi
Ju
Al
Be
30
6
7
5
10
10
5
3
8
5
3
60
121
82
72
43
27
33
24
15
30
19
24
9
2
28
8
62
63
46
40
28
28
19
9
22
21
30
4
4
14
1
4
9
7
7
8
8
2
2
4
3
77
98
60
46
36
53
19
32
29
16
15
30
32
12
10
22
8
8
8
8
7
12
19
12
15
17
66
48
148
88
37
21
3
42
56
26
19
26
2
3
1
12
21
32
40
17
21
27
26
34
11
22
1
5
2
4
1
1
3
7
1
1
3
2
1
1
6
3
1
2
2
8
17
17
17
16
16
6
3
3
5
4
3
1
14
1
12
3
1
2
39
23
19
21
12
5
1
1
18
4
6
18
1
8
4
1
3
4
19
28
56
20
42
14
5
11
10
24
20
1
19
3
3
1
1
4
3
3
7
2
1
3
14
3
8
3
1
19
10
7
4
6
2
6
6
2
3
6
2
Po Sal Sam
15
8
11
5
7
8
12
6
3
8
6
3
3
2
27
10
10
16
14
14
8
14
15
5
7
7
3
1
1
3
1
2
5
3
3
2
1
So
Sy
10 107
1 48
2 31
4 55
2 54
7 69
61
39
4 37
1 15
3
5
2
6
3
1
6
2
5
2
1
1
6
1
2
1
11
23
1
4
18
11
16
11
6
4
18
15
10
1
1
1
1
8
8
1
59
Xanthoria polycarpa (Hoffm.) Th. Fr. ex Rieber 295
Hypogymnia physodes (L.) Nyl.
288
Scoliciosporum chlorococcum (Graewe ex Stenh.) Vêzda 196
Parmelia sulcata Taylor
165
Lecanora hagenii (Ach.) Ach.
156
Physcia tenella (Scop.) DC.
141
Lecanora carpinea (L.) Nyl.
95
Lecidella elaeochroma (Ach.) M. Choisy
89
Physcia stellaris (L.) Nyl.
86
Lecanora symmicta (Ach.) Ach.
77
Lecanora pulicaris (Pers.) Ach.
67
Lecanora expallens Ach.
59
Ramalina farinacea (L.) Ach.
49
Xanthoria parietina (L.) Th. Fr.
48
Catillaria nigroclavata (Nyl.) Schuler
44
Amandinea punctata (Hoffm.) Coppins & Scheid. 41
Xanthoria candelaria (L.) Th. Fr.
41
Melanelia olivacea (L.) Essl.
35
Rinodina pyrina (Ach.) Arnold
34
Hypogymnia tubulosa (Schaer.) Hav.
33
Usnea hirta (L.) Weber ex Wigg.
31
Evernia prunastri (L.) Ach.
30
Melanelia exasperata (De Not.) Essl.
29
Ramalina fraxinea (L.) Ach.
29
Rinodina sophodes (Ach.) A. Massal.
25
Tuckermannopsis chlorophylla (Willd.) Hale
22
Tuckermannopsis sepincola (Ehrh.) Hale
18
Melanelia exasperatula (Nyl.) Essl.
17
Physcia dubia (Hoffm.) Lettau
15
Island
60
Lichen species
N
Island
Ra
Cliostomum griffithii (Sm.) Coppins
13
Physcia adscendens (Fr.) H. Olivier
13
Vulpicida pinastri (Scop.) J.-E. Mattsson & M. J. Lai 13
Lecanora chlarotera Nyl.
12
Phaeophyscia orbicularis (Necker) Moberg
11
Lecanora varia (Hoffm.) Ach.
8
Parmeliopsis ambigua (Wulfen) Nyl.
8
Chaenotheca ferruginea (Turner & Borrer) Mig.
7
Pseudevernia furfuracea (L.) Zopf
7
Ramalina fastigiata (Pers.) Ach.
7
Cyphelium lucidum (Ach.) Ach.
6
Melanelia fuliginosa (Th. Fr.) Th. Fr.
6
Bryoria fuscescens (Gyelnik) Brodo & D. Hawksw. 5
Lecanora leptyrodes (Nyl.) Degel.
5
Physconia distorta (With.) J.R. Laundon
5
1
2
Pe
1
4
3
1
8
5
4
2
5
2
Phorophyte*
Ko
Ro
LM
Pi
3
5
3
5
8
3
4
4
2
7
4
6
8
1
2
2
6
4
1
2
1
1
1
5
4
1
3
5
Ju
6
8
7
7
5
6
1
3
Al
1
Be
1
1
Po Sal Sam
2
1
3
7
3
3
2
1
2
4
2
So
Sy
5
8
4
4
1
1
2
1
1
1
2
3
2
Abbreviations:
N, total number of records (both as main and accompanying species)
Islets: Ra, Rammu; Pe, Pedassaar; Ko, Koipse; Ro, Rohusi; LM, Lõuna-Malusi
Phorophytes: Pi, Pinus sylvestris; Ju, Juniperus communis; Al, Alnus glutinosa; Be, Betula spp.; Po, Populus tremula; Sal, Salix caprea;
Sam, Sambucus racemosa; So, Sorbus aucuparia; Sy, Syringa vulgaris; * records on Calluna and Empetrum are not included.
Folia Cryptog. Estonica
Table 2 (continued)
61
(Dietrich & Scheidegger, 1997). The first part of
our lichen studies on the islets of Kolga bay deals
with distribution and frequency of most common
epiphytic lichens, based on floristic data. The
similarity of epiphytic lichen flora of different
islets is briefly discussed, as compared with the
data on vascular plants and bryophytes.
MATERIAL AND METHODS
The material (820 herbarium specimens) was
collected on the islands of Kolga bay mostly in
1991 and 1993 and is kept in the Herbarium
of Tallinn Botanic Garden (TBA). Each piece of
bark was examined carefully and not only the
species for what it was collected, but all accompanying species were identified or recorded as
well. In all 2890 identifications and checking of
epiphytic lichens (140 species) were made by
the second author. The nomenclature of lichen
species follows mainly Santesson (1993).
The lichens were collected on 9 woody plant
species: Pinus sylvestris (57 trees), Juniperus
communis (10), Sorbus aucuparia (21), Populus
tremula (5), Alnus glutinosa (5), Betula pubescens (2), Salix caprea (5), Syringa vulgaris (1),
Sambucus racemosa (1), altogether on 109 trees
or shrubs.
Some numeric characteristics of the islets
of Kolga bay, the number of epiphytic lichen
species recorded on each islet included, are
represented in Table 1.
RESULTS AND DISCUSSION
The highest number of corticolous lichen species
has been recorded from the islets of Pedassaar
(79) and Rohusi (74) that are covered with forest
in great extent.
Table 2 includes more frequent lichen species (5 or more records), showing their distribution on the islets and on the most common
phorophytes. Sorbus aucuparia, Pinus sylvestris
and Alnus glutinosa are the phorophytes that
carry the highest number of lichen species.
Lecanora hagenii, L. symmicta, Physcia tenella,
Scoliciosporum chlorococcum and Xanthoria polycarpa are ubiquitous lichen species inhabiting
the islets of Kolga bay. They are met on all kinds
of phorophytes on all islets. The named species
belong to the rich bark species (Du Rietz, 1945)
and their occurrence on normally nutrient-poor
and acidic bark (incl. that of conifers) indicates
bark eutrophication under the influence of seawater aerosols and ammonia derived from bird
excrements. Hypogymnia physodes and Parmelia sulcata are also very frequent and are found
on their usual bark substrates, as well as on
boulders and on sand of coastal grey dunes.
Table 3. The similarity of vascular plant flora
(Sørensen’s similarity coefficient, %, left bottom)
and number of species in common (top right) on
four islets of Kolga Bay. Abbreviations of the islet
names see Table 2.
Islet
Ra
Pe
Ko
Ro
Ra
Pe
Ko
Ro
=
56
62
58
109
=
58
61
130
101
=
65
118
103
119
=
Table 4. The similarity of bryophyte flora (Sørensen’s similarity coefficient, %, left bottom)
and number of species in common (top right)
on four islets of Kolga Bay. Abbreviations of the
islet names see Table 2.
Islet
Ra
Pe
Ko
Ro
Ra
Pe
Ko
Ro
=
46
61
50
24
=
64
65
33
33
=
56
24
30
27
=
Table 5. The similarity of epihytic lichen flora
(Sørensen’s similarity coefficient, %, left bottom)
and number of species in common (top right) on
five islets of Kolga Bay. Abbreviations of the islet
names see Table 2.
Islet
Ra
Pe
Ko
Ro
LM
Ra
Pe
Ko
Ro
LM
=
44
53
48
60
26
=
53
87
42
29
44
=
61
50
27
51
49
=
55
20
26
27
32
=
62
Folia Cryptog. Estonica
Owing to careful examination of every
piece of bark, many crustose species, such as
Amandinea punctata, Catillaria nigroclavata,
Cliostomum griffithii, Rinodina pyrina, R. sophodes, that could be easily overlooked in field
are recorded with quite high frequency. Some
easily detectable epiphyte species, characteristic
to pine in the mainland part of Estonia, such
as Chaenotheca ferruginea, Hypocenomyce scalaris (Ach.) M. Choisy, Parmeliopsis ambigua,
Platismatia glauca (L.) W.L. Culb. & C.F. Culb.,
Pseudevernia furfuracea and Vulpicida pinastri
were rarely met on pines on the islets of Kolga
bay.
Similarities of vascular plant and bryophyte
floras are compared between four islands (Tables
3 and 4), Lõuna Malusi is left out. The island is
inhabited with large seabird colonies that have
influenced the vegetation heavily by trampling
and manuring, so the species diversity of ground
vegetation is much lower on Lõuna Malusi than
on other islets (see Table 1). As epiphytic lichens
are more independent on bird activities, Lõuna
Malusi is included into the analysis of similarity
of lichen floras (Table 5).
The data in the Table 3 allows to conclude
that there are no big differences between the
Sørensen’s coefficients of similarity between the
islets in regard of the composition of vascular
plant floras. The most dissimilar are floras of
Rammu and Pedassaar. It concerns vascular
plants (similarity 56 %), as well as bryophytes
(46%) and epiphytic lichens (44%). These two
largest islets have nearly equal areas, but differ largely in landscape diversity and dominant
vegetation types (Ratas et al., 1995). The most
similar are the bryophyte (65%) and especially
the epiphytic lichen (87%) floras of Pedassaar
and Rohusi islets, although the area of Rohusi is
less than 1/8 of Pedassaar. Pine forest is dominant and fairly extensive on both islets and the
same species of deciduous trees are spread on
forest edges along the shoreline.
ACKNOWLEDGEMENTS
Thanks are due to Mari Sarv for identification
and verification of Rinodina species, to Leiti
Kannukene for making available unpublished
data on bryophytes and to Laimdota Truus for
preparing the map. Financial support from the
Environmental Department of Harjumaa County
Government is gratefully acknowledged.
REFERENCES
Dietrich, M. & Scheidegger, C. 1997. Frequency,
diversity and ecological strategies of epiphytic lichens in the Swiss Central Plateau and Pre-Alps.
Lichenologist 29: 237–258.
Du Rietz, E. 1945. Om fattigbark- och rik-barksamhällen. Svensk Bot. Tidskr. 39: 147–148.
Karukäpp, R. & Malkov, B. 1993. Pinnamood. (Summary: Bottom topography). In Geology of the
Estonian shelf (eds Lutt, J. & Raukas, A.), pp.
22–29. Tallinn.
Lepland, A. 1995. Kolga lahe saarte randade geoloogia
(Summary: Coastal geology of the islands of the
Kolga Bay). Eesti Looduseuurijate Seltsi Aastaraamat 76: 109–136.
Lippmaa, T. 1935. Eesti geobotaanika pôhijooni. Acta
Comm. Univ. Tartuensis A 28: 1–151.
Loopmann, A. 1996. Eesti meresaarte nimestik.
Tallinn. 84 pp.
Nilson, E., Kannukene, L., Truus, L., Ratas, U., Puurmann, E. & Tobias, M. 1997. Biological diversity.
In Small islands of Estonia. Landscape ecological
studies (eds Ratas, U. & Nilson, E.) Inst. of Ecol.
Publ. 5: 131–179. Tallinn.
Nilson, E. & Piin, T. 1993. Kuni 300 samblikuliiki.
Eesti Loodus 4: 140.
Ploompuu, T. 1995. Harjumaa klindieelsete saarte
soontaimed (Summary: Vascular plants of the
preglint islands of the Harjumaa county). Eesti
Looduseuurijate Seltsi Aastaraamat 76: 15–49.
Randlane, T., Jüriado, I., Martin, J., Martin, L., Nilson, E. & Temina, M. 1997. Lichens of Naissaar
Island (Gulf of Finland, Estonia). Folia Cryptog.
Estonica 31: 20–25.
Ratas, U., Nilson, E., Truus, L.& Kannukene, L. 1995.
Development of landscapes on the islands of the
Kolga Bay. Eesti Looduseuurijate Seltsi Aastaraamat 76: 137–163.
Rebassoo, H. 1987. Biocoenoses of Eastern Baltic
islets, their composition, classification and protection. II (in Russian). Tallinn. 142 pp.
Santesson, R. 1993. The lichens and lichenicolous fungi
of Sweden and Norway. Lund. 240 pp.
Varep, E. 1964. The landscape regions of Estonia.
TRÜ Toimetised 156, Geograafia-alaseid töid IV:
3–28.
Folia Cryptog. Estonica, Fasc. 32: 63–66 (1998)
Athelia arachnoidea, a lichenicolous basidiomycete in Estonia
Erast Parmasto
Institute of Zoology & Botany, Estonian Agricultural University, 181 Riia St., EE 2400 Tartu, Estonia
Abstract: Athelia arachnoidea (Berk.) Jülich is a frequent lichenicolous basidiomycete that often grows in Estonia in towns
and along roadsides, and destroys epiphytic algae and lichens. In “wild” forests it has not been detected in Estonia. Mycelium
growth is extremely vigorous in winter, at about 1–3 ºC. The fungus distributes probably with small white sclerotia, sporebearing basidia have been noticed only rarely. Anamorphic state of this species, Fibulorhizoctonia carotae (Rader) G. Adams &
B. Kropp is a cold-storage pathogen of carrot; it has not been found in Estonia.
Kokkuvõte: E. Parmasto. Parasiitnahkis (Athelia arachnoidea), samblikel kasvav kandseen Eestis.
Athelia arachnoidea (Berk.) Jülich on Eesti linnades, asulates ja teeäärtes üha sagedasem parasiitseen, mis rõngana laiemaks
kasvades hävitab puutüvedel vetikaid ja mitmeid samblikuliike. Metsades pole sellist kahjustust Eestis seni täheldatud.
Seeneniidistiku kasv on eriti intensiivne talvel sulailmaga, kui õhu temperatuur on vaid 1–3 ºC. Seen levib meil tõenäoliselt
peamiselt väikeste valgete sklerootsiumide abil, eoseid produtseerivaid viljakehasid on täheldatud väga harva. A. arachnoides’e
anamorf Fibularhizoctonia carotae (Rader) G. Adams & B. Kropp tekitab porgandite külmsäilitamisel nende haigestumist;
Eestist teda seni leitud pole.
INTRODUCTION
In Estonian towns and along roadsides, thalli of
Xanthoria parietina, Physcia tenella and several
other lichens are sometimes “moldy”, damaged
by a fungus. I have seen such lichens since
1943 when Hans Trass and me studied lichens
together in our hometown Nõmme and avoided
collecting such “bad” specimens. Only decades
later I found out that the “mold” is not less interesting than its host lichens.
TAXONOMY
Athelia arachnoidea (Berk.) Jülich, Willdenowia Beih. 7: 53 (1972). Basionym: Corticium
arachnoideum Berk., Ann. Mag. Nat. Hist. 13:
345 (1844); syn.: Hypochnus bisporus J. Schröt.,
Pilze Scles. 3 (1): 415 (1888), Athelia epiphylla
Pers., Mycol. Eur. 1: 84 (1822) sensu auct. plur.
Anamorph: Fibularhizoctonia carotae (Rader) G.
Adams & B. Kropp, Mycologia 88 (3): 464 (1996);
basionym: Rhizoctonia carotae Rader, Phytopathology 38: 444 (1948).
Good descriptions and figures: Jülich, 1972:
53, Fig. 8; Eriksson & Ryvarden, 1973: 103, Fig.
37; Poelt, 1975, Fig. 1; Arvidsson, 1976, Fig. 1;
Gilbert, 1988, Fig. 1.
Damaged by the fungus lichens and algae,
but also surrounding tree bark is covered with
thin and loose cobweb-like (arachnoid) white or
slightly creamish mycelium. Later small white
spherical granules (“sclerotia”) 0.2–0.7–(1) mm
in diam. develop on mycelium in groups. Basidioma (fruit-body) of the fungus is similar but
covered with thin pellicle formed by basidia.
Microscopically both mycelium and basidioma
consist of loosely interwoven thin-walled hyphae
3–7 µm in diam., which are branched under
right angle; septa numerous, without clamps or
with few clamps mainly on basal hyphae. The
hyphae attached to and near the algal cells are
up to 8 µm in diam., richly branched, somewhat
torulose and with slightly monilioid short sidebranches. At the margin of a mycelium there
are some straight very long hyphae. Basidia are
clavate, 20–30 × 5–7 µm, with 2–4 (mainly 2)
sterigmata. Spores are narrowly ellipsoid, thinwalled, 7–9–(10) × 3.5–5 µm.
Basidiomata have been described in several
monographs and key books without mentioning
characteristic small white sclerotia. Reason for
such an omitting is simple: only full-developed
basidiomata have been described not paying attention to the surrounding these marginal area
of aerial mycelium.
Hyphae of this species have been described
and figured as smooth, i. e. without crystals. In
all specimens studied by me (about 30), several
or many hyphae of mycelium, but sometimes
64
Folia Cryptog. Estonica
also of basidiomata, are covered with short rodelike crystals of calcium oxalate-dihydrate as
shown in Adams & Kropp (1996: 464, Fig. 5).
RESULTS
Distribution and hosts in Estonia
A. arachnoidea has been found in almost all
suitable places where it was searched for (Parmasto, 1995; Fig. 1). Typical localities are living
trunks of deciduous trees (mainly Tilia cordata)
and dead twigs of Syringa vulgaris on roadsides,
at alleys and in parks. It is growing on many
lichen species and protococcoid algae; Xanthoria
parietina (L.) Th. Fr. and Physcia tenella (Scop.)
DC. are the most common hosts in Estonia; it
has been found also on Anaptychia ciliaris (L.)
Körb., Lecanora conizaeoides Nyl., Melanelia exasperatula (Nyl.) Essl., Parmelia sulcata Taylor,
Physconia distorta (With.) J.R. Laundon.
A. arachnoidea has been seen since the
40ies of this century in Estonia but was never
so widespread as during the last 5–8 years. For
example, in Tartu more and more trees covered
with lichens and algae are infected since the
beginning of the 90ies. This is possibly caused
by relatively warm winters with frequent and
long thaw periods.
Fig. 1. Localities of Athelia arachnoidea in Estonia.
Cycle of development
Late autumn and in winter, mycelium of A.
arachnoidea is frequently seen on tree trunks
as glittering white circles up to 40 cm long (Fig.
2); sometimes these circles unite and form an
infected area up to 5 m along a trunk. When
it is thawing after severe frosts, the circles appear again after some few days. In this stage,
mycelium is web-like, loose, in many cases
with white loose sclerotia turning brownish,
and more compact later. Hyphae are always
covered with numerous crystals; no basidia or
free spores were seen.
We have no collections of this species before
May, but possibly they survive. From July, mycelium can be seen on lichens again. Pellicular
hymenium with developed and spore-bearing
basidia has been seen in 3 specimens (of 30),
namely in August and September. First, usually
small and white sclerotia can be seen rarely in
August, frequently in September and later.
DISCUSSION
Taxonomy of the Athelia epiphylla-complex is
unclear; several species have been described
based on small differences in spore form and
size without any statistical study on variability
of these characters. Under the name A. arachnoidea fungi growing parasitically on lichens or
algae, and on wood (bark) have been united.
There is also another species found sometimes on lichens, A. epiphylla. Main difference
between these species is that basidia of A. arachnoidea have usually two sterigmata (and spores),
A. epiphylla – four. This is a vague character:
most of the specimens growing on lichens and
algae are sterile, i. e. without basidia and spores.
Among Estonian specimens there are some with
almost equal number of 2- and 4-spored basidia
(TAA 152154, 152433).
The specimens collected in Estonia are similar to each other, and support J. Eriksson’s
(in Arvidsson, 1976) and Arvidsson’s idea that
only one species in this species group is growing on lichens and algae, characterized by the
production of sclerotia. Correct name for this
taxon is A. arachnoidea: its type was “creeping over mosses and lichens on fallen sticks”.
However, this species has been found also on
65
Fig. 2. White-margined lesions on algal cover on a lime tree, formed by Athelia arachnoidea. Tartu,
18 Jan 1998. Photo by E. Parmasto.
dead leaves of deciduous trees, and I have seen
it spreading from lichens to surrounding tree
bark and wood.
Diederich (1986) mentioned that he had not
seen specimens with fructifications (= specimens
with developed hymenium) in Luxembourg.
During two-years’ observations in Sheffield
area (Great Britain), Gilbert (1988) did not observe any spores produced by this fungus. In
Estonia, of the 30 specimens studied only one
has quite well developed hymenium. Possibly
the main way of distribution in this species is
by sclerotium-like propagula which are loosely
attached to mycelium, as said already by Poelt
(1975: 7).
There are two ecological peculiarities in
A. arachnoidea. In Estonia, it has been found
only in anthropogenic localities, never in “wild”
forests. According to Arvidsson (1976: 7), “the
parasite appears to be less common outside
towns or in areas with no air pollution”. Poelt
& Jülich (1969) described it as found in Berlin,
but also in forests near the city. Only in England,
the fungus is widespread in open woodland as
well as in valley bottoms (Gilbert, 1998: 183).
Another interesting feature is vigorous
growth of its mycelium at low air temperature
(in Estonia, at about 1–4 ºC in winter). Air pollution and low temperature seem to weaken
lichens and algae infected thereafter with A.
arachnoidea. In England, radial extension of
the mycelium occurred only during the winter
months from November to February (Gilbert,
1988: 185).
As demonstrated by Adams & Kropp (1996),
anamorphic state of this species, Fibulorhizoctonia (Rhizoctonia) carotae is a cold-storage
pathogen of carrot in Europe, North America
and India, developing mainly at temperature
1–3 ºC and 92–96 % relative humidity. These
66
Folia Cryptog. Estonica
environmental conditions are the same as for development of the fungal mycelium and sclerotia
on lichens in Estonia in wintertime. Occurrence
of A. arachnoidea as an agent causing crater rot
of carrot roots in their storage has not yet been
studied in Estonia.
ACKNOWLEDGEMENTS
The author of this paper is greatly indebted
to Dr Tiina Randlane for identification of several
lichen species – hosts of A. arachnoidea, and
to Dr Alan Morton for providing his mapping
program DMAP for WINDOWS.
REFERENCES
Adams, G. C. & Kropp, B. R. 1996. Athelia arachnoidea, the sexual state of Rhizoctonia carotae, a
pathogen of carrot in cold storage. Mycologia 88
(3): 459–472.
Arvidsson, L. 1976. Athelia arachnoidea (Berk.) Jül.
and its influence on epiphytic cryptogams in
urban areas. Göteborgs Svampklubb Årsskrift
1975–76: 4–10.
Diederich, P. 1986. Lichenicolous fungi from the
Grand Duchy of Luxembourg and surrounding
areas. Lejeunia II 119: 1–26.
Eriksson, J. & Ryvarden, L. 1973. The Corticiaceae of
North Europe. Vol. 2. Fungiflora, Oslo. Pp. 57–261,
pl. 1–24.
Gilbert, O. L. 1988. Studies on the destruction of
Lecanora conizaeoides by the lichenicolous fungus Athelia arachnoidea. Lichenologist 20 (2):
183–190.
Jülich, W. 1972. Monographie der Athelieae (Corticiaceae, Basidiomycetes). Willdenowia Beih. 7:
3–283.
Parmasto, E. 1995. Nõiaringid puutüvedel. Eesti
Loodus 1: 23.
Poelt, J. 1975. Basidioflechten eine in den Alpen
lange übersehene Pflanzengruppe. In Jubiläumsjahrbuch 1900–1975, 40. Band des Vereins
zum Schutze der Alpenpflanzen und -Tiere e. V.
München, pp. 1–16.
Poelt, J. & Jülich, W. 1969. Über die Beziehungen
zweier corticioider Basidiomyceten zu Algen. Österr. Bot. Z. 116: 400–410.
Folia Cryptog. Estonica, Fasc. 32: 67 (1998)
The first record of Ochrolechia szatalaënsis in Estonia
Taimi Piin1 & Lea Lensment2
Tallinn Botanic Garden, 52 Kloostrimetsa Rd., EE0019 Tallinn, Estonia
2
Tallinn University of Educational Sciences, 25 Narva Rd., EE0001 Tallinn, Estonia
1
Abstract: Ochrolechia szatalaënsis Verseghy is reported for the first time from Estonia, lichen was found on bark of Acer
platanoides.
Kokkuvõte: T. Piin ja L. Lensment. Sambliku Ochrolechia szatalaënsis esmasleid Eestis.
Esmakordselt teatatakse Ochrolechia szatalaënsis Verseghy leiust Eestis, samblik kasvas vahtra (Acer platanoides) koorel.
Ochrolechia szatalaënsis Verseghy was described from Bulgaria (Verseghy, 1958). An
improved, detailed description was given by I.
M. Brodo (1991).
This corticolous lichen has an oceanic and
suboceanic distribution in Holarctic Kingdom:
Boreal and Tethyan Subkingdoms. O. szatalaënsis prefers bark of conifers (Picea, Abies) in Central and Southern Europe and bark of deciduous
trees (Sorbus, Salix, Betula, Quercus, Corylus)
in Western and Northern Europe (Purvis et al.,
1994). The species occurs in western North
America on bark and wood of different kinds
including Betula, Alnus, Quercus, Pseudotsuga,
and Thuja (Brodo, 1991). The nearest locations
of this species to Estonia are in Gotland, Sweden
(Santesson, 1993).
The second author collected O. szatalaënsis
in the northern part of our country, Raplamaa
Co., in the vicinity of Märjamaa settlement
(60 km south of the Gulf of Finland; 58°54’N
24°26’E) during a bioindication study in 1994.
The specimen examined was found on the stem
bark of Acer platanoides L. (exposition S, at the
height of 1.5 m, not abundant) in Märjamaa Orthodox Cemetery (Herbarium of Tallinn Botanic
Garden, TBA). The associated species included
Lecidella elaeochroma (Ach.) M. Choisy and Pertusaria leucostoma A. Massal.
O. szatalaënsis belongs to the upsaliensisgroup of Ochrolechia following I. M. Brodo (1991).
Its good diagnostic features are the almost white,
thin thallus, white-pruinose apothecia and discs
(all UV+ and C+ yellow). This species is closely
related to O. upsaliensis (L.) Massal., a common
arctic-alpine terricolous lichen species.
We suppose that this species belongs by
its distribution pattern to holarctic suboceanic
element in the lichen flora of Estonia (Trass,
1970).
The detailed description of the specimen will
given in future together with other Ochrolechia
species in Estonia which are just under revision.
The following species of Ochrolechia A. Massal.
have been published for Estonia in former times:
O. alboflavescens (Wulfen) Zahlbr., O. androgyna
(Hoffm.) Arnold, O. arborea (Kreyer) Almb., O.
frigida (Sw.) Lynge, O. microstictoides Räsänen,
O. pallescens (L.) A. Massal. and O. turneri (Sm.)
Hasselrot (Trass, 1970; Ekman et al., 1991).
REFERENCES
Brodo, I. M. 1991. Studies in the lichen genus Ochrolechia. 2. Corticolous species of North America.
Canad. J. Bot. 69: 733–772.
Ekman, S., Fröberg, L., Kärnefelt, I., Sundin, R. &
Thor, G. 1991. New or interesting lichens from
Estonia. Folia Cryptog. Estonica 28: 1-32.
Purvis, O.W., Jørgensen, P. M., Coppins, B. J. 1994.
Ochrolechia szatalaënsis Vers., new to Great Britain and Ireland. Lichenologist 26: 393–397.
Santesson, R. 1993. The lichens and lichenicolous
fungi of Sweden and Norway. SBT-förlaget, Lund.
240 pp.
Trass, H. 1970. The elements and development of the
lichen-flora of Estonia (in Russian). Pap. Bot. 9:
5–233.
Verseghy, K. 1958. Studien über die Gattung Ochrolechia II. Neue Flechten. Ann. Hist.-Nat. Mus.
Natl. Hung. 50: 76–85.
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Folia Cryptog. Estonica, Fasc. 32: 69–73 (1998)
Deterioration of the lichen flora in the National Nature Reserve
Rozsutec (the Malá Fatra Mts, Slovakia)
Ivan Pišút & Anna Guttová
Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-84223, Bratislava, Slovakia
Abstract: So far 273 lichen taxa have been listed from the territory. The analysis of the results gathered in the course of
the 1970s within the area of Vel’ký Rozsutec Mt., the Malá Fatra Mts, demonstrated intensifying antropogenous pressure.
The contemporary research (1996–1997) confirmed 124 species, recorded 55 taxa new to the area and gave evidence of
threat of 59 lichens (21.7%), enrolled in the local red list presented here. The process of impoverishment affects mainly
epiphytes dependent on air humidity and the alpine muscicolous and terricolous taxa concentrated about the summit. Over
the period of elapsed 25 years several toxitolerant, acidofilous and nitrofilous species emerged.
Kokkuvõte: I. Pišút ja A. Guttová. Lihhenofloora vaesustumine Rozsuteci looduskaitsealal (Malá Fatra,
Slovakkia).
Kokku on looduskaitsealalt teada 273 samblikuliiki. 70-ndatel a-tel läbiviidud uurimus viitas tugevale antropogeensele survele.
Käesoleva töö (1996-1997) käigus täheldati 124 liigi esinemist ja leiti 55 piirkonnale uut liiki; 59 samblikuliiki (21,7%)
tuleb pidada ohustatuks ning need lülitati kohalikku punasesse nimekirja. 25 viimase aasta jooksul on lisandunud paljud
vähetundlikud, happe- ja lämmastikulembesed liigid .
INTRODUCTION
The National Nature Reserve (NNR) Rozsutec
(734.74 ha), situated in the northern part of
the Malá Fatra Mts, covers up the geobiological complex of the landscape dominant Vel’ký
Rozsutec Mt. Its crest is built of dolomite and
limestone both forming vivid mosaic of rock faces. The lower parts are of calcareous sandstone
and schist. The vast area stretches within the
altitude limits from 600 to 1610 m. The beech
and fir forest formations, relict mountain maple
woods and mountain pine belt with scattered
spruce keep their natural character to a significant extent (Klinda, 1985). The average annual
precipitation falls vary from 900 to 1000 mm;
the average annual temperature amounts are
from 2 to 5°C (Intribus, 1981).
The NNR Rozsutec is exhaustively examined.
The first notes date back to the second half of
the 19th century (Brancsik, 1862). More intensive investigation started in the 1920s and
1930s (Servít, 1954; Servít and Cernohorský,
1935; Suza, 1926, 1934, 1943, 1944) and
then in the 1960s and 1970s (Pišút, 1966,
1968, 1969, 1971, 1974; Vêzda, 1958, 1965a,
1965b, 1970, 1972, 1973, 1978). The collective
information on the local lichenflora was presented by Pišút (1981). By 1973, 218 species
were recorded (Pišút, 1981, 1992, 1996). In the
meantime, the antropogenous influence upon
this domain, exposed in terms of orography and
tourism, increased.
MATERIAL AND METHODS
The study is based on two field visits in 1996
and 1997. The observations were guided by the
steps given in Pišút (1981). The research conducted after 25 years is comparable. Floristic
knowledge is updated as well. The species with
localities are listed in Pišút & Guttová (1998, in
press). Moreover, the adjacent area (the Vrátna
dolina valley, Štefanová and Zázrivá villages) is
included. Regarding the changes in the diversity
of the lichenflora, which no way could be neglected, the local red list (Table 1) is elaborated.
Material from BRA, SAV and SLO was studied
and revised.
The nomenclature as well as the categories
of threat follow Pišút et al. (1996). The figures in
the Tables 2, 3, 4 giving the number of species
in the NNR and the surroundings, cannot be
added up to obtain the total number, as some
of the taxa occur in both areas.
The specimens collected are at disposal at
the Institute of Botany, Slovak Academy of Sciences, Bratislava (SAV).
70
Folia Cryptog. Estonica
RESULTS AND DISCUSSION
The isolated massif of Vel’ký Rozsutec Mt. (alt.
1606 m) functions as a natural barrier to emission particles transported mostly by northwestern winds. While in the first quarter of the
20th century this factor played minor role (the
only eventual source of pollutants was Zilina, a
town 26 km westward, with a population of less
than 10,000 then), the 1950s were the turning
point. Since 1952 Vel’ký Rozsutec Mt. has been
attacked from smelters in Istebné (10 km SE),
since 1965 from Široká (18 km E) and a couple
of decades by emission generated in chemical,
textile and power supply industries as well as
engineering of 100,000-inhabitant Zilina. Vel’ký
Rozsutec Mt., the distinct elevation, is the
„ideal destination“ for long-distance emissions
originating in the industrial area of Ostrava (80
km NW, Czech Republic) and Silesia (120 km,
Poland).
The field work in the 1970s documented
the impoverished diversity of mostly epiphytic
lichens, which was the great contrast looking
back to 1920s and 1930s. Even in 1973 several more susceptible epiphytes died out (e.g.
Bryoria bicolor, Collema fragrans, C. nigrescens, Evernia divaricata, Hypogymnia bitteri,
Nephroma resupinatum, Normandina pulchella,
Peltigera collina, Ramalina calicaris, Ramalina
farinacea). The others, e.g. Graphis scripta, Lobaria pulmonaria, and the representatives of the
genera Bryoria and Usnea were extremely rare.
The remarkable thing was the extinction of the
lignicolous species Cladonia botrytes, once not
that uncommon.
The contemporary research (1996–1997)
gave evidence of the extinction or alternatively
missing of Belonia herculina, Cetrelia cetrarioides, Hypogymnia bitteriana, Lecanora albella,
L. intumescens, Lobaria pulmonaria, Ochrolechia
alboflavescens, Pertusaria flavida, P. leioplaca,
the Usnea species. The domination of the
antropogenous influences over the epiphytic
lichenflora, with consequences on its diversity,
is supported by the following examples.
1. Suza (1934) examined Populus pyramidalis,
P. tremula and Fraxinus excelsior between the
villages Terchová and Zázrivá (north periphery
of the Rozsutec Mt.), writing down subsequent
taxa: Candelaria concolor, Collema fragrans,
Leptogium saturninum, Parmelia caperata, P.
exasperata, P. glabra, P. subargentifera, P. subaurifera, Phaeophyscia nigricans, P. orbicularis,
Physcia stellaris, Physconia distorta, P. enteroxantha, P. grisea, Xanthoria candelaria, X. fallax,
X. parietina.
In 1996 we recorded here Physconia distorta, Xanthoria parietina (both with high cover
degree), Candelariella reflexa, Lecania cyrtella,
Parmelia submontana, Phaeophyscia orbicularis,
Physcia tenella.
2. As for the north slope of Rozsutec (900–1000
m), Suza (1934) made reference to the occurrence of Lobaria pulmonaria, Nephroma resupinatum (rather abundant), Leptogium saturninum,
Bryoria bicolor, Collema nigrescens, Graphis
scripta, Hypogymnia physodes, Normandina
pulchella, Parmelia glabratula, P. sulcata, Peltigera collina, P. praetextata, Platismatia glauca,
Pyrenula nitida, Ramalina farinacea, all growing
on Fagus sylvatica.
In 1996 Lecanora argentata, L. carpinea,
Graphis scripta (one thallus) and Pyrenula nitida
(one thallus) were put on record on the south
slope of the mountain (Fagus sylvatica). The
east slope was slightly more varied with Cladonia fimbriata, Hypogymnia physodes, Lecanora
conizaeoides, L. pulicaris, Parmelia glabratula, P.
saxatilis, P. sulcata, Parmeliopsis ambigua, Phlyctis argena, Platismatia glauca, Pseudevernia
furfuracea and Pyrenula nitida (rarely).
In contrast to the elevations and the outskirts
of the mountain region, one can find relatively
rich epiphytic lichen flora at the protected lower
altitudes. The Vrátna dolina valley, shielded by
surrounding mountain ridges, illustrates the
fact appropriately. Being a ski-centre, thus influenced by more or less heavy traffic, notwithstanding, one can still find Acrocordia gemmata,
Anaptychia ciliaris, Bacidia rubella, Caloplaca
cerina var. cerina, Caloplaca herbidella, Evernia
prunastri, Lecanora allophana, Leptogium saturninum, Opegrapha varia, Parmelia caperata,
P. flaventior, P. glabra, P. glabratula, P. pastillifera, Pertusaria albescens, Physconia distorta,
P. perisidiosa and Xanthoria parietina growing
on Fraxinus excelsior or Tilia parviflora (Guttová, 1997).
71
The diversity of muscicolous and terricolous
lichens decreases, apparently, mainly on the top
of Vel’ký Rozsutec Mt. and along the paths. In
this case, antropogenous influences are found
the crucial cause, although one cannot omit the
role of emission from near and distant sources.
Enormous rise in the number of tourists (the
Malá Fatra Mts is on the 2nd position, right
after the Vysoké Tatry Mts) has been bringing
about not only direct trampling of the soil cover,
but also a supply of nitrates. Even in the 1970s
Pišút (1981) could not confirm the occurrence
of several species such as Cetraria cucullata,
C. nivalis, Omphalina hudsoniana, Pannaria
pezizoides, Peltigera aphthosa, P. venosa and
Thamnolia vermicularis. The investigation in
1996–1997 confirmed the increase in the antropogenous pressure and further decrease
in lichen diversity. The species Arthroraphis
citrinella, Biatorella hemisphaerica, Caloplaca
sinapisperma, Cladonia macrophylla, Icmadophila ericetorum, Lecanora epibryon, Peltigera
leucophlebia, Pertusaria glomerata, Vulpicida
tubulosus were not found. In total, 59 lichen
species (21.7%) are enrolled in the local red list
(Tab. 1). Total account of the recorded species
is given in Tables 2, 3, 4.
Gradual acidification of the substrata in
the 1970s is documented by the presence of
Scoliciosporum chlorococcum, found at three localities, exclusively on Picea abies at 1200–1300
m. This acidophilous lichen was very rare in
the late 1960s in Slovakia, up to 1970 known
only from 11 grid squares (Pišút, 1985), as well
as Lecanora conizaeoides and Hypocenomyce
scalaris, typical invasively spreading acidophilous, toxitolerant lichens. The former species is
now found on timber, Picea abies, Larix decidua,
Abies alba and Fagus sylvatica at 700–1320 m.
Similarly, the latter one grew on Picea excelsa,
Table 1. Local red list of NNR Rozsutec and the adjacent area
Extinct/missing – Ex
Endangered – E
Vulnerable – V
Rare – R
Belonia herculina
Ochrolechia alboflavescens
Bryoria bicolor
Cetraria cucullata
Pannaria pezizoides
C. nivalis
Cetrelia cetrarioides
Cladonia botrytes
L. intumescens
Peltigera aphthosa
Collema fragrans
C. nigrescens
P. leucophlebia
Lobaria pulmonaria
Evernia divaricata
Hypogymnia bitteri
H. bitteriana
Lecanora albella
Peltigera venosa
Pertusaria flavida
P. glomerata
P. leioplaca
Ramalina calicaris
Thamnolia vermicularis
Acrocordia gemmata
Anaptychia ciliaris
Bacidia rubella
Bryoria capillaris
Caloplaca herbidella
Cetraria sepincola
Evernia prunastri
Graphis scripta
Hypogymnia tubulosa
Leptogium saturninum
Parmelia caperata
P. pastillifera
P. submontana
Pertusaria pertusa
Pyrenula nitida
Ramalina fastigiata
R. fraxinea
Usnea filipendula
Candelaria concolor
Cetraria islandica
Cliostomum corrugatum
Fulgensia bracteata
Chromatochlamys muscorum
Megaspora verrucosa
Physconia perisidiosa
Thelopsis melathelia
Xanthoria fulva
X. polycarpa
Agonimia tristicula
Calicium salicinum
Ochrolechia arborea
Parmelia subargentifera
P. tiliacea
Peltigera monticola
Toninia verrucarioides
Total:
24
18
10
7
72
Folia Cryptog. Estonica
Larix decidua and Abies alba at 900–1100 m.
More than that, other Hypocenomyce taxa are
spreading over Slovakia (H. caradocensis – 2
localities, fertile specimen at one of them. H.
leucococca – 2 localities. H. praestabilis – fertile
specimen, 1 locality).
Table 2. Number of lichens reported in NNR
Rozsutec and the adjacent area prior to 1973
Lichens
Epiphytic
Epilithic
Terricolous
Muscicolous
On plant debris
Epixylic
Total number
NNR
Adjacent
area
Total
number
58
77
35
7
7
3
187
46
4
7
0
1
1
59
86
78
36
7
7
4
218
Table 3. Number of lichens reported from NNR
Rozsutec and the adjacent area 1996–1997
Lichens
Epiphytic
Epilithic
Terricolous
Muscicolous
On plant debris
Epixylic
Total number
NNR
Adjacent
area
Total
number
38
54
12
15
5
10
134
50
13
2
2
0
2
69
71
63
13
16
5
11
179
Table 4. Number of newly recorded taxa in the
years of 1996–1997
Lichens
Epiphytic
Epilithic
Terricolous
Muscicolous
On plant debris
Epixylic
Total number
NNR
Adjacent
area
Total
number
12
14
3
3
2
9
44
12
3
0
1
0
2
20
22
14
3
4
2
10
55
Even the application of the Index of Ecological
continuity – IEK, (Pišút, 1997) confirms not only
decrease in indicatory epiphytic lichen species,
but also deterioration of the territorial quality
in environmental terms. While the occurence
of 15 indicatory species (IEK 15) in the 1920s
and 1930s pointed at a high quality of the forest
formations, the number of 8 in the 1970s (IEK
8) marked a poorer quality. The contemporary
research gives only 4 taxa (IEK 4), which represents the area of poor quality, and supports
the statements on increasing antropogenous
pressure.
ACKNOWLEDGEMENT
Financial support of the Slovak Grant Agency
(Projects Nos. 731 and 1181) is gratefully acknowledged. The authors are intebted to the
head office of the National Park Malá Fatra Mts,
who made the investigation possible.
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der Slowakei]. Bull. Slov. Bot. Spol. (Bratislava)
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Cechoslovakei. IV. Malá Fatra, Velká Fatra und
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(Malá Fatra na Slovensku). Sborn. Muz. Slov.
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Kenntnis des Vorkommens und der Verbreitung
einiger Flechten – Epiphyten in den Karpathen].
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1943: 1–59.
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a Pachyphiale s klícem a prehledem evropských
druhù. Sborn. Vysoké Školy Zemed. Lesn. Ser. C,
1: 21–56.
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Instituto botanico Academiae scientiarum Cechoslovacae, Prùhonice prope Pragam. Fasc. XVII.
(nos. 425–450). Brno. 7 pp.
Vêzda, A. 1965b. Lichenes selecti exsiccati editi ab
Instituto botanico Academiae scientiarum Cechoslovacae, Prùhonice prope Pragam. Fasc. XVIII.
(nos. 451–475). Brno. 7 pp.
Vêzda, A. 1970. Neue oder wenig bekannte Flechten in
der Tschechoslowakei. I. Folia Geobot. Phytotax.
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Vêzda, A. 1972. Lichenes selecti exsiccati editi ab Instituto botanico Academiae scientiarum Cechoslovacae, Prùhonice prope Pragam. Fasc. XLV. (nos.
1101–1125). Brno. 7 pp.
Vêzda, A. 1973. Lichenes selecti exsiccati editi ab Instituto botanico Academiae scientiarum Cechoslovacae, Prùhonice prope Pragam. Fasc. XLVI. (nos.
1126–1150). Brno. 7 pp.
Vêzda, A. 1978. Neue oder wenig bekannte Flechten in
der Tschechoslowakei. II. Folia Geobot. Phytotax.
13: 397–420.
74
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 75–79 (1998)
Red list of Estonian macrolichens
Tiina Randlane
Institute of Botany and Ecology, University of Tartu, 38 Lai St., EE2400 Tartu, Estonia
Abstract: A list comprising 110 endangered macrolichen species in Estonia is presented. The following threat categories
have been applied: 0 – extinct or probably extinct (18 species), 1 – endangered (5), 2 – vulnerable (24), 3 – rare (41), 4 – care
demanding (21), 5 – indeterminate (1).
Kokkuvõte: T. Randlane. Eesti suursamblike punane nimekiri.
Eesti ohustatud suursamblike nimekiri sisaldab 110 liiki. Kasutatud on järgmisi ohukategooriaid: 0 – hävinud või tõenäoliselt
hävinud (18 liiki), 1– eriti ohustatud (5), 2 – ohualtid (24), 3 – haruldased (41), 4 – tähelepanu vajavad (21), 5 – määratlemata (1).
INTRODUCTION
According to our present knowledge the lichen
flora of Estonia consists of about 800 species,
out of which we have verified data about the
distribution of 337 macrolichens. In addition,
the occurence of more than 450 microlichen species is strongly suggested in our territory. The
new checklist of Estonian lichen species will be
completed at the end of this year as a result of
a collective project. The revision of several microlichen genera has not been finished yet and,
therefore, the inclusion of the microlichens into
the red list is not considered to be scientifically
justified at present.
The following red list is based on data
presented in Trass & Randlane (1994) and on
herbarium materials collected or identified later.
Specimens cited are kept at the International
Center for Environmental Biology (ICEB), or
the Institute of Ecology (IE), both in Tallinn; in
the herbaria of the Universities of Helsinki (H),
Tartu (TU) or Uppsala (UPS). A few changes in
the checklist of Estonian macrolichens since
1994 should be pointed out.
1. Several species that were earlier considered to be extinct from the modern lichen
flora of Estonia (Trass & Randlane, 1994), are
now treated as doubtfully recorded. These species are: Bryoria bicolor (Ehrh.) Nyl., Collema
auriforme (With.) Coppins & J.R. Laundon, Collema crispum (Huds.) Weber, Collema fluviatile
(Huds.) Steudel, Dermatocarpon meiophyllizum
Vain., Leptogium byssinum (Hoffm.) Nyl., Leptogium plicatile (Ach.) Leight., Melanelia elegantula (Zahlbr.) Essl., Parmotrema chinense
(Osbeck) Hale & Ahti, Peltula euploca (Ach.)
Poelt, Phaeophyscia constipata (Norrl. & Nyl.)
Moberg, Physcia dimidiata (Arnold) Nyl., Physconia muscigena (Ach.) Poelt, Ramalina lacera
(With.) J.R. Laundon, Umbilicaria grisea Hoffm.,
Umbilicaria proboscidea (L.) Schrad., Umbilicaria
vellea (L.) Hoffm., Usnea articulata (L.) Hoffm.,
Usnea longissima Ach., Xanthoria elegans (Link)
Th. Fr. These taxa had been included in the flora
based on information in old literature (Bruttan,
1870, 1889; Mereschkowsky, 1913; Räsänen,
1931). Unfortunately, we have not seen any
herbarium specimens of them (possibly the
specimens are not preserved) and cannot verify
the identifications according to modern taxonomy. Still, we cannot convincingly exclude
them either, as their occurrence in Estonia
is chorologically possible. Therefore, a special
category of species – doubtful (marked with ?)
– is introduced now in all the lists and papers
concerning the revision of Estonian lichens (see
also Jüriado, 1998; Lõhmus, 1998; Suija, 1998).
In the present treatment doubtful species are
not included in the red list.
2. Five new species have been found during
the last few years: Bryoria furcellata (Fr.) Brodo &
D. Hawksw. (Randlane et al., 1997; ICEB, dup.
in TU), Dermatocarpon luridum (With.) J.R. Laundon (TU), Phaeophyscia chloantha (Ach.) Moberg
(Moberg, 1997; UPS, dup. in TU), Ramalina elegans (Bagl. & Carestia) Jatta (Skytén, 1993;
H), Solorina bispora Nyl. (Moberg, 1997; UPS).
One more species – Ramalina capitata (Ach.) Nyl.
– must be added to the list, according to earlier
literature data (Räsänen, 1931), as doubtful.
76
Folia Cryptog. Estonica
3. One species – Usnea glauca Mot. – is excluded from the list after re-determination (det.
Pekka Halonen 1997 as U. scabrata).
4. Some species reported as extinct in Trass
& Randlane (1994) have recently been found
again: Alectoria sarmentosa (Ach.) Ach. (Martin &
Martin, 1998; ICEB, dup. in TU), Cladonia brevis
(Sandst.) Sandst. (TU), Leptogium gelatinosum
(With.) J.R. Laundon (TU), Melanelia hepatizon
(Ach.) A. Thell (Randlane et al. 1997; ICEB, IE,
dup. in TU). Thus, these taxa are present in our
contemporary lichen flora.
5. Three more species – Nephroma helveticum Ach., Punctelia subrudecta (Nyl.) Krog and
Umbilicaria hyperborea (Ach.) Hoffm. – should
be considered as extinct from Estonia, according
to the latest data.
RED LIST OF ESTONIAN MACROLICHENS
The following threat categories have been applied:
0 – extinct or probably extinct,
1 – endangered,
2 – vulnerable,
3 – rare,
4 – care demanding,
5 – indeterminate.
Criteria for including the species and numeral data are presented in discussion. In nomenclature Trass & Randlane (1994) are mainly
followed; changes in taxonomy or nomenclature
compared to this are pointed out in brackets.
ALECTORIA SARMENTOSA (Ach.) Ach.
2
ANAPTYCHIA RUNCINATA (With.) J.R. Laundon
3
ARCTOPARMELIA INCURVA (Pers.) Hale
4
BRYORIA CHALYBEIFORMIS (L.) Brodo & D. Hawksw.
3
BRYORIA FURCELLATA (Fr.) Brodo & D. Hawksw. 4
BRYORIA IMPLEXA (Hoffm.) Brodo & D. Hawksw.
[incl. B. osteola (Gyeln.) Brodo & D.
Hawksw., B. pseudofuscescens (Gyeln.)
Brodo & D. Hawksw., B. vrangiana (Gyeln.)
Brodo & D. Hawksw.]
4
B RYORIA NADVOR NIKIANA (Gyeln.) Brodo & D.
Hawksw.
2
BRYORIA SIMPLICIOR (Vain.) Brodo & D. Hawksw.
3
CALOPLACA BIATORINA (A. Massal.) J. Steiner
3
CALOPLACA FLAVESCENS (Huds.) J.R. Laundon 3
CALOPLACA THALLINCOLA (Wedd.) Du Rietz
4
CALOPLACA VERRUCULIFERA (Vain.) Zahlbr.
3
CATAPYRENIUM CINEREUM (Pers.) Körb.
3
CATAPYRENIUM LACHNEUM (Ach.) R. Sant. [syn. Placidium lachneum (Ach.) de Lesd.]
3
CATAPYRENIUM PILOSELLUM (Breuss) Breuss [syn.
Placidium pilosellum Breuss]
3
CETRELIA OLIVETORUM (Nyl.) W.L. Culb. & C.F. Culb.
[incl. C. cetrarioides (Duby) W.L. Culb. & C.F.
Culb.]
0
CLADONIA BREVIS (Sandst.) Sandst.
2
CLADONIA CONVOLUTA (Lam.) Anders
3
CLADONIA DECORTICATA (Flörke) Spreng.
4
CLADONIA INCRASSATA Flörke
2
CLADONIA METACORALLIFERA Asahina
3
CLADONIA PARASITICA (Hoffm.) Hoffm.
2
COLLEMA BACHMANIANUM (Fink) Degel.
3
COLLEMA LIMOSUM (Ach.) Ach.
3
COLLEMA NIGRESCENS (Huds.) DC.
0
COLLEMA OCCULTATUM Bagl.
0
COLLEMA PARVUM Degel.
3
COLLEMA SUBNIGRESCENS Degel.
0
COLLEMA UNDULATUM Flot.
3
DERMATOCARPON ARNOLDIANUM Degel.
3
DERMATOCARPON LEPTOPHYLLUM (Ach.) K.G.W. Lång3
DERMATOCARPON LURIDUM (With.) J.R. Laundon 3
ENDOCARPON PSORODEUM (Nyl.) Blomb. & Forss. 2
ENDOCARPON PUSILLUM Hedw.
2
EVERNIA DIVARICATA (L.) Ach.
2
EVERNIA MESOMORPHA Nyl.
4
FLAVOCETRARIA CUCULLATA (Bellardi) Kärnefelt & A.
Thell
2
FLAVOPARMELIA CAPERATA (L.) Hale
0
FULGENSIA FULGENS (Sw.) Elenkin
3
HETERODERMIA SPECIOSA (Wulfen) Trevis
0
HYPERPHYSCIA ADGLUTINATA (Flörke) H. Mayrhofer &
Poelt
0
HYPOCENOMYCE ANTHRACOPHILA (Nyl.) P. James &
Gotth. Schneid.
3
H YPOCENOMYCE SOROPHORA (Vain.) P. James &
Poelt
3
HYPOGYMNIA VITTATA (Ach.) Parrique
2
LASALLIA PUSTULATA (L.) Mérat
4
LECANORA ACHARIANA A.L. Sm.
3
LEMPHOLEMMA ISIDIODES (Arnold) H. Magn.
3
LEPTOGIUM CYANESCENS (Rabenh.) Körb.
2
LEPTOGIUM GELATINOSUM (With.) J.R. Laundon 2
LEPTOGIUM RIVULARE (Ach.) Mont.
1
LEPTOGIUM SCHRADERI (Bernh.) Nyl.
2
LEPTOGIUM SUBTILE (Schrad.) Torss.
0
LEPTOGIUM TENUISSIMUM (Dicks.) Körb.
4
LOBARIA PULMONARIA (L.) Hoffm.
4
77
LOBARIA SCROBICULATA (Scop.) DC.
1
MELANELIA COMMIXTA (Nyl.) A. Thell
3
MELANELIA GLABRA (Schaer.) Essl.
2
MELANELIA HEPATIZON (Ach.) A. Thell
3
MENEGAZZIA TEREBRATA (Hoffm.) A. Massal.
4
MULTICLAVULA VERNALIS R.H. Petersen
2
NEPHROMA ARCTICUM (L.) Torss.
0
NEPHROMA BELLUM (Spreng.) Tuck.
1
NEPHROMA HELVETICUM Ach.
0
NEPHROMA ISIDIOSUM (Nyl.) Gyeln.
2
NEPHROMA LAEVIGATUM Ach.
2
NEPHROMA PARILE (Ach.) Ach.
4
NEPHROMA RESUPINATUM (L.) Ach.
2
PANNARIA LEUCOPHAEA (Vahl) P.M. Jørg. [syn. Fuscopannaria leucophaea (Vahl) P.M. Jørg.] 2
PANNARIA PEZIZOIDES (Weber.) Trevis.
2
PARMELIA FRAUDANS (Nyl.) Nyl.
3
PARMELIELLA TRIPTOPHYLLA (Ach.) Müll. Arg.
2
PARMELINA TILIACEA (Hoffm.) Hale
4
PELTIGERA DEGENII Gyeln.
3
PELTIGERA ELISABETHAE Gyeln.
0
PELTIGERA SCABROSA Th. Fr.
3
PELTIGERA VENOSA (L.) Hoffm.
3
PHAEOPHYSCIA CHLOANTHA (Ach.) Moberg
0
PHAEOPHYSCIA ENDOPHOENICEA (Harm.) Moberg 3
PHYSCIA MAGNUSSONII Frey
3
PHYSCIA SEMIPINNATA (J.F. Gmelin) Moberg
2
PILOPHORUS CEREOLUS (Ach.) Th. Fr.
3
PUNCTELIA SUBRUDECTA (Nyl.) Krog
0
PYCNOTHELIA PAPILLARIA Dufour
4
RAMALINA CALICARIS (L.) Fr.
2
RAMALINA ELEGANS (Bagl. & Carestia) Jatta
5
RAMALINA SILIQUOSA (Huds.) A.L. Sm.
3
RAMALINA SINENSIS Jatta
4
RAMALINA THRAUSTA (Ach.) Nyl.
4
SOLORINA BISPORA Nyl.
3
SOLORINA SACCATA (L.) Ach.
4
SOLORINA SPONGIOSA (Ach.) Anzi
4
SPHAEROPHORUS GLOBOSUS (Huds.) Vain.
1
STEREOCAULON EVOLUTUM Graewe
0
STEREOCAULON INCRUSTATUM Flörke
4
STEREOCAULON VESUVIANUM Pers.
3
TONINIA LOBULATA (Sommerf.) Lynge [syn. Mycobilimbia lobulata (Sommerf.) Hafellner] 0
TONINIA VERRUCARIOIDES (Nyl.) Timdal
3
UMBILICARIA CINERASCENS (Arnold) Frey
3
UMBILICARIA CYLINDRICA (L.) Duby
3
UMBILICARIA DECUSSATA (Vill.) Zahlbr.
3
UMBILICARIA HYPERBOREA (Ach.) Hoffm.
0
USNEA BARBATA (L.) F.H. Wigg.
0
USNEA DIPLOTYPUS Vain.
3
USNEA GLABRATA (Ach.) Vain.
1
USNEA SCABRATA Nyl. (incl. U. prostrata Vain., U.
rugulosa Vain., U. sylvatica Mot.)
4
VULPICIDA TUBULOSA (Schaer.) J.-E. Mattsson &
M.J. Lai
4
XANTHORIA CALCICOLA Oxner
4
XANTHORIA FALLAX (Hepp) Arnold
2
XANTHORIA LOBULATA (Flörke) Hellb. [syn. Caloplaca
lobulata (Flörke) de Lesd.]
0
XANTHORIA SOREDIATA (Vain.) Poelt
3
DISCUSSION
The present red list of macrolichens was compiled as a part of a larger project “Red Data Book
of Estonia. Threatened Fungi, Plants and Animals” (Lilleleht, in press) which hopefully will be
published in 1998. Threat categories applied in
this project were discussed on several meetings
of the Nature Conservation Commission of the
Estonian Academy of Sciences, consisting of representatives of different specialities. As a result,
the desision was made to apply the categories
that were already in use in the Red Data Books
of neighbouring countries (Andrušaitis, 1996;
Aronsson et al., 1995; Ingeloeg et al., 1993;
Rassi et al., 1992), with minor changes.
This is the first officially recognised red list
of Estonian lichens (available also in internet
http://www.ut.ee/lichens/red_list.html), although some earlier publications also present
relevant information: the list of 93 Estonian lichens in need of protection (Trass & Randlane,
1986) and the list of 38 extinct macrolichens in
Estonia (Trass & Randlane, 1987). A list of 40
macrolichen species that should be enclosed in
the Red Data Book of Estonia was printed in an
encyclopaedic reference book a few years ago
(Eesti A ja O, 1993).
The new list, based on latest verified data, includes 110 species, which comprises 33% of
the total macrolichen flora known in Estonia
at present. The distribution of taxa between
the threat categories and the criteria for their
including are as follows.
Category 0 – extinct or probably extinct
– includes 18 species (16 % of the total number
of lichens in the red list). Herbarium material
verifies earlier occurrence of all these species in
Estonia; in addition, there are no new records
78
Folia Cryptog. Estonica
since 1950. Several taxa of this category (e. g.
Hyperphyscia adglutinata, Nephroma helveticum, Umbilicaria hyperborea) have been collected
from the present territory of Tallinn (Nõmme, Rahumäe, Kopli, Lasnamäe) at the end of the last or
at the beginning of this century. These habitats
and localities have been certainly destroyed by
now. Some species have earlier also been found
in other towns (Phaeophyscia chloantha from
Kunda – a small but heavily polluted town in
norteastern Estonia; Cetrelia olivetorum and Flavoparmelia caperata from the closest vicinity of
Tartu). A few extinct species are recorded from
the islands of Saaremaa (Kuressaare, Orissaare,
peninsula of Sõrve), Abruka (Heterodermia speciosa) or Muhu (Toninia lobulata).
Category 1 – endangered – includes 5 species (5 %). All these species are very rare in
Estonia (found only once after 1950) and seriously threatened to a certain extent. Leptogium
rivulare is a rare species in the whole world,
confined to the periodically inundated trees or
occasionally rocks, on the banks of sluggish rivers and ponds in northern Europe and northeastern USA and Canada. It has disappeared
from several localities in Sweden, Finland and
Russia as well as in France, Canada and USA
(Jørgensen & James, 1983; Jørgensen, 1994;
Sierk, 1964). Only a few current localities are
known, one of them in Estonia: Pärnumaa
Co., Lihula, river Kasari, leg. H. Trass 1957,
det. T. Randlane 1986, verified G. Degelius
1991 (Randlane, 1987). This is a lichen that
strongly depends on changes of water level and
is threatened by the destruction of such habitats. Sphaerophorus globosus is an arctic-alpine
lichen growing in Estonia on its southern limit,
inhabiting here open localities – alvars – on the
Saaremaa Island. The general threat to alvars
– becoming overgrown with bushes and trees
as a result of ceased grazing – is dangerous to
several rare lichens of that habitat, including S.
globosus. Lobaria scrobiculata, Nephroma bellum
and Usnea glabrata grow in old virgin forests and
are mainly endangered by forestry.
Category 2 – vulnerable – includes 24 species (22 %). Lichens belonging to this category
are either quite rare or sparsely distributed in
Estonia but certainly sensitive to the environmental changes and, therefore, they may fall
into the previous category in the future. The
major threats are various: destruction of very
specialised habitats (Cladonia parasitica, Pannaria pezizoides), vulnerability due to growth on
the very limit of distribution area (Flavocetraria
cucullata), air pollution and cutting of forests
(Alectoria sarmentosa, Evernia divaricata, Ramalina calicaris).
Category 3 – rare – includes 41 species (37
%). All macrolichens that are recorded from one
or two localities only (and which are not included
in the first three categories) are treated as rare.
The main threats are not identified, the taxa
are considered endangered by the occasional
destruction of localities.
Category 4 – care demanding – includes
21 species (19 %). These taxa are distributed
sparsely or even widely in Estonia but they
are evidently sensitive to various factors and
their decline has been observed during last
decades.
Category 5 – indeterminate – comprises 1
species (1 %) at present (Ramalina elegans).
The distribution of this taxon in North Europe
was described only recently (Skytén, 1993); its
general distribution area and precise ecological
demands are not clear yet.
The most important sites of endangered lichens
in Estonia are: alvar communities, deciduous
forests and wooded meadows, sandy dunes, calcareous or granite rocks (often on the seashore).
Maintaining and undisturbing these habitats is
the main strategy in protection of endangered
lichens.
ACKNOWLEDGEMENTS
This research is a part of the project that was
ordered and financed by the Ministry of Environment and the Foundation of Environment.
The author is especially grateful to Hans Trass
for the information and prolonged consultations
during the last decades. All lichenologists in Tartu and Tallinn are thanked for discussions on
the subject. Jan-Eric Mattsson is acknowledged
for kindly revising the manuscript.
REFERENCES
Andrušaitis, G. (ed). 1996. Latvijas sarkana gramata.
I. Senes un kerpji [Red Data Book of Latvia. Vol. I.
Fungi and lichens]. Riga. 202 pp.
Aronsson, M., Hallingbäck, T. & Mattsson, J.-E. (eds.)
1995. Rödlistade växter i Sverige 1995 [Swedish
Red Data Book of Plants 1995]. ArtDatabanken,
Uppsala. 272 pp.
79
Bruttan, A. 1870. Lichenen Est-, Liv- und Kurlands.
Arch. Naturk. Liv-, Ehst- u. Kurl. Zweite Serie 7:
163–326.
Bruttan, A. 1889. Nachtrag zu den Lichenen Liv-,
Est- und Kurlands. Sitz.-ber. d. Nat.-forsch. Ges.
bei der Univ. Dorpat 8: 444–449.
Eesti A ja O. 1993. Eesti Entsüklopeedia Kirjastus,
Tallinn.
Ingeloeg, T., Andersson, R. & Tjernberg, M. (eds.)
1993. Red Data Book of the Baltic Region. I. Lists of
threatened vascular plants and vertebrates. Swedish Threatened Species Unit, Uppsala. 95 pp.
Jørgensen, P. M. 1994. Further notes on european taxa
of the lichen genus Leptogium, with emphasis on
the small species. Lichenologist 26: 1–29.
Jørgensen, P. M. & James, P. W. 1983. Studies on
some Leptogium species of Western Europe. Lichenologist 15: 109–125.
Jüriado, I. 1998. Review of the Lecanora subfusca
group in Estonia. Folia Cryptog. Estonica 32:
15–20.
Lilleleht, V. (ed.) 1998. Eesti Punane raamat. Ohustatud seened, taimed ja loomad [Red Data Book of
Estonia. Threatened Fungi, Plants and Animals].
Tallinn (in press).
Lõhmus, P. 1998. List of Estonian Caliciales. Folia
Cryptog. Estonica 32: 43–46.
Martin, L. & Martin, J. 1998. Epiphytic macrolichens
in Estonian forests. Folia Cryptog. Estonica 32:
47–55.
Mereschkowski, K. 1913. The checklist of lichens in the
Baltic provinces (in Russian). Kazan.
Moberg, R. 1997. Phaeophyscia chloantha and Solorina
bispora new to Estonia. Folia Cryptog. Estonica
31: 36–37.
Randlane, T. 1987. Leptogium rivulare (Ach.) Mont. – a
new rare lichen species in Estonia. Folia Cryptog.
Estonica 25: 8–11.
Randlane, T., Jüriado, I., Martin, J., Martin, L., Nilson, E. & Temina, M. 1997. Lichens of Naissaar
Island (Gulf of Finland, Estonia). Folia Cryptog.
Estonica 31: 20–25.
Rassi, P., Kaipiainen, H., Mannerkoski, I. & Ståhls, G.
1992. Uhanalaisten eläinten ja kasvien seurantatoimikunnan mietintö [Report on the monitoring
of threatened animals and plants in Finland].
Komiteanmietintö 1991, 30: 1–328.
Räsänen, V. 1931. Die Flechten Estlands. Helsinki.
163 pp.
Sierk, H. A. 1964. The genus Leptogium in North America North of Mexico. Bryologist 67: 245–317.
Skytén, R. 1993. Ramalina elegans, new to Sweden
and Norway. Graphis Scripta 5: 93–95.
Suija, A. 1998. Lecideoid lichens from Estonia – an
annotated checklist. Folia Cryptog. Estonica 32:
107–112.
Trass, H. & Randlane, T. 1986. Estonian lichens in
need of protection (in Russian). Folia Cryptog.
Estonica 21: 1–3.
Trass, H. & Randlane, T. 1987. Extinct macrolichens
of Estonia. Folia Cryptog. Estonica 25: 1–7.
Trass, H. & Randlane, T. (eds) 1994. Eesti suursamblikud. Tartu. 399 pp.
80
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 81–84 (1998)
Lichenological collections in TU
Andres Saag, Tiina Randlane & Ave Suija
Institute of Botany and Ecology, University of Tartu, 38 Lai St., EE2400 Tartu, Estonia
Abstract: Lichenological collections in TU include about 62 000 specimens. They are arranged into three main sections:
Estonian herbarium (ca. 24 000 specimens), geographical collections (ca. 30 000 specimens) and a comparison herbarium
(ca. 8 000 specimens). The specimen databasing has been started in the Estonian herbarium using the computer program
BRAHMS. The genera, species of which have been databased already, are listed. Exsiccate materials and types that are
located in TU are also listed.
Kokkuv›te: A. Saag, T. Randlane & A. Suija. Lihhenoloogilised kollektsioonid Tartu Ülikoolis.
Tartu Ülikooli (TU) lihhenoloogiline herbaarium sisaldab umbes 62 000 eksemplari. Samblike kollektsioon on jaotatud kolme
ossa: Eesti samblike herbaarium (u. 24 000 eksemplari), geograafilised kogud (u. 30 000 eksemplari) ja võrdlusherbaarium (u.
8 000 eksemplari). Alustatud on Eesti samblike herbaareksemplaride andmebaasi loomist, kasutades programmi BRAHMS.
Loetletud on andmebaasi juba sisestatud perekonnad. Samuti on esitatud meie herbaariumis paiknevate tüüpmaterjalide ja
eksikaatide loendid.
INTRODUCTION
A thorough review revealing the present state
of all herbaria of higher plants in Estonia
(including private herbaria and collections of
local museums and nature reserves) has been
recently presented by Kukk (1997). Lichenological collections are much less popular and
kept mainly in the research institutes. There
are five intitutions in Estonia where scientific
lichen herbaria are kept: International Center
for Environmental Biology (ICEB), Institute
of Ecology (IE), State Nature Museum (TAL),
Tallinn Botanic Garden (TBA) – all four located
in Tallinn – and University of Tartu (TU). Private lichenological collections are unusual in
Estonia. The greatest private lichenological herbarium, founded in 1930s, belonged to Heinrich
Aasamaa, a former lecturer of the University of
Tartu. The conditions of these materials were
unknown for decades. A few years ago it became
evident that the collections, left in a farm near
Tartu in the 1950s, had been destroyed. Heinrich Aasamaa, being over 80, started collecting
lichens and preparing a herbarium again. All
his present collections (including herbarium of
higher plants of about 30 000 sheets) are kept
at his home in Lasnamäe, Tallinn.
Lichenological herbarium of the Institute of
Botany and Ecology, University of Tartu (TU) is
the oldest and biggest of all those listed above.
The different collections of this herbarium, and
the curative work in TU are shortly characterized
in the present paper.
RESULTS AND DISCUSSION
Lichenological collections in TU include about 62
000 specimens. There are three main sections:
Estonian herbarium – ca. 24 000 specimens
Geographical collections – ca. 30 000 specimens
Comparison herbarium (incl. exciccatae) –
ca. 8 000 specimens.
All materials are arranged in the alphabetical
order of genera – inside each section – and in
the alphabetical order of species – inside the
genera.
The Estonian herbarium of TU is the biggest
collection of Estonian lichens in the world (about
24 000 specimens). Smaller collections of Estonian lichens are kept in other local herbaria as
well – ICEB, IE, TAL, TBA – and to a lesser extent
also in H, LD, RIG, S and UPS. Managing of
Estonian herbarium has been our priority task
during last four years. It has been re-arranged
recently according to the modern taxonomy (accepting small genera of parmelioid, cetrarioid,
lecideoid etc. lichens) in the course of verifying
82
Folia Cryptog. Estonica
all specimens to compile the new checklist of
Estonian lichens. Specimen databasing has
been started using the computer program
BRAHMS (Botanical Research And Herbarium
Management System). The system was initially
worked out by Denis Filer , Oxford Forestry Institute. Versions 3.3 and 3.4 have been applied
for managing lichenological herbarium in TU.
A module known as RDE (Rapid Data Entry),
linked to BRAHMS, has been used to enter various information connected with the herbarium
specimens. Configuration of RDE files has been
edited according to our special needs of lichenological herbarium. Every file includes data about
specimens of one species. Characters stored are
the following for every specimen: collector and
field number, number of the specimen in herbarium, additional collectors, genus, species,
author(s), infraspecific taxa, date of collection,
geographical locality, ecological group (based on
substratum), longitude, latitude and altitude
of the site of collection, local grid, herbarium
name where the collection and its duplicates are
kept, determiner and the date of identification,
presence of fruiting bodies and parasymbionts,
vitality, other ecological data, data of chemical
and anatomical studies, other notes by the determiner (Saag, 1996). After importing all these
data into BRAHMS, general specimen cataloguing, production of different categories of labels,
citation lists of specimens, floristic checklists
and other research proceedings become possible. Mapping program DMAP (Morton, 1997)has
been used together with BRAHMS to produce
distribution maps. At the moment, data of about
5000 lichen specimens of Estonia (materials of
all species from genera Acarospora, Adelolecia,
Amandinea, Amygdalaria, Biatora, Buellia,
Calicium, Carbonea, Cecidonia, Chaenotheca,
Chaenothecopsis, Cyphelium, Diplotomma, Farnoldia, Fuscidea, Graphis, Ionaspis, Lecanora,
Lecidea, Lecidella, Microcalicium, Mycocalicium, Miriquidica, Phaeocalicium, Placynthiella,
Polysporina, Porpidia, Protoparmelia, Psilolechia,
Pyrrhospora, Rimularia, Sagiolechia, Sarcogyne, Schaereria, Sclerophora, Sphaerophorus,
Stenocybe, Tephromela, Thelomma, Trapelia,
Trapeliopsis, Tremolecia, Rinodina) have been
entered into RDE and imported into BRAHMS.
The majority of data have been entered into the
database by Inga Jüriado, Piret Lõhmus, Mari
Sarv and Ave Suija; the technical managing of
the Estonian herbarium has mainly been carried
out by Inga Jüriado.
The oldest lichen specimens in the Estonian
herbarium date from the end of the last century.
They were collected by Andreas Bruttan, altogether about 260 species, from Abruka, Aegviidu, Haaslava, Hiiumaa, Muhu, Saaremaa, Selja
(Lääne-Virumaa), Tallinn, Tartu, Valga, Virtsu,
Viru-Nigula. Part of his herbarium (179 sheets,
with several species on each sheet, collected
from the present territories of both Latvia and
Estonia) are kept in RIG (Piterans, 1996). Other
older collections of Estonian lichens in TU were
raised by Paul Wasmuth (about 200 species),
and by Konstantin Mereschkowski (only a few
specimens preserved). Both persons worked
during the first two decades of this century
in Tallinn and its vicinity. Numerous materials (about 170 species) collected from Estonia
(Abruka, Keila, Kunda, Lihula, Rakvere, Paldiski, Pärnu, Saaremaa, Tallinn, Tartu, Viljandi,
Vilsandi, Väike-Maarja) by Veli Räsänen in 1927
and 1929, as well as some specimens collected
by L. Kari, K. Linkola and E. Häyrén, are kept
partly in TU and partly in H. Smaller collections
of Hilja Lippmaa, Voldemar Reinthal, Jaak Ruubel, Gustav Vilbaste, that date mainly from the
1930s, are also deposited in this section. In the
post-war period the biggest lichen collections
have been raised by Hans Trass.
Geographical collections raised by the coworkers of our institute (altogether about 30
000 specimens) are arranged into some bigger
geographical units:
I Extra-europian countries (mainly materials
from Canada)
II Europian countries
1. Foreign countries (mainly materials from
Finland and Sweden)
2. Former republics of the USSR (Latvia,
Lithuania, European part of Russia)
III Caucasus (Azerbaijan, Georgia)
IV Central Asia (Kazakhstan, Kirghizia, Tadjikistan, Turkmenistan, Uzbekistan)
V Siberia
VI Lake Baical region
VII Russian Far East.
83
Two latter collections are the most numerous. Field works in the vicinity of the Lake
Baical (Khamar-Daban Mountain Range in the
Irkutsk Territory and Buryatian Autonomic
Republic) were carried out in 1979–1982 by
a group of students and reseachers (A. Pärn,
K. Zobel, A. Roosma, T. Randlane etc.) led by
Hans Trass. More than 1000 sample quadrats
were analysed using air pollution monitoring
methods and about 10 000 specimens were
collected. This is the biggest geographical collection of lichens in TU. Herbarium of lichens
from the Russian Far East consists of ca. 8 500
specimens, collected from the Chabarovsk and
Primorje Territories, Peninsulas of Kamtchatka
and Tshukotka, Wrangel Island – by H. Trass,
A. Roosma, T. Randlane, S. Pärn-Eilart, V. Masing etc.
The following type materials are located in TU:
ASAHINEA CULBERSONIORUM Trass – holotype
CETRARIOPSIS LAII A. Thell & Randlane – holotype
CETRELIA ORIENTALIS Randlane & Saag – holotype
CETRELIA PSEUDOCOLLATA Randlane & Saag – holotype
CETRELIOPSIS PAPUAE Randlane & Saag – isotype
CLADONIA ALINII Trass – holotype
CLADONIA FAVILLICOLA Trass – holotype
CLADONIA TRASSII Ahti – isotype
HETERODERMIA INTERMEDIA Trass – holotype
RAMALINA STEVENSIAE Elix - isotype
TUCKNERARIA AHTII Randlane & Saag – isotype
XANTHORIA OXNERI S. Kondratyuk & Poelt – isotype
Comparison herbarium consists of ca. 8 000
specimens and has been maintained mainly by
exchanging specimens with or receiving gifts from
other herbaria. The following exsiccatae are represented in our collections rather numerously:
Herbarium Lichenum Fenniae (1875, 1882) and
Nylander & Norrlin, Herbarium Lichenum Fenniae (continued in 1921); Lichenotheca Fennica;
Lichenes Fenniae Exsiccati; Plantae Graecenses
(Lichenes); Lichenes Canadenses Exsiccati; Prof.
C. Mereschkowsky. Lichenes Rossiae Exsiccati;
V. P. Savicz. Lichenotheca Rossica; J. Suza: Lichenes Bohemoslovakiae; Reliquiae Suzaianae
(Table 1). Several other exsiccate materials are
represented in TU incompletely or in a few numbers only: Flora Exsiccata Austro-Hungarica;
Flora Hungarica Exsiccata; J. A. Elix: Lichenes
Australasici Exsiccati; Lichenes Americani Exsiccati; Lichenes Norvegiae Mediterranei Exsiccati; Syo Kurokawa: Lichenes Rariores et Critici
Exsiccati; Reliquiae Tuckermanianae etc.
During the last few years our herbarium
has been supplemented by a numerous and
valuable collection of lichens from the southern hemisphere (Australia, New Zealand, Papua New Guinea, Antarctica, Argentina, Brasil,
Chile) for which we are deeply indebted to our
compatriot Dr Heinar Streimann, curator of the
cryptogamic herbarium in Australian National
Herbarium (CANB). These specimens have also
been added to the comparison herbarium.
Lichenes Fenniae Exsiccati (Räsänen, Helsinki)
Fasc. I
1–2, 4–5, 7–22, 25–50
Fasc. II
51–89, 98, 100
Fasc. III
101–113, 116–123, 125–150
Fasc. IV
151–164, 167–185, 190–192,
194, 197–200
Fasc. V
201–226, 229–240, 244–245,
250
Fasc. VI
251–266, 268, 270, 275–300
Fasc. VII
301–347, 349
Fasc. VIII
351–362, 364–367, 369, 385,
393–394
Fasc. IX
403
Fasc. X
462–477, 500
Fasc. XI
501–515, 517–550
Fasc. XV
701–734, 736–750
Fasc. XVI
751–784, 786–800
Fasc. XVII
801–835, 837–850
Fasc. XVIII
851–860, 862–885, 887–900
Fasc. XIX
901–907, 909–945, 947, 949
Fasc. XX
952–960, 963–981, 983–1000
Lichenotheca Fennica (Räsänen, Kuopio)
Fasc. I
2, 4, 19, 20
Fasc. II
26–27, 39–43
Fasc. III
53, 64.
Fasc. IV
77–78, 81–83
Fasc. V
104, 121–122
Fasc. VI
126, 129–132, 147
Fasc. VII
153, 171–172
Fasc. VIII
181–183, 189–200
Fasc. IX
201, 225
Fasc. X
228–229, 232–233, 247–248
Fasc. XI
268–275
Fasc. XII
276–280, 291.
Fasc. XIII
301–325
Table 1. Exsiccate materials in TU
Name of exsiccatae
Series
Numbers present in TU
84
Folia Cryptog. Estonica
Fasc.
Fasc.
Fasc.
Fasc.
Fasc.
Fasc.
XIV
XV
XVI
XVII
XVIII
XIX
326–332, 334–350
351–361, 363–375
376–400
401–425
426–449
451–459, 461–465, 467–472,
474
Fasc. XX
476–493, 496–497, 499–500
Fasc. XXI
501–519, 521–523
Fasc. XXII
526–550
Fasc. XXIII
551–572, 574–575
Fasc. XXIV
576–580, 582–586, 588–590,
592–600
Fasc. XXV
601–608, 610–620, 622–625
Fasc. XXVI
626–628, 630–636, 638–649
Fasc. XXVII
651–652, 654–675
Fasc. XXVIII
676–680, 682–700
Fasc. XXIX
701–712, 714–722, 724–725
Fasc. XXX
726–744, 746–749
Fasc. XXXI
751–773, 775
Fasc. XXXII
776–800
Fasc. XXXIII
801–822, 824–825
Fasc. XXXIV
826–828, 830–850
Fasc. XXXV
851–875
Fasc. XXXVI
876–890, 892–900
Fasc. XXXVII
901–925
Fasc. XXXVIII
926–950
Fasc. XXXIX
951–975
Fasc. XL
976–991, 993–1000
Fasc. XLI
1001–1025
Fasc. XLII
1027–1034, 1036–1049
Fasc. XLIII
1051–1075
Fasc. XLIV
1076–1100
Fasc. XLV
1101–1113, 1115–1125
Fasc. XLVI
1126–1150
Fasc. XLVII
1151–1163, 1165–1175
Fasc. XLVIII
1176–1200
Fasc. XLIX
1201–1203, 1206, 1209–1210,
1213, 1215, 1218, 1220–1225
Fasc. L
1226–1250
Fasc. LI
1251–1252, 1254, 1256–1273,
1275
Fasc. LII
1276–1289, 1291–1300
Herbarium Lichenum Fenniae
Fasc. II
51–59, 61–66, 68–100
Fasc. III
101–132, 134–150
Fasc. IV
151–160, 162–186, 188–200
Fasc. VI
252–273, 276–280, 282–300
Fasc. VII
301–317, 319, 321–328, 330–
334, 336–350
Fasc. VIII
351–393, 395–400
Nylander & Norrlin, Herbarium Lichenum Fenniae
(continuatio)
451–502, 504–522, 524–566,
568–586, 588–736, 738–807
Plantae Graecenses. Lichenes
1–17, 20–43, 45–49, 51–60,
62–65, 200, 239, 250–288,
290–419, 421–447, 480
Lichenes Canadenses Exsiccati
1–6, 8–15, 17–18, 20–27,
29–227, 229–250
Prof. C. Mereschkowsky. Lichenes Rossiae exciccati
2–4, 6–8, 12–24, 27–29, 31,
33–35, 37–51, 53–55, 59,
62–63, 65–71, 73–74
V. P. Savicz. Lichenotheca Rossica
Decas III–XIII
21–70, 81–89, 91–109, 111–114,
116, 118–129
J. Suza: Lichenes Bohemoslovakiae
3, 6, 11, 37, 42, 45–50, 67–68,
71–72, 74, 76, 78–79, 86–87,
92–93, 97, 106–108, 110–114,
116, 118, 120, 126, 131, 135,
137–139, 143, 146, 148, 155–
156, 158, 161, 164–165, 167,
169, 175, 179, 183, 193, 199,
205, 209, 214, 215, 222–223,
225, 227, 229–232, 234–236,
251, 254, 260, 262, 266,
270, 275–277, 285, 289–291,
294–296, 298–300
Reliquiae Suzaianae.
1–28, 30–62, 64–100
REFERENCES
Kukk, T. 1997. Eesti soontaimede herbaariumid – hetkeseis ja perspektiivid. Rukkilill 5: 4–17.
Morton, A. 1997. DMAP for Windows. Version 6.4.
Piterans, A. 1996. Lichenological herbaria of Latvia. In
Fungi and lichens in the Baltic region (ed. Vimba,
E.), pp. 76–77. Riga.
Saag, A. 1996. Usage of computer program BRAHMS
in lichenological studies. In Fungi and lichens
in the Baltic region (ed. Vimba, E.), pp. 78–79.
Riga.
Folia Cryptog. Estonica, Fasc. 32: 85–96 (1998)
Time-space analyses of the British lichen flora, with particular
reference to air quality surveys
Mark R. D. Seaward
Department of Environmental Science, University of Bradford, Bradford BD7 1DP, U.K.
Abstract: The spread of air pollution, particulary sulphur dioxide, throughout the British Isles and elsewhere over the past
200 years is accurately reflected by the decline of lichens. Lichens have proved highly versatile for monitoring not only rising
but also decreasing levels of air pollution in general and, more recently, qualitative differences brought about by changes in
energy policies, clean air legislation and agricultural practices. The use of lichens as biological monitors in air quality surveillance
over wide geographical areas, where physicochemical equipment is often inadequate or indeed non-existent, is demonstrated
through time-space analyses of distribution data derived from the British Lichen Society’s Mapping Scheme.
Kokkuvõte: M. R. D. Seaward. Briti lihhenofloora aeg-ruumilised analüüsid, rõhuasetusega õhu kvaliteedi
hinnangule.
Õhu saaste, eriti vääveldioksiidi levik kõikjal Briti saartel viimase 200 a. jooksul peegeldub samblike vähenemises. Samblikud
on osutunud mitmekülgseteks indikaatoriteks, võimaldades täheldada mitte ainult õhu saastatuse tõusu ja langust üldjoontes,
vaid ka viimasel ajal ilmnenud kvalitatiivseid erinevusi, mida põhjustavad muudatused energiapoliitikas ning põllumajanduses.
Demonstreeritakse samblike kasutamist õhu kvaliteedi biomonitoridena samblike levikuandmete aeg-ruumiliste analüüside
abil.
INTRODUCTION
The value of lichens as monitors of air pollution
is undisputed and has captured the attention
not only of lichenologists but also of biological and environmental scientists in general, as
reflected by the endless flood of publications
on this topic. This paper provides an historical
review of the methodologies developed, and the
different circumtances in which they have been
employed, in order to monitor quantitative and
qualitative changes in the British isles.
Although several early British botanists
noted that lichens did not thrieve in polluted
atmospheres (e.g. Borrer c. 1812 in Turner &
Borrer, 1839; Grindon, 1859; Macmillan, 1861),
it was the famous Finnish lichenologist William
Nylander, at that time working in Paris, who
suggested that lichens might be used as indicators of air quality: his studies in the Jardin
du Luxembourg showed first the decline in
(Nylander, 1866) and later the disapperance of
(Nylander, 1896) its epiphytic flora. More than
90 years were to elapse before epiphytic lichens
returned to the Jardin du Luxembourg (Seaward
& Letrouit-Galinou, 1991).
British air pollution studies involving lichens in the second half of the 19th century
and the first half of the 20th century, with few
exceptions, were mainly concerned with casual
observations on the impoversihment of urban
floras or on the general decline in species over
local areas (reviewed in Seaward, 1993). Such
studies were instrumental in establishing baseline data which could be used later in time-space
analyses. Hawksworth et al. (1973), for example,
incorporated species data of earlier fieldwork in
their reconstruction of the change in air quality
in Epping Forest, near London, over the period
1784–1970; based on the nature and diversity
of the epiphytic flora, and employing the scale
devised by Hawksworth and Rose (1970), mean
winter sulphur dioxide levels increased from less
than 30 to 70–125 µg/m3 over that time. Further
studies by Hawksworth and McManus (1992)
have shown a remarkable recovery of this flora
consequent upon environmental amelioration.
As well as their value in time-space analyses,
such distribution studies made it possible for
the first time to compare the severity, extent and
pattern of air pollution between cities (Hawksworth, 1971, 1973; Laundon, 1973).
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Folia Cryptog. Estonica
Several highly influential papers appeared in
the 1950s which were to provide a springboard
for the future development of lichen – air pollution studies. The first of these, by Jones (1952),
contained details of lichen floras of tree boles at
sites in England where air pollution levels were
known. This made it possible for the first time to
construct pollution monitoring scales based on
the composition of the lichen flora. Based upon
this rudimentary scale, many more sophisticated bioindicational scales were developed. Prior
to this period, an ‘air pollution hypothesis’ had
evolved to explain the paucity of lichens in urban
areas and the gradual decline in the European
lichen flora generally since the Industrial Revolution at the end of the 18th century. However,
the constituents of air pollution responsible for
these phenomena had not yet been determined;
indeed, Jones (1952) and others had indicated
smoke and/or soot deposition to be the main
cause. Today, sulphur dioxide is recognized as
being the major culprit affecting lichen vigour.
However, other air pollutants are often implicated, and ever-changing environments, the
exacerbatory/synergistic/buffering effects of
two or more constituents of a complex pollution
burden also affect lichen performance.
It was two publications in 1958 by the
Dutch and Swedish scientists Barkman and
Skye respectively which most influenced future
research on lichens and air pollution in Britain
and elsewhere. Since most field studies on this
subject relied on the morphological nature and
ecological performance of epiphytic lichens,
Barkman’s (1958) monumental treatise on the
phytosociology and ecology proved of immense
value in establishing protocols for description
and measurement. Skye (1958) recognized sulphur dioxide to be the overriding factor affecting
the lichen flora around an oil-shale works. This
and his detailed investigation of the compostion and distribution of lichens in and around
Stockholm (Skye 1968) provided models for a
very large number of subsequent investigations.
Casual observations of differences in the composition and numbers of species between lichen
floras in polluted and unpolluted areas in earlier
studies had been replaced by a simplistic zonal
interpretation based on the concept of ‘desert’,
‘struggle’ and ‘normal’ devised by Sernander
(1926), to be superseded in its turn, largely as
a consequence of Skye’s work, by more detailed
zonal studies in the 1960s. Investigations in
Britain at that time were centred on individual
industrial plants, towns or cities, or on urban/
industrial complexes, but regional and national
studies were to follow with the establishment of
the British Lichen Society’s Distribution Maps
Scheme which commenced in September 1963
(Hawksworth & Seaward, 1990); by applying
Sernander’s concept of ‘desert’ to these maps it
has been possible to rank species susceptibility
to air pollution (see below).
The key paper on zonal mapping was by
Hawksworth and Rose (1970) who devised a
qualitative scale using epiphytic lichens for
the estimation of sulphur dioxide air pollution.
Their scheme, designed for use in England and
Wales, is based on simple phytosociological
analyses which delineate zones equated with
mean winter sulphur dioxide levels. This scale
has stood the test of time, proving of immense
value in air pollution monitoring throughout the
British Isles, as well as forming the basis, often
only requiring minor adaptation, for scales in
Europe and elsewhere. It has been extensively
used to monitor the extent or spread of air pollution, particulary sulphur dioxide; however, in
areas experiencing reduced levels of, or qualitative changes in, pollution as a result of clean
air legislation, it may be less effective (Seaward,
1987; Van Haluwyn & Lerond, 1988; Richardson, 1988). It should be noted that its biological
scale is logarithmic in nature (Seaward, 1993,
1997): although major reductions in the sulphur
dioxide level at the polluted end of the scale may
have little or no effect on lichen species diversity,
moving from a polluted to a less polluted situation exponentially increases the effect on the
lichen vegetation, so that even a minor increase
in sulphur dioxide would dramatically reduce
species diversity.
The establishment of baseline data from
zonal studies and transect work employing strict
methodological protocols, regulary updated to
provide valid data, is highly effective in timespace analyses aimed at demonstrating environmental changes, for good or ill, or in predictive
interpretation through modelling (HendersonSellers & Seaward, 1979; Seaward, 1997).
87
AMELIORATING ENVIRONMENTS
The undoubted improvement in the lichen flora
in many parts of the British Isles during recent
years has been clearly demonstrated (Henderson-Sellers & Seaward, 1979; Seaward, 1980,
1993, 1997; Bates & Farmer, 1992; Rose &
Hawksworth, 1981; Hawksworth & McManus,
1989). However, it is evident from the above
works that the nature of a re-establishing lichen
flora differs from that formerly wiped out by air
pollution. From our knowledge of the behaviour of lichens subjected to sulphur dioxide air
pollution, it has been possible to rank species
according to their sensitivity to this pollutant,
but variations in replacement assemblages will
necessitate modifications to some bioindicational scales to account for these hierarchical
changes.
The distribution of Usnea spp. in the British
Isles illustrates the type of change which has occured since about 1800. This genus, at one time
widespread and luxuriant, had over the course
of the next 160 or more years almost entirely
disappeared from a major area of England and
Table 1. Sensitivity to air pollution of selected species ranked according to scarcity in a prescribed
region (1287 grid squares 10 km × 10 km) of Britain, the list commencing and finishing with the
most and least sensitive species respectively
% no. of squares from which sp.
is absent (with ranking)
Lobaria pulmonaria
Ramalina fraxinea
Thelotrema lepadinum
Lecanactis abietina
Parmelia perlata
Physcia aipolia
Graphis elegans
Ramalina fastigiata
Graphis scripta
Enterographa crassa
Pertusaria hymenea
Parmelia revoluta
Calicium viride
Lecidella elaeochroma
Usnea subfloridiana
Hypogymnia tubulosa
Parmelia subaurifera
Parmelia caperata
Platismatia glauca
Cliostomum griffithi
Lecanora chlarotera
Parmelia subrudecta
Ramalina farinacea
Parmelia glabratula
Evernia prunastri
Lecanora expallens
Buellia punctata
Hypogymnia physodes
Parmelia sulcata
Lecanora conizaeoides
in Sept. 1992
in Nov. 1997
96.04 (1)
91.06 (2)
82.67 (3)
79.72 (4)
77.78 (5)
77.54 (6)
76.61 (7)
74.90 (8)
73.89 (9)
73.19 (10)
71.95 (11)
69.77 (12)
66.43 (13)
64.18 (14)
62.08 (15)
60.76 (16)
54.86 (17)
54.47 (18)
53.61 (19)
50.12 (20)
48.41 (21)
45.53 (22)
42.81 (23)
41.96 (24)
41.86 (25)
30.92 (26)
29.76 (27)
21.06 (28)
20.05 (29)
1.24 (30)
95.80 (1)
90.21 (2)
81.04 (3)
77.00 (4)
71.48 (8)
75.29 (5)
75.06 (6)
72.96 (7)
71.41 (9)
71.25 (10)
70.40 (11)
64.96 (12)
64.72 (13)
61.07 (14)
54.31 (15)
52.76 (16)
41.10 (21)
48.02 (18)
50.04 (17)
46.08 (19)
42.89 (20)
40.71 (22)
35.35 (24)
35.51 (23)
23.00 (25)
16.94 (27)
21.13 (26)
15.31 (28)
12.12 (29)
0.85 (30)
% change
- 0.24
- 0.85
- 1.63
- 2.72
- 6.30
- 2.25
- 1.55
- 1.94
- 2.48
- 1.94
- 1.55
- 4.81
- 1.71
- 3.11
- 7.67
- 8.00
- 13.76
- 6.45
- 3.57
- 4.04
- 5.52
- 4.82
- 7.46
- 6.45
- 8.86
- 13.98
- 8.63
- 5.75
- 7.93
- 0.39
88
Folia Cryptog. Estonica
Wales covering at least 68,000 km2 (Fig. 1), and
at least 6,000 km2 of lowland Scotland, mainly
as a result of the increase in atmospheric pollution. However, since the implementation of
the Clean Air Acts of 1956 and 1968 in Britain,
sulphur dioxide concentrations have declined
dramatically in urban/industrial areas, with
less dramatic reductions, or indeed occasional
rises due to changes in pollutant dispersion
techniques, in rural areas, so that much of
Britain is now experiencing a more homogeneous distribution of this pollutant. The re-establishment of Usnea spp. (mainly U. subfloridana),
ususally on Fraxinus and Salix, at numerous
sites throughout the British Isles during recent
years may largely reflect decreases in pollution
levels. There is some evidence that Ramalina
farinacea, Evernia prunastri and Bryoria fuscescens are exhibiting a similar response. It would
appear, however, that the stability of these taxa
of these sites is tenuous, since small thalli may
only succeed in establishing themselves for one
or two years. Nevertheless, it is now proving possible to monitor ameliorating environments on
a relatively large scale through the use of lichen
mapping (Fig. 1).
Table 1 has been assembled from detailed
comparative studies of the distributions of selected lichen species within a prescribed region
of 128,700 km 2 (Fig. 2): comparisons have
been made between (a) field data derived from
the British Lichen Society’s Mapping Scheme
up to May 1980, some of which were used
in the preparation of the first volume of the
Lichen Atlas (Seaward & Hitch, 1982) and (b)
that derived from the current (November 1997)
database to demonstrate the large-scale monitoring capabilities of lichens. These data show
the strong correlation between a lichen’s sensitivity to air pollution (mainly sulphur dioxide)
and the area from which it had declined, as can
be seen from the distributions of, for example,
Ramalina farinacea (Fig. 3), Usnea spp. (Fig.
1) and Parmelia perlata (Fig. 4), their ‘deserts’
in c. 1970 extending over 45,000, 68,000 and
88,000 km2 respectively. However, it is also clear
that rankings (cf. Seaward, 1992, Table 1) are
changing due to the different abilities of species
to re-establish themselves in areas experiencing
atmospheric ameloration. Hence, since the previous analysis undertaken less than five years
ago, some species, Lecanora expallens, Parmelia
subaurifera, Evernia prunastri, etc. (see Table 2)
have a greater capacity to adapt to these new
regimes.
Regional or ‘blanket’ pollution over wide
areas has reduced lichen diversity, but has favoured expansion of a relatively small number
of species formerly having narrower ecological
requirements and more restricted distributions,
such as Parmeliopsis ambigua (Fig. 5), Scoliciosporum chlorococcum (Fig. 6) and Xanthoria
polycarpa (Fig. 7). Some species which are
moderately tolerant of sulphur dioxide air pollution have been able to spread in areas where
other species have been eliminated. Parmelia
laciniatula and P. elegantula, first recorded in
the British Isles as recently as 1933 and 1965
respectively, are two such species and detailed
distributional studies show them to be on the
increase due to both quantitative and qualitative
changes in air pollution (Seaward, 1993).
‘Acid rain’ can affect lichens both directly or
indirectly through the acidification of substrata.
Parmeliopsis ambigua (Fig. 5), for example,
which was formerly to be found mainly on
decorticate conifer wood in Scotland, has, over
the past few decades, undergone a remarkable
expansion onto deciduous trees in moderately
polluted areas throughout Britain (cf. Seaward,
1992, 1993), in response to increased bark acidification. The development of more oligotrophic
epiphytic lichen floras, including such species as
Micarea nitschkeana (Fig. 8), has been reported
in the British Isles as a result of changes in the
nature of pollutant emissions, but their present
distribution is poorly understood; since many
of the microlichens in these floras are difficult
to determine, particulary in the field, their use
as monitors of ‘acid rain’ will be limited. Furthermore, deposition of nitrates via acid rain,
emanating from NOx emissions, can produce a
fertilization effect (see below), as yet little studied, which encourages numerous lichen species
to flourish, at least over the short term.
Changes in the present lichen composition
in many areas of the British Isles are often as
much a reflection of increased eutrophication as
a reduction in sulphur dioxide levels. Numerous
species which are now spreading into suburbs
are equally successful elsewhere but formerly
had narrower ecological requirements and more
restricted distributions. They are probably fa-
Fig. 1. Distribution of Usnea spp. in the British Isles (January 1998) to show its dissappearance from an area of c. 68,
000 km2 during the period c. 1800-1970, and its subsequent
re-establishment at numerous sites (in 150 recording units
10 km × 10 km) within that area; open dot = pre-1960 (usually 19th c.) records, filled dot = 1960 onwards records.
Fig. 2. Prescribed region of Britain (mainly England), much
of which has been subjected to increasing air pollution over
the past two centuries. The distributions of selected and
potentially widespread species from this delimited area
(1287 recording units 10 km × 10 km) have been ranked
in Table 1 according to their susceptibility to air pollution,
based on the number of recording units from which they
are currently absent (i.e. lichen ‘desert’).
89
90
Folia Cryptog. Estonica
Fig. 3. Distribution of Ramalina farinacea in the British
Isles (January 1998) to show its dissappearance from an
area of c. 45, 000 km2 during the period c. 1800-1970, and
its subsequent re-establishment at numerous sites (in 173
recording units 10 km × 10 km) within that area (cf. Seaward
& Hitch, 1982, map 140); open dot = pre-1960 (usually 19th
c.) records, filled dot = 1960 onwards records.
Fig. 4. Distribution of Parmelia perlata in the British Isles
(January 1998) to show its dissappearance from an area
of c. 88, 000 km2 during the period c. 1800-1970, and its
subsequent re-establishment at numerous sites (in 82 recording units 10 km × 10 km) within that area (cf. Seaward
& Hitch, 1982, map 102); open dot = pre-1960 (usually 19th
c.) records, filled dot = 1960 onwards records.
Fig. 5. Distribution of Parmeliopsis ambigua in the British
Isles (January 1998) to show how it is extending its geographical range in response to increased bark acidification
(to pH 3.0), at a rate of more than 11 recording units (10
km × 10 km) per annum over the past 15 years (cf. Seaward
& Hitch, 1982, map 116; Seaward, 1992).
Figur 6. Distribution of Scoliciosporum chlorococcum in
the British Isles (January 1998) to show its spread into
both rural and suburban areas in response to changes in
atmospheric pollution, being monitored at a rate of c. 26
recording units (10 km × 10 km) per annum over the past
9 years.
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Folia Cryptog. Estonica
Table 2. Ranking of selected species (see Table
1) according to their ability to recolonize areas
experiencing amelioration of air pollution within
a prescribed region of Britain (see Fig. 2) over
the period September 1992 and November 1997.
Lecanora conizaeoides, with almost complete
coverage of the area under study in September
1992, has been omitted from analysis
Species significantly extending their distribution
by 7.0 % or more
Lecanora expallens
Parmelia subaurifera
Evernia prunastri
Buellia punctata
Hypogymnia tubulosa
Parmelia sulcata
Usnea subfloridiana
Ramalina farinacea
Species extending their distribution by 4.5 to
6.5 %
Parmelia caperata
Parmelia glabratula
Parmelia perlata
Hypogymnia physodes
Lecanora chlarotera
Parmelia subrudecta
Parmelia revoluta
Species extending their distribution by only 2.0
to 4.5 %
Cliostomum griffithii
Platismatia glauca
Lecidella elaeochroma
Lecanactis abietina
Graphis scripta
Physcia aipolia
Species scarcely extending their distribution
Ramalina fastigiata
Enterographa crassa
Calicium viride
Thelotrema lepadinum
Pertusaria hymenea
Graphis elegans
Ramalina fraxinea
Lobaria pulmonaria
voured by reduced competition, but urban and
industrial (mainly alkaline) dusts, acid rain
derived from NOx emissions, soil fertilizers and
other agrochemicals promote interesting lichen
floras (Seaward, 1997). A fertilization effect on
the lichen flora may manifest itself as a consequence of a reduction in the sulphur dioxide
burden, but there are instances where this
effect generated from factory emissions may
ameliorate a local sulphur dioxide problem
(Seaward, 1990).
The recent spread of some species into city
suburbs and elsewhere may be promoted by
windborne nutrient-enriched dusts (Seaward,
1997). By 1973, Ramalina farinacea had disappeared from c. 47,300 km2 of England (Fig.
3), an area incorporating 457 of the 10 km × 10
km recording units (Seaward, 1992); since then,
this taxon has been recorded from 78 of these
units, showing its successful re-colonization not
only of sites formerly experiencing sulphur dioxide levels of 60 µm/m3 or more, but also those
where barks of trees and shrubs have become
moderately nutrient-enriched. The increasing
occurrence of Xanthoria polycarpa throughout
the British Isles (Fig. 9), where mean winter sulphur dioxide levels have fallen below 60 µg/m3,
is due in no small measure to agrochemicals
which have promoted its growth even in the
inner suburbs of most large towns and cities
(Seaward, 1997). In 1980, it had been recorded
from 580 recording units, and by 1991, it occurred in 869 (Seaward, 1992); in June 1993,
this figure has risen to 966 (Seaward, 1993) and
by November 1997 had reached 1249.
From an ever-present rich air spora, only
a few species are capable of surviving the
crucial stages of establishment, germination
and development in polluted environments.
Once established, such toxitolerant species are
strongly competitive, with a high reproductive
capacity and a tenacious hold on the substrata
they colonize; they can create monovegetational
cover, even when a reduction in air pollution
level would presuppose reinvasion by formerly
successful species. This is illustrated by the rise
to dominance of Lecanora conizaoides throughout the British Isles, and much of Europe, over
the past half century. It was first recorded in
Britain in about 1860, since when its spread has
been dramatic: by 1953, it occupied c. 121,800
km2; by 1973, it occupied c. 185,300 km2, and
by 1997 this had extended to c. 207,900 km2
Fig. 7. Distribution of Parmelia elegantula in the British Isles (January 1998) to show how it is extending its
geographical range, in response to qualitative changes in
atmospheric pollution, at a rate of more than 4 recording
units (10 km × 10 km) per annum over the past 15 years
(cf. Seaward & Hitch, 1982, map 94).
Fig. 8. Distribution of Micarea nitschkeana in the British Isles (January 1998) to show how it is extending its
ecological amplitude and geographical range, probably in
response to qualitative changes in atmospheric pollution,
at a rate of c. 7 recording units (10 km × 10 km) per annum
over the past 9 years; open dot = pre-1960 (usually 19th c.)
records, filled dot = 1960 onwards records.
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Folia Cryptog. Estonica
(Fig. 10). Air polluted environments provide an
ideal milieu for the spread of this taxon, and
even amelioration, resulting from the implementation of clean air policies, has not as yet
broken its monopoly of substrata.
CONCLUDING REMARKS
Profitable lines of enquiry into determining
the effects on lichens of the new air pollution
regimes should include long-term field techniques involving stringent ecological and phytogeographical criteria. To a large extent, such
studies have been made possible in the British
Isles through a comprehensive ongoing programme of detailed lichen mapping carried out
by members of the British Lichen Society since
1963. Although detailed lichen atlases (eg. Seaward & Hitch, 1982; Seaward, 1995 onwards)
contribute significantly to the documentation
of atmospheric regimes, mapping programmes
must be supported by extensive field studies in
a wide range of selected habitats in critical areas
where more immediate changes can be scientifically diagnosed by means of bioindicational
techniques complemented by adequate monitoring equipment. Intensive lichen monitoring
is a necessary component of any programme
aimed at effective long-term observation of air
pollution.
There is a wealth of evidence to demonstrate the selective decline of lichen species over
much wider areas during the past two centuries, mainly attributable to the rise in sulphur
dioxide levels, although other pollutants and
changing forestry and agricultural practices
are also implicated. Lichen mapping is often
a race against time, particulary in those areas
where clean air legislation and changes in energy policies and industrial practices have been
implemented: both qualitative changes in, and
dramatic reductions in the level of, air pollution
are reflected in the consequent reinvasion by
lichens. At the same time, a less dramatic, but
nevertheless significant, rise in pollution level
over wide geographical areas due to changes in
pollutant dispersion techniques has resulted in
the loss of a number of lichen species, although
others have been able to exploit/tolerate the new
atmospheric regimes.
Lichens have been used as monitors of environmental conditions for many years. How-
ever, any successful use of lichen bioindicators
must depend upon the rigorous definition of
the monitoring scales adopted and a complete
analysis of all the environmental parameters
which affect their propagation, establishment
and subsequent growth. The use of bioindicators has the obvious advantage of permitting
long-term monitoring without widespread
establishment and maintenance of costly and
sophisticated equipment. Unfortunately, most of
the existing bioindication scales rely on species
diversity counts, or at least on a fairly detailed
understanding of taxonomy, thereby making
the techniques as difficult to employ as sophisticated measuring equipment. It is therefore a
serious cause for concern that taxonomic skills
are no longer considered an essential component
of many biological educational programmes in
Britain and elsewhere.
Physical and/or chemical surveys do
not necessarily measure those air pollutants
which are harmful to biological materials.
On-site measuring devices are limited both
in number and scope; furthermore, they are
unevenly spread over a country, usually being
concentrated in urban and industrial complexes
(Seaward, 1993). Not only the implementation of
a clean air policy resulted in widespread ‘blanket pollution’, but the location of industry and
power stations away from cities has also added
to the atmospheric burden over wide areas.
The use of corroborative information derived
from bioindication surveys to support the limited data available from rural sites is strongly
recommended. It will never be completely possible to replace direct physical and chemical
measurements of air pollutant concentrations
by the use of bioindicators; nevertheless, both
approaches are necessary for making detailed
or large-scale surveys of the distribution of air
pollutants, where the extensive use of technical
equipment is costly or impractical.
DEDICATION
This paper is respectfully dedicated to Professor Hans Trass whose extensive ecological
and phytogeographical studies of the Estonian
lichen flora have made it possible to exploit
fully their potential to monitor his country’s
environment.
Fig. 10. Distribution of Lecanora conizaeoides in the British
Isles (January 1998), being rarely encountered to the west
and north of the grid lines indicated prior 1950.
95
Fig. 9. Distribution of Xanthoria polycarpa in the British
Isles (January 1998) to show how it is extending its geographical range, in response to nutrient-enrichment of
bark via agrochemicals, alkaline dusts and acid rain from
NO3 emissions, at a rate of more than 40 recording units
(10 km × 10 km) per annum over the past 15 years (cf.
Seaward & Hitch, 1982, map 176; Seaward, 1992) ); open
dot = pre-1960 (usually 19th c.) records, filled dot = 1960
onwards records.
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Folia Cryptog. Estonica
REFERENCES
Barkmann, J. J. 1958. Phytosociology and Ecology of
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Folia Cryptog. Estonica, Fasc. 32: 97–105 (1998)
Bryological comparisons between Australia and northern Europe
Heinar Streimann
Centre for Plant Biodiversity Research, Cryptogamic Herbarium, Australian National Herbarium,
Box 1600, Canberra, A.C.T., 2601, Australia
Abstract: Comparison and discussion on the similarities between the bryophyte floras of Australia and northern Europe
– Denmark, Estonia, Finland, Iceland, Norway, Russia (NW) and Sweden – are made. Currently there are about 112 moss and
14 liverwort species that are common to both Australia and northern Europe, i.e. 10% and 3% respectively of the relevant
Austalian floras. The majority of these species are restricted to Southeast Australia (including Tasmania). Of the 126 species
compared 8 (2 liverwort) are cosmopolitan species, and 13 (1 liverwort) species may possibly be so; 12 species (all mosses)
are recent introductions by man to Australia, and a further 9 are considered as possible introductions; 72 (6 liverwort) species
are considered as bipolar disjunctive, including the recently confirmed Scorpidium scorpioides. Comments are also included on
the Australian distribution and habitat preferences for many of these species.
Kokkuvõte: H. Streimann. Austraalia ja Põhja-Euroopa brüofloorade võrdlus.
Käsitletakse Austraalia ja Põhja-Euroopa (Taani, Eesti, Soome, Islandi, Norra, Loode-Venemaa ja Rootsi) brüofloorade
sarnasust. 112 lehtsambla ja 14 helviksambla liiki (vastavalt 10% ja 3%) on ühised Austraaliale ja Põhja-Euroopale. enamus
neist liikidest levib Kagu-Austraalias (k.a. Tasmaania). Ühistest liikidest 8 (2 helviksammalt) on kindlalt ja 13 (1 helviksammal)
tõenäoliselt kosmopoliidid; 12 liiki (kõik lehtsamblad) on teadaolevalt ja 9 arvatavalt tulnuksamblad Austraaliasse; 72 (6
helviksammalt) on levikult bipolaarsed, sh. hiljuti tuvastatud Scorpidium scorpioides. Paljude liikide puhul tuuakse andmeid
nende kasvukohtade ja leviku kohta Austraalias.
INTRODUCTION
Very few studies have been undertaken to compare the bryophytes of the southern hemisphere
to other regions of the world, mainly because of
the lack of bryological information from the Australian region. Schofield investigated the bipolar
mosses in the southern hemisphere, especially
New Zealand (1974), while Piippo & Koponen
(1997) investigated the affinities of the western
Melanesian moss flora to Australia. The affinities
of Asiatic and Australian liverwort floras were
investigated by Piippo (1992).
This investigation was primarily initiated by
numerous queries in discussions with northern
European and Australian bryologists who were
interested in the similarities of the bryoflora between the two regions. This work compares the
similarities in the bryophyte floras of Australia
and northern Europe, especially Scandinavia
and northern European Russia (Fig. 1). Northern
European Russia comprises the regions of European Arctic, Northwest and Northeast Russia
and northern Ural Mountains (Ignatov & Afonina, 1992). Many of these similarities no doubt
will apply also to other northern regions.
Many people, including plant botanists, are
not aware of how widespread many cryptogams
found in Australia are and they are surprised to
learn that Australia shares many species with
colder northern regions. The bryophyte relationships of tropical Australia with Southeast Asia,
and temperate Victoria and Tasmania. with the
Nothofagus forests of South America are easier
to understand than relationships to the colder
northern hemisphere.
The vegetation and climate differences
between these two geographically distant areas are considerable. Although Australia has
warm to hot arid to semi-arid regions, snow
is a regular feature for part of the year in the
sub-alpine regions of New South Wales, Victoria
and Tasmania. Sub-alpine in this work refers
to the shrub and herb dominated areas above
the tree line.
The Australian vascular flora has adapted
to the climatic extremes and generally poor soils
which resulted in a distinctive vegetation. While
vascular plant geographic distribution does not
extend far from Australia, endemic species comprise 85% of the flora (Anon., 1996). Distribution
of the moss flora is more widespread with only
98
Folia Cryptog. Estonica
Fig. 1. Territories of compared bryophyte floras in northern Europe and Australia.
99
about 38 % endemism (Streimann & Curnow,
1989, with subsequent unpublished amendments due to revisions).
Vascular plant species diversity is greatest
in Queensland (Hnatiuk, 1990) because of its
Nothofagus, tropical to sub-tropical forests,
monsoon scrub, arid western regions and the
higher peaks which have a distinctive flora.
However, New South Wales has the richest recorded bryophyte diversity because it also has
sub-alpine vegetation and Lord Howe Island in
the Pacific Ocean.
Bryophyte studies in Australia lag far behind
those of the vascular flora. Exploration, though
woefully inadequate, still reveals species previously unreported from Australia. Revisions will
possibly decrease the number of endemic species, especially from tropical Australia as many
of those species may prove to be geographical
variations of widespread Southeast Asian species. The number of bryophyte species may
remain stable or decrease. Some names may
still be mis-applied in Australia e.g. Bryoerythrophyllum recurvirostre, Pohlia bulbifera, while
some large groups e.g. Bryum, Pottiaceae and
Sphagnum are still inadequately understood in
Australia. Because of our limited knowledge of
Australian (except possibly Tasmanian) cryptogams, the diversity and, geographic relationships
have not been investigated fully.
This study is based mainly on data from Damsholt et al. (1969), Frisvoll et al. 1995, Hallingbäck & Söderström (1987), Ignatov & Afonina
(1992), Ingerpuu et al. (1994), Jóhannsson
(1983), Kannukene et al. (1997), Konstantinova et al. (1992), Koponen et al. (1977), Nyholm (1986, 1989, 1993) for northern Europe.
For Australia Scott & Bradshaw (1986) and
Streimann & Curnow (1989) were used. All
Australian species of doubtful occurrence were
ignored.
It must be borne in mind that the analyses
of Australian bryophytes, and especially liverworts, are incomplete as most old collections
lack adequate data and some recent collectors
do not record habit and habitat data. Also many
species and areas are still under collected and
therefore their range, habitat preferences and
distribution are not known with certainty.
Bryophyte numbers for the countries in this
study are given in Table 1. The number of Australian moss species in common with the various
European countries studied is much greater
than was expected, but still represents only a
small component of the Australian moss flora.
However, when the number of species common
with individual northern European countries or
regions are compared they comprise 14–21% of
moss flora and only 5–9% of the liverwort flora
(Table 2).
Table 1. Numbers of bryophyte species in Australia and northern Europe
Country
Australia
Denmark
Estonia
Finland
Iceland
Norway
Russia (NW)
Sweden
Mosses
Liverworts
1150
436
405
591
416
755
604
753
436
146
116
226
154
282
221
266
Table 2. Percentage of the bryophyte flora of
each northern European country also occurring
in Australia
Country
Denmark
Estonia
Finland
Iceland
Norway
Russia (NW)
Sweden
Mosses (%) Liverworts (%)
21
19
16
14
15
15
15
8
9
6
6
5
6
5
Most species (Table 3) in common with northern
Europe are concentrated in the cooler moister
habitats of Southeast Australia (including
Tasmania) where sub-alpine regions occur, Kosciuszko National Park, and include areas such
as the High Plains of Victoria and Highlands of
New South Wales, Victoria and Tasmania.
100 Folia Cryptog. Estonica
Table 3. Number of moss species in Australian
states common with northern Europe
State
Mosses
Queensland
New South Wales
Aust. Cap. Terr.
Victoria
Tasmania
Macquarie Is.
South Australia
Western Australia
Northern Territory
28
78
54
75
74
23
47
30
6
A further 35 moss and 16 liverwort genera are
common to both regions, even though the species differ.
BIOGEOGRAPHICAL COMPARISONS OF
MOSSES
The moss species common to Australia and
northern Europe that have been analysed include cosmopolitan, disjunctive bipolar and
introduced species. For these species, and the
majority of the remaining species, comments
are made on their Australian range and habitat
preferences.
Cosmopolitan mosses
Such species generally thrive in disturbed or
open areas and many follow man and his habitation. They are mostly hardy species. It is nearly
impossible to ascertain if they were introduced
prior to European settlement of Australia.
Often there are various interpretations as to
what constitutes a cosmopolitan species and
an introduced species. Species can be spread
unintentionally by birds, animals and man, and
this can be considered as a natural function of
plant migration. Species in Australia considered
as cosmopolitan are: Bryum argenteum, B. caespiticum, B. capillare, Funaria hygrometrica, Leptobryum pyriforme, Polytrichum juniperinum.
Other species that may fit into this category
are: Amblystegium serpens, Aulacomnion palustre, Brachythecium plumosum, Bryum dichotomum, Ceratodon purpureus, Grimmia laevigata,
G. pulvinata, Polytrichum commune, Schistidium
apocarpum, Tortula muralis, T. papillosa which
is generally found in abundance on exotic trees
and Weissia controversa.
Introduced species
Schofield (1974) lists 14 moss species introduced by man to New Zealand, the majority
of which also occur in Australia. There are 9
species in Australia that are bipolar as well as
introductions and they are generally considered
in this section. The following 12 species are considered native to the northern hemisphere, but
introduced to Australia: Barbula unguiculata,
Brachythecium albicans, B. mildeanum, Bryum
radiculosum, B. rubens, Calliergonella cuspidata,
Phascum cuspidatum, Pottia truncata, Pseudoscleropodium purum, Pterygoneurum ovatum,
Rhytidiadelphus triquetrus, Tortula laevipila.
Catcheside (1980) suggests that the following species may have also been introduced to
Australia: Acaulon mediterraneum, Aloina rigida,
Amblystegium serpens, Barbula hornschuchiana,
Brachythecium rutabulum, Kindbergia praelonga,
Physcomitrium pyriforme, Pottia davalliana, Pottia starckeana.
These introductions can be assumed to be
the result of European settlement in Australia as
the majority of collections were found in anthropogenic habitats or near to them. Further field
studies are required to resolve their status.
Rhytidiadelphus triquetrus is a recent introduction to Australia and New Zealand (Dalton,
1997). It is extremely common in the wet, western Tasmania. This species aggressively invades
lawns (especially golf courses), exposed waste
land and roadsides where grass cutting apparently assists in dispersal.
Two Australasian species have taken the reverse
track and have found the European environment
suitable and reached the southern part of the
study area. They are Campylopus introflexus
(Denmark, Sweden, Norway) and Orthodontium
lineare (Denmark, Sweden).
Bipolar disjunctive mosses
Du Rietz (1940) defined these taxa as “distributed both in the boreal and austral zone but absent from the tropical lowlands, with or without
101
intermediate populations in tropical mountain
areas”. This definition was used by Schofield
(1974) and has been followed here.
Schofield & Crum (1972) discussed disjunction in bryophytes and commented comprehensively on the various explanations that
were offered for this distribution pattern. They
mentioned only two species relevant to Australia, Sanionia uncinata and Pseudoscleropodium
purum, and provided distribution maps.
Subsequently Schofield (1974) analysed the
bipolar disjunctive mosses of New Zealand, but
such studies have not been undertaken previously for Australian bryophytes. He included
distribution maps for some species, of which
Aulacomnium palustre, Ceratodon purpureus,
Climacium dendroides, Desmatodon heimii,
Orthodontium lineare, Pottia truncata, Sanionia
uncinata and Ulota phyllantha are also found in
Australia, but the latter not for New Zealand.
Schofield (1974) listed Plagiothecium denticulatum (Hedw.) B.S.G., but most New Zealand and
all Australian collections have proved to be P.
novae-zealandiae Broth. (Ireland, 1992). Similarly Mnium rostratum Schrad. in Australasia
proved to be Plagiomnium novae-zealandiae
(Col.) T. Kop. (Koponen, 1981).
New Zealand bipolar mosses numbered 83
species (Schofield, 1974), but for Australia the
number is about 66, of which the following have
not been reported for New Zealand: Bartramia
halleriana, Bryum alpinum, B. creberrianum, Calliergon sarmentosum, Drepanocladus sendtneri,
Meesia triquetra, Racomitrium heterostichum,
R. sudeticum, Scorpidium scorpioides, Ulota
phyllantha.
An interesting distribution is that of Climacium dendroides which so far has been
reported only from the Dargo High Plains in
Victoria. It is not common there, but the two
colonies found were in seepage areas and quite
large. The only other southern hemisphere occurrence is in New Zealand. The recent discovery
of Scorpidium scorpioides in Tasmania (Dalton
et al., 1991) was unexpected as it does not occur in New Zealand. The species was common
in continuous colonies around the perimeters
of small, shallow alkaline ponds. It may prove
to be more common than present reports suggest. It was not known from New Zealand when
Schofield (1974) reported on the New Zealand
bipolar mosses.
Bartramia halleriana, Conostonum
tetragonum, Polytrichastum alpinum are absent
from Denmark and Estonia because they are
mountain species. Bryum creberrimum, which
prefers basic substrates is absent from Estonia.
Pterygoneurum ovatum is absent from Estonia,
Finland and Iceland. Bryum sauteri is present
in Norway and Iceland only, while Grimmia
laevigata is reported only from Sweden and
Denmark.
Species widespread in Australia
Widespread species are predominantly in the
families Bryaceae or Pottiaceae and occur in
open or disturbed areas. They include: Aloina
aloides var. ambigua, Barbula hornschuchiana,
Bryum creberrimum, B. dichotomum, B. torquescens, Campylopus pyriformis, Grimmia pulvinata,
Hedwigidium integrifolia, Pottia davalliana,
Pterygoneurum ovatum, Tortula papillosa and
Weissia controversa. Of them Grimmia pulvinata and Hedwigidium integrifolia prefer drier
granitic rocks at medium altitudes, generally
in woodlands.
Physcomitrella patens occurs in moist areas
of southern Australia and in northern Europe
it occurs on mud or silt, but is absent from
Iceland.
Cold climate and high altitude species
Australian regions where the climate would
resemble that of northern Europe are the Australian Alps (New South Wales, Victoria), the
highland areas of these two states plus Tasmania and Macquarie Island.
Tasmania. Tasmania has a cooler and moister
climate than most of mainland Australia. This
is reflected in the high number of species restricted to that state only in Australia, but also
common with New Zealand. Species known
only from Tasmania and occurring in northern
Europe are: Buxbaumia aphylla, Dicranella
schreberiana, Fissidens adianthoides, Grimmia
funalis, Kiaeria starkei, Polytrichum formosum,
Racomitrium heterostichum, Rhytidiadelphus
triquetrus, Scorpidium scorpidioides, Sphagnum
compactum.
Macquarie Island. Ulota phyllantha is the only
species restricted to Macquarie Island where
102 Folia Cryptog. Estonica
it forms compact cushions on rocks. It was
reported (Seppelt, 1978) from a raised beach
terrace and apparently it is salt tolerant.
Southeast Australian species that extend to
Macquarie Island are: Amblystegium serpens,
Brachythecium rutabulum, B. salebrosum, Desmatodon heimii, Drepanocladus aduncus, Kindbergia praelonga, Orthodontium lineare, Sanionia
uncinata, Schistidium apocarpum, .
Widespread and common southern Australian species that are absent from humid tropical areas but extend to Macquarie Island are
Campylopus introflexus, Ceratodon purpureus,
Grimmia laevigata and Hypnum cupressiforme.
This latter species is not common in Western
Australia. Another common and widespread
Australian species found on the Island is Polytrichum juniperinum.
High altitude species. The highest proportion
of species which also occur in northern Europe
reach their best development in the Australian
Highlands and the alps, mostly preferring the
moister areas. McVean and Weber were the first
to collect widely in the Kosciuszko National Park.
J.H. Willis included mosses amongst his plant
studies on the Bogong High Plains of Victoria.
In my continuing studies in the High Plains of
Victoria several species have been found that
are more common than previously reported
(e.g. Meesia triquetra reported as rare by Scott
& Stone, 1976). In Tasmania many of these species would occur at lower altitudes.
High altitude species that are confined to
swampy or moist areas include: Brachythecium
rivulare, Bryum algovicum, B. muhlenbeckii, B.
pseudotriquetrum, B. subapiculatum, B. tenuisetum, Calliergon sarmentosum, Calliergonella
cuspidata, Campylium polygamum, Climacium
dendroides, Drepanocladus aduncus, D. sendtneri, Leptodictyum riparium, Meesia triquetra,
Pohlia bulbifera, P. cruda, Sanionia uncinata,
Scorpidium scorpioides, Warnstorfia exannulata, W. fluitans.
Pohlia nutans and Pohlia wahlenbergii can
also occur at mid-elevations where they can
withstand slightly drier conditions.
Mosses found predominantly on rocks or
rock-faces include: Distichium capillaceum,
Fissidens adianthoides, Grimmia trichophylla,
Orthotrichum rupestre, Racomitrium heteros-
tichum, R. sudeticum, Schistidium rivulare,
Tortula ruralis.
Racomitrium lanuginosum is common in
northern Europe. In Australia it is represented
by the ssp. pruinosum Wils. It is scattered and
moderately common in the sub-alpine regions,
but rarely forming large continuous colonies.
Sanionia uncinata is found in moister areas on
the bases of treelets and shrub stems, on logs
and along water course banks.
Other species
Some species barely manage to reach the European study area, possibly because they prefer
the milder climates. Two species are found only
in southern Sweden where they are not very
common – Acaulon mediterraneum and Pottia
starckeana. Acaulon mediterraneum occurs on
lawns and river flats while the latter species is
possibly introduced to Australia (Catcheside,
1980), being common in southern Australia on
disturbed soil and in urban habitats. Bryum
torquescens is rare in Denmark and southern
Sweden while reasonably common in Australia
on coastal sand dunes. Pottia davalliana which
occurs in disturbed situations in southern
Australia is not common in northern Europe,
especially Estonia.
Aloina rigida is restricted to South Australia
where it occurs on calcareous soils in dry disturbed habitats. Catcheside (1980) suspects that
it may be introduced. In northern Europe it is
found in Denmark, Estonia, Sweden and Norway, possibly in the warmer regions. An exclusively limestone moss in Australia, as in Europe,
is Orthotrichum cupulatum which is restricted to
mid-elevations in southeast Australia. Encalypta
vulgaris occurs in drier southern Australia and
is found on the ground, rocks and rock crevices,
often also on limestone.
Bryum alpinum is known from one Victorian
mountain locality from rocks (type unknown)
in a creek while in Europe it prefers wet acid
rocks. B. dichotomum occurs on clayey or silty
soils, rocks, moist and sheltered areas and on
disturbed soils in towns. Hedwigia ciliata prefers
drier acid rocks and is common at mid-elevations in paddocks and Eucalyptus woodlands,
while in Europe it is found on rocks in paddocks.
Several species that are found in both northern Europe and Australia occur in sub-tropical
103
Queensland. Didymodon vinealis and Trichostomum brachydontium, are found in the drier
interior from southern Queensland to Western
Australia. Anoectangium aestivum has been
reported from eastern Victoria, Southeast New
South Wales and from tropical coastal Queensland. Another species that reaches Queensland,
and is reasonably common in moister Southeast Australia, is Neckera pennata. Campylopus
flexuosus is restricted to northeast Queensland
where it occurs above 760 m, but is more common on the higher mountains.
COMPARISONS OF LIVERWORTS
Fewer species (Table 1), and a smaller proportion of liverworts are common between the two
study areas (Table 2). Of the 436 species found
in Australia 26 genera extend into the northern
European study area, with only 10 genera with
14 species common to the two regions.
Cosmopolitan species are not common in
Australia with only 2 species considered cosmopolitan. Lunularia cruciata is common along
water courses, generally in disturbed, manmade habitats, but also in various vegetation
types that often dry out. Ricciocarpus natans
is common on still lakes, especially those with
bird populations (Scott, 1985). Schuster (1992)
considers Pallavicinia lyellii to be sub-cosmopolitan being confined to temperate regions. In
Australia it occurs in moister forested areas at
lower elevations in Tasmania and Victoria.
Scott (1985) considered Marchantia polymorpha to be recently introduced, but CANB
collections annotated by Bischler-Causse (1989)
indicate that this species occurs in natural vegetation, as well as man made habitats between
600–1600 m in moister Southeast Australia.
Six species can be considered to be bipolar
disjunctive and are confined to Southeast Australia and Tasmania. Calopogeia sphagnicola is
confined to Tasmania. Cephaloziella arctica is
found in a wide range of vegetation types, generally on burnt logs or soil, between 80–1400
m. Lophocolea bidentata ranges from Eucalyptus
forests to sub-alpine heath, from near sea-level
to 1520 m, and extends to grasslands on Macquarie Island. Lophocolea minor prefers moist,
often calcareous rocks, in temperate forest and
on semi-exposed ground at low to mid-altitudes.
Metzgeria furcata is found in various forest
types and commonly on road cuttings between
20–1600 m, and extends to grasslands on Macquarie Island. Riccia sorocarpa prefers soil and
rocks in drier areas at lower elevations.
Two species are restricted to warmer,
semi- to arid regions at lower altitudes. Riccia
cavernosa occurs on clay or sand. Riccia bifurca
is found generally in moister regions, often in
towns while in northern Europe it occurs also
at low to mid-altitudes.
Diplophyllum obtusifolium is confined to the
sub-alpine areas where it occurs on rock ledges.
Metzgeria conjugata is found in New South Wales
and Tasmania, but no further data were available.
LIST OF BRYOPHYTES OCCURRING IN
AUSTRALIA AND NORTHERN EUROPE
# bipolar species
introduced species
! cosmopolitan species
Mosses
ACAULON MEDITERRANEUM Limpr.
ALOINA ALOIDES (K.F. Schultz) Kindb.
ALOINA RIGIDA (Hedw.) Limpr.
# AMBLYSTEGIUM SERPENS (Hedw.) B.S.G.
# AMBLYSTEGIUM VARIUM (Hedw.) Lindb.
ANOECTANGIUM AESTIVUM (Hedw.) Mitt.
# AULACOMNIUM PALUSTRE (Hedw.) Schwaegr.
BARBULA HORNSCHUCHIANA K.F. Schultz
# BARBULA UNGUICULATA Hedw.
# BARTRAMIA HALLERIANA Hedw.
# BRACHYTHECIUM ALBICANS (Hedw.) B.S.G.
BRACHYTHECIUM MILDEANUM (Schimp.) Schimp.
ex Milde
# BRACHYTHECIUM PLUMOSUM (Hedw.) B.S.G.
BRACHYTHECIUM RIVULARE B.S.G.
# BRACHYTHECIUM RUTABULUM (Hedw.) B.S.G.
# BRACHYTHECIUM SALEBROSUM (Web. & Mohr)
B.S.G.
# BRYOERYTHROPHYLLUM RECURVIROSTRUM (Hedw.)
Chen
# BRYUM ALGOVICUM Sendtn. ex C. Muell.
# BRYUM ALPINUM Huds. ex With.
! BRYUM ARGENTEUM Hedw.
! BRYUM CAESPITICIUM Hedw.
! BRYUM CAPILLARE Hedw.
# BRYUM CREBERRIMUM Tayl.
104 Folia Cryptog. Estonica
#
#
#
#
#
#
#
#
#
#
#
#
#
#
#
#
#
#
#
!
#
#
#
#
#
#
#
#
#
!
#
#
#
#
#
BRYUM DICHOTOMUM Hedw.
BRYUM INTERMEDIUM (Brid.) Bland.
BRYUM MUHLENBECKII B.S.G.
BRYUM PSEUDOTRIQUETRUM (Hedw.) Gaertn.
BRYUM RADICULOSUM Brid.
BRYUM RUBENS Mitt.
BRYUM SAUTERI B.S.G.
BRYUM SUBAPICULATUM Hampe
BRYUM TENUISETUM Limpr.
BRYUM TORQUESCENS BB. & S.
BUXBAUMIA APHYLLA Hedw.
CALLIERGON SARMENTOSUM (Wahlenb.) Kindb.
CALLIERGONELLA CUSPIDATA (Hedw.) Loeske
CAMPYLIUM POLYGAMUM (B.S.G.) J. Lange & C.
Jens.
CAMPYLOPUS FLEXUOSUS (Hedw.) Brid.
CAMPYLOPUS INTROFLEXUS (Hedw.) Brid.
CAMPYLOPUS PYRIFORMIS (K.F. Schultz) Brid.
CERATODON PURPUREUS (Hedw.) Brid.
CLIMACIUM DENDROIDES Web. & Mohr
CONOSTOMUM TETRAGONUM (Hedw.) Lindb.
DESMATODON HEIMII (Hedw.) Mitt.
DICRANELLA SCHREBERIANA (Hedw.) Dix.
DICRANUM SCOPARIUM Hedw.
DIDYMODON VINEALIS (Brid.) Zander
DISTICHIUM CAPILLACEUM (Hedw.) B.S.G.
DREPANOCLADUS ADUNCUS (Hedw.) Warnst.
DREPANOCLADUS SENDTNERI (Schimp.) Warnst.
ENCALYPTA VULGARIS Hedw.
EURHYNCHIUM SPECIOSUM (Brid.) Jur.
FISSIDENS ADIANTHOIDES Hedw.
FISSIDENS BRYOIDES Hedw.
FUNARIA HYGROMETRICA Hedw.
FUNARIA MUEHLENBERGII Turn.
GRIMMIA LAEVIGATA (Brid.) Brid.
GRIMMIA PULVINATA (Hedw.) Sm.
GRIMMIA TRICHOPHYLLA Grev.
GYMNOSTOMUM AERUGINOSUM Sm.
HAPLOHYMENIUM PSEUDOTRISTE (C. Muell.) Broth.
HEDWIGIA CILIATA (Hedw.) P. Beauv.
HEDWIGIDIUM INTEGRIFOLIUM (P. Beauv.) Dix.
HYMENOSTYLIUM RECURVIROSTRUM (Hedw.) Dix.
HYPNUM CUPRESSIFORME Hedw.
KIAERIA STARKEI (Web. & Mohr) I. Hag.
KINDBERGIA PRAELONGA (Hedw.) Ochyra
LEPTOBRYUM PYRIFORME (Hedw.) Wils.
LEPTODICTYUM RIPARIUM (Hedw.) Warnst.
MEESIA TRIQUETRA (Richter) Ångstr.
NECKERA PENNATA Hedw.
ORTHODONTIUM LINEARE Schwaegr.
ORTHOTRICHUM CUPULATUM Brid.
O R T H O T R I C H U M R U P E S T R E Schleich. ex
Schwaegr.
# PHASCUM CUSPIDATUM Hedw.
PHYSCOMITRELLA PATENS (Hedw.) B. & S.
PHYSCOMITRIUM PYRIFORME (Hedw.) Brid.
POHLIA BULBIFERA (Warnst.) Warnst.
#POHLIA CRUDA (Hedw.) Lindb.
# POHLIA NUTANS (Hedw.) Lindb.
# POHLIA WAHLENBERGII (Web. & Mohr) Andr.
# POLYTRICHASTRUM ALPINUM (Hedw.) G.L. Sm.
# POLYTRICHUM COMMUNE Hedw.
# POLYTRICHUM FORMOSUM Hedw.
! POLYTRICHUM JUNIPERINUM Hedw.
POLYTRICHUM PILIFERUM Hedw.
POTTIA DAVALLIANA (Sm.) C. Jens.
POTTIA STARCKEANA (Hedw.) C. Muell.
# POTTIA TRUNCATA (Hedw.) B. & S.
# PSEUDOSCLEROPODIUM PURUM (Hedw.) Fleisch.
# PTERYGONEURUM OVATUM (Hedw.) Dix.
# RACOMITRIUM HETEROSTICHUM (Hedw.) Brid.
# RACOMITRIUM LANUGINOSUM (Hedw.) Brid.
# RACOMITRIUM SUDETICUM (Funck) B. & S.
RHYTIDIADELPHUS TRIQUETRUS (Hedw.) Warnst.
# SANIONIA UNCINATA (Hedw.) Loeske
SCHISTIDIUM APOCARPUM (Hedw.) B. & S.
# SCHISTIDIUM RIVULARE (Brid.) Podp.
# SCORPIDIUM SCORPIOIDES (Hedw.) Limpr.
SPHAGNUM COMPACTUM Lam. & DC.
SPHAGNUM CUSPIDATUM Ehrh. ex Hoffm.
SPHAGNUM PALUSTRE L.
# TORTULA LAEVIPILA (Brid.) Schwaegr.
# TORTULA MURALIS Hedw.
# TORTULA PAPILLOSA Wils.
TORTULA RURALIS (Hedw.) Gaertn., Meyer &
Schreb.
T RICHOSTOMUM BRACHYDONTIUM Bruch
# ULOTA PHYLLANTHA Brid.
WARNSTORFIA EXANNULATA (B.S.G.) Loeske
# WARNSTORFIA FLUITANS (Hedw.) Loeske
WEISSIA CONTROVERSA Hedw.
WEISSIA RUTILANS (Hedw.) Lindb.
Liverworts
# CALYPOGEIA SPHAGNICOLA (Arn. & Perss.) Warnst.
& Loeske
# CEPHALOZIELLA ARCTICA Bryhn & Douin
DIPLOPHYLLUM OBTUSIFOLIUM (Hook.) Dum.
# LOPHOCOLEA BIDENTATA (L.) Dum.
# LOPHOCOLEA MINOR Nees
! LUNULARIA CRUCIATA (L.) Lindb.
MARCHANTIA POLYMORPHA L.
METZGERIA CONJUGATA Lindb.
# METZGERIA FURCATA (L.) Dum.
PALLAVICINIA LYELLII (Hook.) Carruth.
RICCIA BIFURCA Hoffm.
105
RICCIA CAVERNOSA Hoffm.
# RICCIA SOROCARPA Bisch.
! RICCIOCARPUS NATANS (L.) Corda
ACKNOWLEDGEMENTS
This study resulted from the kind invitation to
contribute to the anniversary volume dedicated
to Prof. Hans Trass. I had the good fortune to
enjoy his company, on several visits to Estonia
and discuss our mutual interests in lichens.
Therefore I feel honoured to be able to pay my
respects to his scholarship.
I am grateful to Drs E. Urmi (Zürich), J. Enroth
(Helsinki) and A. Frisvoll (Trondheim) for assistance with publications and data. Sincerest
thanks are also extended to Prof. W. Schofield
(Vancouver), Dr H.P. Ramsay (Sydney) and
M.Sc. N. Ingerpuu for comments suggesting improvements to the work. Dr R. Ochyra
(KRAM) assisted with further information, while
J. Johnston provided helpful editing assistance
and suggestion to improve the content.
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106 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 107–112 (1998)
Lecideoid lichens from Estonia – an annotated checklist
Ave Suija
Institute of Botany and Ecology, University of Tartu, 38 Lai St., EE2400 Tartu, Estonia
Abstract: 53 species, previously placed in the genus Lecidea are recognized in Estonia. Six of them – Cecidonia umbonella
(Nyl.) Triebel & Rambold, Lecidea plana (J. Lahm) Nyl., Lecidella scabra (Taylor) Hertel & Leuckert, Placynthiella dasaea (Stirt.)
Tønsberg, Placynthiella oligotropha (J.R. Laundon) Coppins & P. James and Trapeliopsis pseudogranulosa Coppins & P. James are
new to the country.
Kokkuvõte: A. Suija. Lecidea-taolised kooriksamblikud Eestis.
Eesti lihhenoflooras on 53 varem perekonda Lecidea kuulunud liiki. Neist kuut – Cecidonia umbonella (Nyl.) Triebel & Rambold,
Lecidea plana (J. Lahm) Nyl., Lecidella scabra (Taylor) Hertel & Leuckert, Placynthiella dasaea (Stirt.) Tønsberg, Placynthiella oligotropha
(J.R. Laundon) Coppins & P. James and Trapeliopsis pseudogranulosa Coppins & P. James märgitakse esmakordselt.
INTRODUCTION
MATERIALS AND METHODS
Since the last and the most complete checklist of
Estonian lichens (Trass, 1970) until now, many
important changes in lichen systematics have
been taken place. For example, the catalogue by
Trass contained 45 species in the genus Lecidea,
while only four of them belong to the genus today. All other species have been placed into 15
different genera. The term “lecideoid lichens”
was proposed by H. Hertel (1984) to unite all
taxa previously treated in the genus Lecidea.
53 lecideoid lichen species representing 20
genera plus eight species with uncertain position
are listed in the present paper. 43 species occur
in the contemporary lichen flora of Estonia; six
(Amygdalaria panaeola, Adelolecia kolaensis,
Fuscidea cyathoides, “Lecidea” hypopta, “L.”
sphaerella, Trapelia coarctata) are considered
to be extinct (records only from before 1950).
Fuscidea lygaea, “Lecidea” albofuscescens,
Lecidea sarcogynoides and Pyrrhospora cinnabarina have been included in the checklist
because of information in old literature (Bruttan,
1870; Mereschkowski, 1913; Räsänen, 1931;
Trass, 1970), whereas there are no herbarium
specimens. Fuscidea kochiana, “Lecidea” fuscescens, “L.” fuliginosa, “L.” olivascens and Lecidella
anomaloides ought to be excluded from the
checklist (Trass, 1970), as they appeared to be
misidentifications. Six species – Cecidonia umbonella, Lecidea plana, Lecidella scabra, Placynthiella dasaea, P. oligotropha and Trapeliopsis
pseudogranulosa are new for Estonia.
This study was carried out in 1995–1997 and
was based on the herbarium collections, kept
in the University of Helsinki (H), International
Center for Environmental Biology (ICEB) in
Tallinn, Ecological Institute (IE) in Tallinn,
University of Riga (RIG), Estonian Nature Museum (TAL), Tallinn Botanic Garden (TBA), the
Institute of Botany and Ecology of the University
of Tartu (TU) and Uppsala University (UPS). The
references in literaure have also been taken into
account: Bruttan (1870), Ekman et al. (1991),
Mereschkowsky (1913), Moberg (1992), Räsänen
(1931) and Trass (1967, 1970).
The morphological characters of taxa were
examined using a stereo lens MBS-9, the anatomical characters using a light microscope
MBI-3. Cross sections of fruitbodies were made
by hand and then mounted in water, 10% KOH
or Lugol’s solution. The “spot tests” were made
on the thallus or medulla using 10% KOH (K),
sodium hypochlorite (C), p-phenylendiamine
in ethanol (Pd) or Lugol’s solution (I). Chemical analyses of secondary products of lichens
were carried out according to standardized TLC
methods described by Culberson & Kristinsson
(1970) and Culberson (1972).
The following keys were used to identify
the specimens: Foucard (1990), Gowan & Ahti
(1993), Hertel (1995), Magnusson (1952b),
Printzen (1995), Purvis et al. (1992), Tønsberg
(1992), Vainio (1934) and Wirth (1995).
108 Folia Cryptog. Estonica
CHECKLIST OF TAXA
Genera and species are listed alphabetically.
Nomenclature follows Santesson (1993), but
recent changes are also taken into account.
Species, the systematical position of which is
uncertain, are marked with quotation marks (“
”). Synonymes, which have been used in Estonian lichenological literature, are also included.
New taxa for Estonia are given in bold, doubtful
species are marked with a question mark (?),
exctinct species with an asterisk (*).
* ADELOLECIA KOLAENSIS (Nyl.) Hertel & Rambold
(syn. Lecidea conferenda Nyl.) – on siliceous rocks. Probably extinct in Estonia,
only one collection known from the first part
of century: Harjumaa Co., Kunda (58°29’N,
26°32’E), in the seashore, V. Räsänen, 14
June 1927 (H).
* AMYGDALARIA PANAEOLA (Ach.) Hertel & Brodo
(syn. Lecidea panaeola Ach.) – on siliceous
rocks. Probably extinct in Estonia, only
one collection known from the last century:
Saaremaa Co., Saaremaa Is., A. Bruttan,
No.141 (RIG).
BIATORA CHRYSANTHA (Zahlbr.) Printzen (syn. Lecidea epixanthoidiza auct. non Nyl.) – on
mosses. Very rare, only one record known:
Läänemaa Co., at the ornithological field station at Puhtu (58°34’N, 23°35’E) (S) (Ekman
et al., 1991).
B IATORA EF FLORESCENS (Hedl.) Erichsen [syn.
Lecidea epixanthoidiza Nyl., L. efflorescens
(Hedl.) Erichsen] – on the bark of coniferous
trees. Rather rare: known from eight localities all over the country.
B IATORA HELVOLA Körb. [syn. Lecidea helvola
(Körb.) Th. Fr.] – on the bark of coniferous
trees, rarely on deciduous trees. Rather
common: about 20 localities known in different parts of Estonia.
BIATORA OCELLIFORMIS (Nyl.) Arnold [syn. Lecidea
atroviridis (Arnold) Th. Fr.] – on the bark
of deciduous and coniferous trees. Rare:
known from four localities. 1. Saaremaa
Co., Saaremaa Is., Viidumäe Nature Reserve,
sq. no.227 (58°16’N, 21°58’E), T. Randlane,
4541, 26 Aug. 1976 (TU); 2. Tartumaa Co.,
Vasula, Vasula forest (58°28’N, 26°40’E),
E. Parmasto, 4539, 21 Nov. 1948 (TU); 3.
Tartumaa Co., Tähtvere distr., sq. no.53
(58°25’N, 26°35’E), P. Pott, 4540, 10 July
1995 (TU); 4. Põlvamaa Co., near the Ahja
River, Otten’s mill (58°07’N, 27°03’E), H.
Trass, 4542, 13 July 1990 (TU).
BIATORA VERNALIS (L.) Fr. [syn. Lecidea vernalis (L.)
Ach.] – on the ground and on mosses. Rather
rare: known from six scattered localities.
CARBONEA VITELLINARIA (Nyl.) Hertel [syn. Lecidea
vittelinaria (Flörke) Körb.] – lichenicolous
on the thallus of Candellariella sp. Rare:
known from five localities in western and
northern Estonia 1. Saaremaa Co., Harilaid
(58°29’N, 21°51’E), H. Trass, 4479, 21 June
1959 (TU); 2. Läänemaa Co., Kaseküla alvar
(58°38’N, 23°34’E), H. Trass, 4481, 4482, 10
Aug. 1991 (TU); 3. Harjumaa Co., Rammu
Is. (59°34’N, 25°14’E), E. Nilson, July 1993
(IE); 4. Harjumaa Co., Rammu Is., T. Piin,
No. 33(91), 1991 (TBA); 5. Harjumaa Co.,
Pedassaar Is. (59°31’N, 25°23’E), T. Piin,
No.183(91), 1991 (TBA); 6. Harjumaa. Co.,
Tallinn, Nõmme (59°23’N, 24°42’E), J. Ruubel No.116, June 1931 (TAL).
CECIDONIA UMBONELLA (Nyl.) Triebel & Rambold
– lichenicolous on the thallus of Lecidea
lapicida var. lapicida. Very rare, known
from one locality: Saaremaa Co., Harilaid
islet (58°29’N, 25°51’E), H. Trass, 4772, 22
June 1959 (TU).
FARNOLDIA JURANA (Schaer.) Hertel (syn. Lecidea
albosuffusa Th. Fr., L. jurana Schaer.) – on
calcareous substrata, particulary limestone.
Rather rare: known from seven localities in
coastal areas of northern and western Estonia.
FUSCIDEA ARBORICOLA Coppins & Tønsberg – on
the bark of deciduous and coniferous trees.
Rare, but possibly overlooked: known from
three localities. 1. Ida-Virumaa Co., Soldina
– Auvere (59°20’N, 28°00’E), T. Piin, 4587, 9
Aug. 1961 (TU); 2. Viljandimaa Co., Olustvere (58°30’N, 25°30’E), T. Piin, 4588, 11
Aug. 1963 (TU); 3. Raplamaa Co., Märjamaa
distr., near Paluküla, sq. no.162, (58°52’N,
24°22’E), A. L. Sõmermaa, 4787, 18 Aug.
1967 (TU).
* FUSCIDEA CYATHOIDES (Ach.) V. Wirth & Vêzda
[syn. Lecidea cyathoides (Ach.) Ach.] – on
siliceous rocks. Probably extinct in Estonia,
only one collection known from the last century: Saaremaa Co., A. Bruttan (RIG, TU).
109
? FUSCIDEA LYGAEA (Ach.) V. Wirth & Vêzda (syn.
Lecidea lygaea Ach.) – Mereschkowski (1913)
observed in Saaremaa, but the herbarium
specimen has not been preserved.
LECIDEA CONFLUENS (Weber) Ach. – on siliceous
rocks. Rare: known from four localities
in western Estonian islands. 1. Hiiumaa
Co., Kassari, Sääre tirp (58°46’N, 22°48’E),
T. Randlane, 4570 17 June 1984 (TU); 2.
Saaremaa Co., Saaremaa Is., Viidumäe Nature Reserve, Viidumägi (58°18’N, 22°25’E),
V. Räsänen, 4569, 30 July 1929 (TU, H);
3. Saaremaa Co., Saaremaa Is., Randvere
(58°18’N, 22°24’E) H. Trass, 4755, 13 Aug.
1964 (TU); 4. Saaremaa Co., A. Bruttan
No.141 (RIG).
This species has two chemotypes (Hertel
1995) – chemotype I (confluentic and stictic acid) and chemotype II (confluentic acid).
Secondary chemistry of all above mentioned
specimens was analysed and only chemotype II was detected.
LECIDEA FUSCOATRA (L.) Ach. – on siliceous rocks.
Rather common: more than 20 localities in
northern and western Estonia known.
LECIDEA LAPICIDA (Ach.) Ach. – both varieties – var.
lapicida and var. pantherina are found on
siliceous rocks.
– var. LAPICIDA. Rather rare: known from seven
localities in western and northern Estonian
islands.
– var. PANTHERINA Ach. Rather common: known
from 20 localities in western and northern
Estonia.
L ECIDEA PLANA (J. Lahm) Nyl. – on siliceous
rocks. Rare: known from three localities. 1.
Hiiumaa Co., Tahkuna peninsula, H. Trass,
4745, 4748, 4751, 4 July 1959 (TU); 2. Põlvamaa Co., Värska (57°58’N, 27°38’E), Soe,
2248, 9 Sept. 1965 (TU); 3. Põlvamaa Co.,
Räpina, Pikaliiva sands, H. Trass, 4744, 8
July 1958 (TU).
? LECIDEA SARCOGYNOIDES Körb. – on siliceous
rocks. Bruttan (1870) has observed it in
Saaremaa, but unfortunately the collection
is not preserved.
LECIDEA TESSELLATA Flörke var. TESSELLATA – on
siliceous rocks. Rather rare: known from
seven localities mainly in coastal areas of
western and northern Estonia.
According to Hertel (1995), the species has
two chemotypes – chemotype I (confluentic
acid) and chemotype II (confluentic and
norstictic acid). Three specimens were
analysed and both chemotypes were detected. 1. Harjumaa Co., Tallinn, Kakumägi
(59°27’N, 24°35’E), K. Mereschkowski, 4565,
1911 (TU) (chemotype I); 2. Saaremaa Co.,
Vahase islet (58°09’N, 22°29’E), H. Lippmaa,
4567, 24 Aug. 1937 (TU) (chemotype II); 3.
Läänemaa Co., Lihula (58°41’N, 23°50’E), V.
Räsänen, 4566, 3 Aug. 1927 (TU) (chemotype I).
? “LECIDEA” ALBOFUSCESCENS Nyl. Trass (1967)
has referred four localities for this species.
Two specimens (Tartumaa Co., Meeksi and
Harjumaa Co., Tallinn-Mustamäe) appeared
to be misidentifications and the other two
(Saaremaa Co., Randvere and Läänemaa
Co., Tauksi Islet) samples are missing.
“LECIDEA” ALBOHYALINA (Nyl.) Th. Fr. [syn. Biatora
albohyalina (Nyl.) Bagl. & Car.] – on the
bark of deciduous trees. Very rare, found
only once: Läänemaa Co., near the Puhtu
ornithological station (58°34’N, 23°33’E), R.
Sundin No. 226a, 11–12 June 1989 (S) (Ekman et al. 1991).
“LECIDEA” BOTRYOSA (Fr.) Th. Fr. (syn. Biatora botryosa Fr.) – on decaying wood. Rare: known
from two localities in southern Estonia. 1.
Jõgevamaa Co., Puurmani, Põltsamaa water-meadow (58°34’N, 26°18’E), H. Trass,
4316, 4317, 4318, 9 June 1958 (TU); 2.
Viljandimaa Co., Tipu (58°21’N, 21°60’E),
L. Martin, Aug. 1995 (ICEB).
“L ECIDEA ” ERYTHROPHAEA Flörke [syn. Lecidea
tenebricosa (Ach.) Nyl.] – on the bark of deciduous trees and wood. Rare: known from
three localities in northern and western Estonia. 1. Harjumaa Co., Tallinn, Mustamägi
(59°24’N, 24°38’E), E. Parmasto, 4473, 20
Apr.1947 (TU); 2. locality unknown, A.
Bruttan, 4319, (TU), 3. Läänemaa Co.,
near Puhtu ornithological station (58°34’N,
23°33’E), 11–12 June 1989 (S) (Ekman et
al 1991); 4. Läänemaa Co., Kõinastu Islet
(58°38’N, 23°02’E), 14 July 1989 (S) (Ekman
et al., 1991).
* “LECIDEA” HYPOPTA Ach. [syn. Lecanora hypopta
(Ach.) Vainio] – on wood. Probably extinct
in Estonia, mentioned twice before 1950:
110 Folia Cryptog. Estonica
1. Locality unknown, J. Seim, 4723 (TU)
2. Harjumaa Co., Kunda, V. Räsänen
(Räsänen, 1931).
“LECIDEA” NYLANDERI (Anzi) Th. Fr. – on the bark
of Pinus silvestris. Rare, but possibly
overlooked, known from three localities:
1. Raplamaa Co., Märjamaa distr., near
Paluküla, sq. no.162 (58°52’N, 24°22’E),
A. L. Sõmermaa, 4787, 18 Aug. 1967 (TU);
2. Saaremaa Co., Saaremaa Is., Odalätsi
(58°28’N, 22°08’E), T. Randlane, 4528, 12
July 1989 (TU); 3. Saaremaa Co., Saaremaa
Is., near Kuressaare (58°16’N, 22°28’E), A.
Bruttan (RIG).
* “LECIDEA” SPHAERELLA Hedl. [syn. Lecidea sylvana
(Körb.) Th. Fr.] – on the bark of decideous
trees. Probably extinct in Estonia, the only
collection dates from the end of the last century: Võrumaa Co., near the lake Pühajärve,
A. Bruttan 4320 (TU).
“LECIDEA” TURGIDULA Fr. – on the bark of Pinus silvestris, rarely on wood. Common: more than
40 localities all over the country known.
LECIDELLA ELAEOCHROMA (Ach.) M. Choisy – on
the bark of deciduous trees, especially on
smooth-barked trees (Alnus sp., Sorbus aucuparia etc.), rarely on conifers and wood.
Very common in all parts of Estonia.
LECIDELLA EUPHOREA (Flörke) Hertel [incl. L. achristotera (Nyl.) Hertel & Leuckert, L. laureri
(Hepp) Körb.] – on the bark of deciduous
trees, rarely on coniferous trees and wood.
Very common in all parts of Estonia.
LECIDELLA SCABRA (Taylor) Hertel & Leuckert – on
siliceous rock. Very rare: known from one
locality: Saaremaa Co., Harilaid (58°29’N,
21°51’E), E. Nilson, Sept. 1993 (IE).
The specimen has both soralia (greenish,
more or less discrete) and apothecia. The
anatomy of fruitbodies is typical for the
genus Lecidella (lax paraphyses, broad-ellipsoid spores etc.). This species is widespread in northern and central parts of
Europe (Tønsberg, 1992; Wirth, 1995 etc.).
LECIDELLA STIGMATEA (Ach.) Hertel & Leuckert – on
calcareous rocks, more rarely on siliceous
rocks, grows both on natural and anthropogenous substrata. Common: more than
30 localities known, mainly in northern and
western Estonia.
MIRIQUIDICA DEUSTA (Stenh.) Hertel & Rambold
(syn. Lecidea deustata Zahlbr.) – on siliceous rocks. Very rare, only one locality
known: Saaremaa Co., Saaremaa Is., Atla
alvar (58°18’N, 21°54’E), H. Trass, 4501, 5
Aug. 1991 (TU).
The specimen contains lobaric acid.
PLACYNTHIELLA DASAEA (Stirt.) Tønsberg – on decaying wood. Very rare, but possibly overlooked, known from two localities: 1. Hiiumaa Co., Hanikatsi Islet (58°47’N, 23°02’E),
T. Randlane, 4243, 9 June 1982 (TU); 2.
Saaremaa Co., Saaremaa Is., Odalätsi
(58°23’N, 22°09’E), T. Randlane, 4226, 12
July 1989 (TU).
PLACYNTHIELLA ICMALEA (Ach.) Coppins & P. James
– on decaying wood, on plant debris, on bare
ground. Very common: more than 50 localities known, mainly northern, western and
southern Estonia.
PLACYNTHIELLA OLIGOTROPHA (J.R. Laundon) Coppins & P. James – on humus-rich ground
(heathlands, cut-over areas, peat-bogs etc.).
Rather common: known from 15 localities
all over the country.
PLACYNTHIELLA ULIGINOSA (Schrad.) Coppins & P.
James – on decaying wood, on plant debris,
on humus-rich and sandy ground. Common:
known from over 20 localities in western,
northern and southern Estonia
PORPIDIA CINEREOATRA (Ach.) Hertel & Knoph (syn.
Lecidea cinereoatra Ach.) – on siliceous
rocks. Very rare: known from two localities.
1. Harjumaa Co., Pedassaar Is. (59°31’N,
25°23’E), T. Piin, 244(91), 1991 (TBA); 2.
Saaremaa Co., Saaremaa Is., A. Bruttan,
4612 (TU).
PORPIDIA CRUSTULATA (Ach.) Hertel & Knoph [syn.
Lecidea crustulata (Ach.) Spreng.] – on siliceous rocks, especially on pebbles. Common: known from 20 localities in northern,
western and southeastern Estonia.
PORPIDIA MACROCARPA (DC.) Hertel & A.J. Schwab
[syn. Lecidea macrocarpa (DC.) Steud., L. nigrocruenta Anzi] – on siliceous rocks. Rather
rare: known from nine scattered localities.
PORPIDIA TUBERCULOSA (Sm.) Hertel & Knoph (syn.
Lecidea sorediza Nyl.) – on siliceous rocks.
Rare: known from three localities: 1. Harjumaa Co., Rammu Is. (59°34’, 25°12’E), E.
Nilson, July 1993 (IE); 2. Saaremaa Co.,
Saaremaa Is. Viidumäe Natural Reserve,
111
Tornimägi, sq. no.182 (58°16’N, 22°08’E),
T. Randlane, 4605, 18 Aug. 1976 (TU); 3.
Saaremaa Co., Kõinastu Islet (58°38’N,
23°02’E), 15 June 1989 (LD) (Ekman et
al., 1991).
PSILOLECHIA LUCIDA (Ach.) M. Choisy [syn. Lecidea
lucida (Ach.) Ach.] – on rocks, on the roots,
on the bark of Pinus silvestris etc. Grows
preferably in shaded areas. Rare: known
from five localities in southern and eastern Estonia. 1. Ida-Virumaa Co., Kauksi
(58°59’N, 27°15’E), A. L. Sõmermaa, 4741,
9 Aug. 1966 (TU); 2. Tartumaa Co., Tartu
(58°23’N, 26°44’E), A. Bruttan (RIG); 3.
Tartumaa Co., Alam-Pedja Nature Reserve,
Laeva distr., Karussaare, sq. no. 182/7
(58°26’N, 26°16’E), P. Lõhmus, 4742, 5
Okt. 1996 (TU); 4. Valgamaa Co., Kääriku
(58°01’N, 26°24’E), H. Trass 4262 12 July
1964 (TU); 5. Viljandimaa Co., Tipu distr.,
sq. no. 27 (58°21’N, 25°06’E), P. Lõhmus,
4743, 28 July 1994 (TU).
? PYRRHOSPORA CINNABARINA (Sommerf.) M. Choisy
[syn. Lecidea cinnabarina Sommerf., Protoblastenia cinnabarina (Sommerf.) Räsänen].
Räsänen (1931) observed it in Lääne-Virumaa, Väike-Maarja (under the name Protoblastenia cinnabarina), but the herbarium
specimen has not been preserved.
PYRRHOSPORA QUERNEA (Dicks.) Körb. [syn. Lecidea
quernea (Dicks.) Ach.] – on the bark of deciduous trees, rarely on the bark of coniferous trees. Rather rare: known from nine
localities in western and central Estonia.
RAMBOLDIA INSIDIOSA (Th. Fr.) Hafellner (syn. Lecidea insidiosa Th. Fr.) – lichenicolous on
the thallus of Lecanora varia. Rather rare:
known from six localities mainly in western
and northern Estonia.
RIMULARIA FURVELLA (Nyl.) Hertel & Rambold – lichenicolous on various lichens on siliceous
rocks. Rare: known from four localities: 1.
Läänemaa Co., Kaseküla alvar (58°38’N,
23°34’E), H. Trass, 4729, 10 Aug. 1991 (TU);
2. Saaremaa Co., Saaremaa Is., Vilsandi Nature Reserve (58°23’N, 2150’E), T. Randlane,
4561, 21 June 1980 (TU); 3. Saaremaa Co.,
Saaremaa Is., Viidumäe Nature Reserve, on
the border of sq. no. 17 and 164 (58°18’N,
22°06’E), T. Randlane, 2240 11 Aug. 1976
(TU); 4. Saaremaa Co., Abruka Is. (58°09’N,
22°31’E), R. Moberg (UPS).
RIMULARIA INSULARIS (Nyl.) Rambold & Hertel [syn.
Lecidea insularis (Nyl.) Rambold & Hertel,
L. intumescens (Flörke) Nyl.] – lichenicolous
on the thallus of Lecanora rupicola. Rather
common: known from 12 localities mainly
in western and northern Estonia.
SCHAERERIA FUSCOCINEREA (Nyl.) Clauzade & Roux
[syn. Lecidea griseoatra (Hoffm.) Flot.,
Schaereria tenebrosa (Flot.) Hertel & Poelt]
– on siliceous rocks. Rather rare: known
from seven localities in western and northern Estonia.
* TRAPELIA COARCTATA (Sm.) M. Choisy [syn. Lecidea
coarctata (Turner) Nyl.] – on siliceous rocks.
Probably extinct in Estonia, collected only
once in the last century: Saaremaa Co.,
Saaremaa Is., A. Bruttan, 4261 (TU).
T RAPELIA PLACODIOIDES Coppins & P. James – on
brick. Very rare, known from one locality:
Läänemaa Co., Osmussaar Is. (59°18’N,
23°22’E), I. Jüriado, 4260, 29 July 1993
(TU).
T RAPELIOPSIS FLEXUOSA (Fr.) Coppins & P. James
[syn. Lecidea flexuosa (Fr.) Nyl.] – on decaying wood (stumps, old roofs etc.) and
on the ground. Common: more than 30 localities known in western, northern, central
and southwestern Estonia.
T RAPELIOPSIS GRANULOSA (Hoffm.) Lumbsch (syn.
Lecidea granulosa (Ehrh.) Ach.) – on humusrich ground, on plant-debris, on mosses.
Very common: more than 50 localities
known all over Estonia.
TRAPELIOPSIS PSEUDOGRANULOSA Coppins & P. James
– on peaty ground, on humid and shaded
areas. Very rare, known from one locality:
Tartumaa Co., Tähtvere, on the bank of a
drainage ditch (58°24’N, 23°22’E), A. Suija,
4573, 4574, 12 June 1996 (TU).
T REMOLECIA ATRATA (Ach.) Hertel – on siliceous
rocks, usually on stone pebbles. Rare:
known from four localities in northern and
western Estonia. 1. Harjumaa Co., Äksi Is.
(59°36’N, 25°06’E), P. Põldmaa, 4503, 27
Aug. 1960 (TU); 2. Harjumaa Co., Rammu
Is. (59°34’, 25°12’E), T. Piin, No.58(91),
1991 (TBA); 3. Harjumaa Co., Naissaar Is.
(58°34’N, 24°31’E), E. Nilson, Aug.1993 (IE);
4. Saaremaa Co., Saaremaa Is., Harilaid
112 Folia Cryptog. Estonica
(58°29’N, 21°51’E), E. Nilson, Sept.1993 (IE);
5. Saaremaa Co., Saaremaa Is., Harilaid, H.
Trass, 4492, 6 Aug. 1991 (TU); 6. Saaremaa
Co., Saaremaa Is., Harilaid, T. Randlane,
4493, 15 June 1984 (TU).
ACKNOWLEDGEMENTS
My sincere thanks are due to Ljudmilla Martin,
Roland Moberg, Eva Nilson, Taimi Piin, Alfons
Piterans, Tiina Randlane, Andres Saag, Marina
Temina and Orvo Vitikainen. Prof. Dr. Hannes
Hertel and Dr. Tør Tønsberg are thanked for
identifying some of the specimens, Dr. G. Rambold for reviewing the manuscript.
This study is part of the project “Composition and analysis of Estonian lichen flora” and
is financially supported by the Estonian Science
Foundation (Grant No. 1297).
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Arch. Naturk. Liv-, Ehst-, u. Kurl. 7 : 163–326.
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data for the identification of lichen products by a
standardized thin-layer chromatographic method.
J. Chromatogr. 72 : 113–125.
Culberson, C. F. & Kristinsson, H. 1970. A standardized method for the identification of lichen
products. J. Chromatogr. 46 : 85–93.
Ekman, S., Fröberg, L., Kärnefeldt, I., Sundin, R. &
Thor, G. 1991. New or interesting lichens from
Estonia. Folia Cryptog. Estonica 28: 1–32.
Foucard, T. 1990. Svensk skorplavs flora. Lund. 306
pp.
Gowan, S. P. & Ahti, T. 1993. Status of the lichen
genus Porpidia in Eastern Fennoscandia. Ann.
Bot. Fenn. 30 : 53–75.
Hertel, H. 1984. Über saxicole, lecideoide Flechten
der Subantarktis. Beih. Nova Hedwigia 79 :
400–492.
Hertel, H. 1995. Schlüssel für die Arten der Flechtenfamilie Lecideaceae in Europa. Bibl. Lichenol.
58 : 137–180.
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in Scandinavia and Finland. 2. Non-saxicolous
species. Svensk Bot. Tidskr. 46 (3–4) : 313–323
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Baltic provinces (in Russian). Kazan.
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from Estonia. Folia Cryptog. Estonica 29: 28–30.
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James, P. W., Moore, D. M. 1992. The Lichen
Flora of Great Britain and Ireland. Natural History
Museum Publications, London. 710 pp.
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163 pp.
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fungi of Sweden and Norway. SPT -förlaget.
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Trass, H. 1967. Analysis of the lichen-flora of Estonia
(in Russian). Doctoral dissertation. Tartu State
University.
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lichen-flora of Estonia (in Russian). Pap. Bot. 9:
5–233.
Tønsberg, T. 1992. The sorediate and isidiate, corticolous crustose lichens in Norway. Sommerfeltia
14 : 1–331.
Vainio, E. A. 1934. Lichenographa Fennica IV. Lecideales II. Acta Soc. Fauna Fl. Fenn. 57 (2) : 1–506.
Wirth, V. 1995. Flechtenflora. Ulmer. 666 pp.
Folia Cryptog. Estonica, Fasc. 32: 113–122 (1998)
Phylogenetic relationships of some cetrarioid species in British
Columbia with notes on Tuckermannopsis
Arne Thell
Department of Systematic Botany, Lund University, Östra Vallgatan 18–20, S-223 61 Lund, Sweden
Abstract: Phylogenetic relationships of seven cetrarioid lichens – Ahtiana sphaerosporella, Esslingeriana idahoensis, Kaernefeltia
merrillii, Tuckermannopsis americana, T. platyphylla, T. subalpina and Vulpicida canadensis – occurring in British Columbia are
analysed together with seven other cetrarioid species using parsimony analysis. The phylogenetic hypothesis based on ITS
sequences suggest that ascus characters cannot be used as characters to distinguish genera within cetrarioid lichens but
pycnoconidial shape is representative for larger groups. Tuckermannopsis as a polyphyletic genus is regarded as a key group
for a better phylogenetic understanding of cetrarioid lichens.
Kokkuvõte: A. Thell. Mõnede tsetrarioidsete liikide fülogeneetilised seosed Briti Kolumbias, rõhuasetusega perekonnale Tuckermannopsis.
Analüüsitakse seitsme Briti Kolumbias (Kanada) esineva tsetrarioidse samblikuliigi – Ahtiana sphaerosporella, Esslingeriana
idahoensis, Kaernefeltia merrillii, Tuckermannopsis americana, T. platyphylla, T. subalpina jaVulpicida canadensis – omavahelisi fülogeneetilisi seoseid, samuti seoseid seitsme muu tsetrarioidse liigiga, kasutades parsimoonset analüüsi. ITS sekventsidel põhineva
fülogeneetilise hüpoteesi kohaselt ei saa eoskoti tunnuseid kasutada tsetrarioidsete samblike perekondade eristamisel, kuid
püknokoniidide kuju on sobiv tunnus suuremate rühmade eristamiseks. Polüfüleetilist perekonda Tuckermannopsis võib käsitleda kui võtmerühma tsetrarioidsete samblike fülogeneesi paremaks mõistmiseks.
INTRODUCTION
The lichen flora of British Columbia, acknowledged for its richness, comprises to 35 out of approximately 135 cetrarioid species in the latest
world list (Goward et al., 1994; Randlane et al.,
1997). In autumn 1996, I had the opportunity to
collect fresh material of cetrarioid lichens in British Columbia. During a stay at the University of
British Columbia, I started molecular work on
the group. The internal transcribed spacer (ITS)
regions of the mycobiont’s nuclear ribosomal
DNA (rDNA) gene appeared to be particularly
suitable for comparing closely related species
and genera. This gene has already been used in a
study of populations of Platismatia s.l. and other
cetrarioid species (Thell et al., 1998; Thell &
Miao in prep.). Several sequences resulting from
these studies (Cetraria islandica, C. sepincola,
Flavocetraria cucullata, Platismatia glauca, Tuckermannopsis chlorophylla and Vulpicida pinastri)
were selected for a comparison with new sequences from seven boreal to subalpine species
from British Columbia (Ahtiana sphaerosporella,
Esslingeriana idahoensis, Kaernefeltia merrillii,
Tuckermannopsis americana, T. platyphylla, T.
subalpina and Vulpicida canadensis). As part
of a larger survey of the phylogeny of cetrarioid
lichens, this investigation focuses mainly on
the relationships of some species assigned to
Tuckermannopsis.
The genus Tuckermannopsis was introduced
by Gyelnik (1933), who distinguished it from
Nephromopsis by the absence of pseudocyphellae on the lower surface. The generic name was
not commonly used by lichenologists until it was
reinstated by Lai (1981), who combined several
taxa in Tuckermannopsis. Hale (in Egan, 1987),
Weber (in Egan, 1991), Kurokawa (1991) and
Kärnefelt et al. (1993) transferred additional taxa
to the genus. Several of these taxa have recently
been assigned to other genera, i. e. Ahtiana, Allocetraria, Kaernefeltia, Melanelia and Vulpicida
(Mattson & Lai, 1993; Thell, 1995, Thell et al.,
1995b, c; Thell & Goward, 1996). Currently, 11
taxa are assigned to Tuckermannopsis but still
today the genus appears heterogeneous.
Cetrarioid lichens have been divided into
three categories and may also include parmelioid
genera (Kärnefelt et al., 1992). Two of these
groups, centered around Cetraria s. str. and
114 Folia Cryptog. Estonica
Tuckermannopsis, included taxa almost exclusively with an origin in Cetraria. The third group
contains taxa with asci of the Melanelia type
which is present in the majority of the Parmeliaceae (Thell et al., 1995b).
MATERIAL AND METHODS
I. Selected material
The lichen samples used for the study and
cited below were collected in British Columbia
1995–96 and are stored together with DNA isolations in the herbaria LD, TDI (TerraGen Diversity
Inc.) and UBC.
Ahtiana sphaerosporella: Joffre Lake, Miao
& Taylor, DNA#AT83 (TDI#211). Esslingeriana
idahoensis: Clearwater, Goward TG-961354,
DNA#AT145 (UBC); Goward TG-961348,
DNA#AT146 (UBC). Kaernefeltia merrillii: Clearwater, Thell BC-9698, DNA#AT190; Whistler,
Thell & Veer BC-9670, DNA#AT71 (LD). Tuckermannopsis americana: Joffre Lake, Miao &
Taylor, DNA#AT82 (TDI#210); Wells Gray Park,
Goward TG-961350 (UBC). Tuckermannopsis
platyphylla: Whistler, Thell & Veer BC-9675,
DNA#AT75 (LD); Kamloops, Thell & Veer BC9643, DNA#AT43 (LD). Tuckermannopsis subalpina: Burke Mtn., Thell BC-9606, DNA#AT109
(LD). Vulpicida canadensis: Kamloops, Thell &
Veer BC-96250, DNA#AT36 (LD); Jackson Flats,
Miao, DNA#VM174 (TDI#146).
II. DNA preparation
The DNA was isolated following the procedure
of Thell et al. (1998). Fragments were amplified
with the primers ITS5 and ITS4 (White et al.,
1990) using AmpliTaq DNA Polymerase (5 U/
µl) according to the manufactorers protocol.
All samples were PCR-amplified and agarose
purified twice. A PCR was performed for 30
cycles with the following cycling conditions:
denaturation at 94°C for 1 min., annealing at
48°C for 1 min., and extension at 72°C for 45
seconds. The second PCR differed from the first
by an elevated annealing temperature of 53°C
(Thell et al. 1998).
Both complementary strands were sequenced using Dye Terminator Cycle Sequencing Ready Reaction Kit (Perkin Elmer) according
to the manufactorers instructions. ITS5, ITS4,
ITS3 and ITS2 were used as sequencing primers (White et al., 1990). Cycle sequencing was
carried out for 25 cycles with denaturation at
96°C for 10 seconds, annealing temperature
at 50°C for 5 seconds, and extension at 60°C
for 4 minutes. The DNA was cleaned through
CENTRI-SEP columns (Princeton Separations,
Inc.). Sequenced fragments were analysed using an ABI Prism 377 DNA automatic sequencer
(Perkin Elmer).
III. Alignment and phylogenetic analysis
The ITS sequences were aligned with
SeqApp/CAP 2, followed by a slight adjustment
by hand (Table 1) and transferred to PAUP 3.0
(Gilbert, 1993; Huang, 1992). The consensus
tree from 500 bootstrap replicates is presented.
This tree was obtained using the branch and
bound option (Fig. 1).
RESULTS
I. The sections of the ITS region and flanking parts
The amplified fragment of nuclear rDNA including the ITS region is about 560 base pairs
long in cetrarioid lichens, and can be divided
into five easily distinguished sections: The last
part of the SSU rDNA gene (1) at the 5’ end followed by the highly variable ITS 1 section (2),
the conserved 5.8 S rDNA (3), the ITS 2 section
(4), which is somewhat less variable than ITS 1,
and, finally the 5’-end of the large conserved LSU
rDNA gene (5). In the alignment, the five sections are recognized at the following sites (Table
1): 1: 1–33, 2: 34–223, 3: 224–381, 4: 382–536
and 5: 537–577. Group I introns of c. 220 nt
length, which are excluded from the analyses,
were found close to the end of the small subunit
ribosomal DNA gene at position 1516 in four
species (Gargas et al., 1995).
II. Variation in ITS1 and ITS2
The variation within and between species is
restricted to the ITS 1 and ITS 2 regions, which
are rapidly degraded during rRNA processing
(between the bases 33 and 224 and 380–535).
As in related investigations (Thell et al., 1998;
Thell & Miao in prep.), a larger variation was
found in the ITS 1 when compared with ITS 2.
115
In this study, infraspecific variation was found
at 16 positions in the ITS 1 section but only at
two in ITS 2 (Table 1).
III. The sequenced species and infraspecific
variation
1. AHTIANA SPHAEROSPORELLA (Müll. Arg.) Goward.
Only one specimen, which included a group
I intron, was available for molecular studies
from this species (DNA#AT83).
2. ESSLINGERIANA IDAHOENSIS (Essl.) Hale & M.J.
Lai. Two samples (DNA#AT145, 146) from
different localities in the same area had
identical sequences.
3. K AERNEFELTIA MERRILLII (Tuck.) A. Thell &
Goward. The two studied specimens
(DNA#AT71, 190) were morphologically
similar. Their sequences, however, differed
at several positions: 113, 114, 146, 181 and
187 in the ITS 1 section, and at 456 and 527
in the ITS 2 section (Table 1).
4. TUCKERMANNOPSIS AMERICANA (Spreng.) Hale
[syn. Cetraria ciliaris Ach. var. halei (W. L.
Culb. & C. F. Culb.) Ahti]. The following sites
were different in two sequenced specimens:
34, 99, 113, 136, 138, 143, 166, 201, 206
and 211. Group I introns were found in both
sequences (DNA#AT82, 148).
5. TUCKERMANNOPSIS PLATYPHYLLA (Tuck.) Hale.
The ITS region differed only at one position
in both samples sequenced: 168 (Table 1).
Group I introns were detected in both sequences (DNA#AT43, 75).
6. TUCKERMANNOPSIS SUBALPINA (Imshaug) Kärnefelt.
A group I intron was detected in the single
sample sequenced (DNA#AT109).
7. VULPICIDA CANADENSIS (Räsänen) J.-E. Mattsson
& M.J. Lai. The two sequences (DNA#AT36
and DNA#VM174) were identical (Table 1).
DISCUSSION
Using Platismatia as the outgroup taxon, two
main group where apparent in the ITS phylogeny (Fig. 1). One of these, supported by a bootstrap value of 71, includes Cetraria islandica,
C. sepincola, Vulpicida canadensis, V. pinastri
and Tuckermannopsis subalpina.
Vulpicida contains two distinct species
groups. One group, containing the North American endemics V. canadensis and V. viridis, is
Fig. 1. The bootstrap consensus tree based
on ITS sequences from Table 1. Tree length =
262, CI = 0.537, generated using the branch
and bound option in PAUP 3.0. The numbers
above or to the right of the branches are substitution numbers per branch. Branch lengths
are proportional to the substitution numbers.
Bootstrap percentages (from 500 replicates) are
in bold face below the branches.
characterized by citriform conidia and asci with
less amyloid and smaller tholi. The other group,
including four species, produces sublageniform
conidia and asci showing a strong amyloid reaction in the tholus (Mattsson & Lai, 1993;
Mattsson, 1993). These two groups, represented
by one species each in the analysis appeared as
a monophyletic genus.
The two species of Cetraria included, C. islandica and C. sepincola, did not form a monophyletic group. This is not surprising because
only C. islandica is considered as a Cetraria in
a strict sense (Kärnefelt et al., 1993). Cetraria
sepincola has asci of Lecanora type (Melanelia form) instead of Cetraria type (Thell et al.,
1995b). However, more species related to C.
sepincola, e.g. C. fendleri and C. weberi, need
to be analysed before an alternative taxonomy
might be considered. At least, a position in
Tuckermannopsis for C. sepincola, as proposed
by Hale (in Egan, 1987), is not supported by
ITS data. C. islandica and C. sepincola are on
a polytomous branching with Vulpicida (Fig. 1)
116 Folia Cryptog. Estonica
Table 1. ITS sequences of seven cetrarioid species from British Columbia. Sequences of Esslingeriana idahoensis and Vulpicida canadensis were identical within the species
* Infraspecific variation in Kaernefeltia merrillii, Tuckermannopsis americana and T. platyphylla
117
Table 1 (continued)
118 Folia Cryptog. Estonica
Table 1 (continued)
119
Table 1 (continued)
120 Folia Cryptog. Estonica
and the ITS data suggests that Vulpicida and
Cetraria s. str. are quite close. This is surprising
considering the chemical differences.
Tuckermannopsis subalpina forms a basal
clade in the first main group. Interestingly,
Kärnefelt (1979) pointed out earlier that Cetraria species and Masonhalea are closely
related to Tuckermannopsis subalpina (earlier
Cetraria subalpina), but more recent studies on
reproductive characters suggested a position in
Tuckermannopsis for this species together with
closely related T. inermis (Kärnefelt et al., 1993).
However, both taxa belong to neither Tuckermannopsis nor Cetraria, as suggested by ITS
data as well as by morphological characters:
an one-layered cortex, spherical ascospores and
asci with large axial bodies does not support a
position in Cetraria, whereas the pycnocondial
type – a long, narrow shape with sharp ends
– differs from the dumbbell shape characteristic
for the genus Tuckermannopsis (Kärnefelt, 1979;
Kärnefelt et al., 1993).
The other main group of cetrarioid taxa
includes Ahtiana sphaerosporella, Kaernefeltia
merrillii, Tuckermannopsis americana, T. chlorophylla, Flavocetraria cucullata, the monotypic
genus Esslingeriana and Tuckermannopsis
platyphylla. This group is supported by a bootstrap value of 51 (Fig. 1).
Tuckermannopsis platyphylla has a characteristic appearance with large, almost Umbilicaria-like papillate and tuberculate lobes
(Esslinger, 1973). Tuckermannopsis orbata is
presumably the most closely related species
but T. platyphylla may also be difficult to separate from broad-lobed specimens of Kaernefeltia
merrillii, as well. The bootstrap consensus tree
would support a new generic position for T.
platyphylla (Fig. 1), but additional taxa have to
be examined before an alternative systematic
position is proposed.
Esslingeriana idahoensis, the rarest of the
lichens represented here, is an example of a
cetrarioid lichen with both laminal and marginal
apothecia and pycnidia. Its morphology is very
distinct, characterized by a foliose habit, a grey
upper surface and large, dark pigmented cells
in the lower cortex. Atranorin and endocrocin
are distinct secondary compounds (Esslinger,
1971; Kärnefelt et al., 1992). The long branch
of 28 steps agrees with an isolated position for
this monotypic genus.
The genus Flavocetraria is characterized by
asci of Cetraria type and the genus was earlier
presumed to have close affinities with Cetraria s.
str. (Kärnefelt et al., 1992). The unexpected position of F. cucullata presented here indicates that
Cetraria is quite distinct from Flavocetraria. The
pycnoconidia, however, are dumbbell shaped,
a character shared with all taxa in this second
main group of the bootstrap consensus tree (Fig.
1). Preliminary data suggest that Flavocetraria
cucullata and F. nivalis group together as a
monophyletic genus (Thell & Miao in prep.).
The sister group of Flavocetraria cucullata included British Columbian species in a
polytomous branching (Fig. 1). One of these
species is Tuckermannopsis americana, which
is represented by two specimens on the tree.
This taxon is indistinguishable from the type
species, T. ciliaris, and sometimes recognized
only at variety level, as Cetraria ciliaris Ach.
var. halei (W. L. Culb. & C. F. Culb.) Ahti (in
Brodo, 1984: 100; Goward et al., 1994). It is
recognized by presence of alectoronic acid and
occasionally by α-collatolic acid (Culberson &
Culberson, 1967). Similar as for the chemical
species of Cetrelia, the species status for this
taxon is uncertain. The four species composing
the T. ciliaris group, T. americana, T. ciliaris, T.
microphyllica and T. orbata, are closely allied according to morphology and secondary chemistry
(Culberson & Culberson, 1967).
The bootstrap consensus tree (Fig.1) suggests that if Tuckermannopsis chlorophylla with
world-wide distribution and probably also the
related eastern Asian T. gilva and T. ulophyllodes are also maintained in Tuckermannopsis,
then the genera Ahtiana and Kaernefeltia have
to be included as well. Ahtiana sphaerosporella
is morphologically distinct form other groups
and was originally distinguished from Parmelia
as a monotypic genus – primarily on the basis of
small and spherical ascospores (Goward, 1985),
whereas Ahtiana aurescens and A. pallidula were
transferred later from Cetraria (Thell et al.,
1995a). Ahtiana is presumably most closely allied to the likewise yellow, Eastern Asian genus
Tuckneraria which differs in the presence of cilia
and pseudocyphellae (Randlane et al., 1994).
The type species of Tuckneraria and Allocetraria
must certainly be sequenced to obtain a more
detailed picture of the evolution of the group.
121
Kaernefeltia was recently described and differs
from other cetrarioid genera by its unique combination of characters (Thell & Goward, 1996).
However, there is considerable morphological
variation in the genus and the growth form of
Kaernefeltia merrillii varies from broadly foliose
to subfruticose. Unfortunately, suitable material
for molecular studies was not available from the
type species K. californica.
At present the data show that Tuckermannopsis is not only poorly delimited to members
of other genera in the current circumscription
but species assigned to Tuckermannopsis are
polyphyletic according to the ITS data. Also,
the status of several other genera is difficult to
ascertain until more species assigned to Tuckermannopsis are sequenced. Even though this
taxon is heterogeneous, it appears to be a key
group for a better phylogenetic understanding of
cetrarioid lichens. To improve the phylogenetic
hypothesis, further cetrarioid genera also need
to be compared with non-cetrarioid representatives from different segregates of the Parmeliaceae. This is especially important since gross
morphology or chemistry are not supported as
characters to separate larger units in neither
parmelioid nor cetrarioid lichens (for chemistry
compare with Poelt & Leuckert, 1993). As a consequence, generic rearrangements in cetrarioid
lichens should be carried out with great care
until a more detailed phylogeny is at hand.
ACKNOWLEDGEMENTS
I was delighted to contribute in this special
volume in honour of Prof. Hans Trass, who has
played an important role in the fruitful research
coorperation between lichenologists in Tartu and
Lund. My sincere thanks to Martin Grube, Vivian Miao, Tiina Randlane and Andres Saag for
reading and commenting on the manuscript.
Fellowships covering expenses connected
with this study were received from the Swedish
Royal Academy of Sciences, Per-Erik Lindahls
Foundation and from the European Comission, Human Capital and Mobility ”Large Scale
Facility” contract no. ERBCHGECT940065 between the EU and the University of Helsinki,
Department of Ecology and Systematics.
REFERENCES
Brodo, I. M. 1984. Lichenes Canadenses Exsiccati:
Fascicle III. Bryologist 87: 97–111
Culberson, W. L. & Culberson, C. F. 1967. A new
taxonomy for the Cetraria ciliaris group. Bryologist 70: 158–166.
Egan, R. S. 1987. A fifth checklist of the lichen-forming, lichenicolous and allied fungi of the continental United States and Canada. Bryologist 90:
77–173.
Egan, R. S. 1991. Changes to the ”fifth checklist of the
lichen-forming, lichenicolous and allied fungi of
the continental United States and Canada. Bryologist 94: 396–400.
Esslinger, T. L. 1971. Cetraria idahoensis, a new species of lichen endemic to western North America.
Bryologist 74: 364–369.
Esslinger, T. L. 1973. Chemical and taxonomic studies
on some corticolous members of the lichen genus
Cetraria in western North America. Mycologia 65:
602–613.
Gargas, A., DePriest, P. T. & Taylor, J. W. 1995. Positions of multiple insertions in SSU rDNA of lichenforming fungi. Mol. Biol. Evol. 12: 208–218.
Gilbert, D. 1993. SeqApp, a biological sequence editor and analysis program. Indiana University,
Bloomington.
Goward, T. 1985. Ahtiana, a new lichen genus in the
Parmeliaceae. Bryologist 88: 367–371.
Goward, T., McCune, B. & Meidinger, D. 1994. The
lichens of British Columbia. Illustrated keys. Part
1. Foliose and squamolose species. Ministry of
Forests Research Program. 181 pp.
Gyelnik, V. 1933. Lichenes varii novi critique. Acta pro
Fauna et Fl. Univ. 2, 1 (5–6): 3–10.
Huang, X. 1992. A contig assembly program based on
sensitive detection of fragment overlaps. Genomics 14: 18–25.
Kärnefelt, I. 1979. The brown fruticose species of
Cetraria. Opera Bot. 46, 150 pp.
Kärnefelt, I., Mattsson, J.-E. & Thell, A. 1992. Evolution and phylogeny of cetrarioid lichens. Pl. Syst.
Evol. 183: 113–160.
Kärnefelt, I., Mattsson, J.-E. & Thell, A. 1993. The lichen genera Arctocetraria, Cetraria and Cetrariella
(Parmeliaceae) and their presumed evolutionary
affinities. Bryologist 96: 394–404.
Kurokawa, S. 1991. Japanese species and genera of
the Parmeliaceae. J. Jap. Bot. 66: 152–159.
Lai, M. J. 1981 [1980]. Studies on the cetrarioid lichens in Parmeliaceae of east Asia (1). Quart. J.
Taiwan Mus. 33: 215–229.
Mattsson, J.-E. 1993. A monograph of the genus
Vulpicida. Opera Bot. 119: 1–61.
Mattsson, J.-E. & Lai, M. J. 1993. Vulpicida, a new
genus in Parmeliaceae (Lichenized Ascomycetes).
Mycotaxon 49: 425–428.
122 Folia Cryptog. Estonica
Poelt, J. & Leuckert, C. 1993. Substitution and supplementary addition of secondary products in
the evolution of lichenized Ascomycotina. Bibl.
Lichenol. 53: 201–215.
Randlane, T., Saag, A., Thell, A. & Kärnefelt, I. 1994.
The genus Tuckneraria Randlane & Thell – a new
segregation in the family Parmeliaceae. Acta Bot.
Fenn.150: 143–151.
Randlane, T., Saag, A. & Thell, A. 1997. A second
updated world list of cetrarioid lichens. Bryologist
100: 109–122.
Thell, A. 1995. A new position of the Cetraria commixta
group in Melanelia (Ascomycotina, Parmeliaceae).
Nova Hedwigia 60: 407–422.
Thell, A., Berbee, M. & Miao, V. 1998. Phylogeny of the
genus Platismatia based on rDNA ITS sequences
(Lichenized Ascomycotina). Crypto-gamie, Bryol.Lichénol. (in press)
Thell, A. & Goward, T. 1996. The new cetrarioid
genus Kaernefeltia and related groups in the
Parmeliaceae (Lichenized Ascomycotina). Bryologist 99: 125–136.
Thell A., Goward, T., Randlane, T., Kärnefelt, I & Saag,
A. 1995a. A revision of the North American genus
Ahtiana (Parmeliaceae). Bryologist 98:596–605.
Thell, A., Mattsson, J.-E. & Kärnefelt, I. 1995b. Lecanoralean ascus types in the lichenized families
Alectoriaceae and Parmeliaceae. Crypt. Bot. 5:
120–127.
Thell A., Randlane, T., Kärnefelt, I., Gao, X. & Saag,
A. 1995c. The lichen genus Allocetraria (Ascomycotina, Parmeliaceae). In Flechten Follmann.
Contributions to lichenology in honour of Gerhard
Follmann (eds Daniels, F. J. A., Schultz, M. &
Peine, J.), pp. 353–370. University of Cologne,
Germany.
White, T. J., Burns T., Lee, S. & Taylor, J. 1990. Amplification and direct sequencing of fungal ribosomal
DNA genes for phylogenetics. In PCR protocols: A
guide to methods and applications (eds Innis, M.,
Gelfand, J., Sninsky, J. & White, T.), pp. 315–322.
Academic Press, Orlando, Florida.
Folia Cryptog. Estonica, Fasc. 32: 123–125 (1998)
16 lichens new to Estonia
Göran Thor & Anders Nordin
Department of Systematic Botany, Uppsala University, Villavägen 6, S-752 36 Uppsala, Sweden
Abstract: An investigation of three wooded meadows in Saaremaa and Läänemaa Counties and short visits at some other
localities in 1991 resulted in the finding of 16 lichen species new to Estonia, Buellia violaceofusca, Caloplaca herbidella, Cyphelium
sessile, Fuscidea pusilla, Gyalecta truncigena, Lauderlindsaya acroglypta, Lecanora strobilina, Lepraria eburnea, Lepraria lobificans, Leptogium
teretiusculum, Micarea peliocarpa, Ochrolechia subviridis, Opegrapha herbarum, O. sorediifera, Rinodina efflorescens and Ropalospora viridis.
Kokkkuvõte: G. Thor ja A. Nordin. 16 Eestile uut samblikku.
Uurimistööde tulemusena kolmel Saaremaa ja Läänemaa puisniidul ning mõnedes teistes kasvukohtades 1991. a-l leiti 16
Eestile uut samblikuliiki: Buellia violaceofusca, Caloplaca herbidella, Cyphelium sessile, Fuscidea pusilla, Gyalecta truncigena, Lauderlindsaya
acroglypta, Lecanora strobilina, Lepraria eburnea, Lepraria lobificans, Leptogium teretiusculum, Micarea peliocarpa, Ochrolechia subviridis,
Opegrapha herbarum, O. sorediifera, Rinodina efflorescens jaRopalospora viridis.
INTRODUCTION
In August 24–30, 1991 the lichen flora of three
wooded meadows in Estonia was investigated,
two of which were situated in the island of
Saaremaa and one on the Estonian coast c. 30
km E of Saaremaa (Fig. 1, localities no. 1, 4,
5). Some additional localities were also shortly
visited, including two in the vicinity of Tartu,
where the excursion ended up (Fig. 1, localities
2, 3, 6, 7).
Traditionally managed wooded meadows in
Sweden usually have a species rich lichen
flora, including several red-listed species. This
is mainly due to the continuity in tree growth
in combination with the open forest structure,
characteristic of traditional management practices. The red-listed species are mainly found
on broad-leaved deciduous trees (Thor, 1998).
Also the Estonian wooded meadows investigated
proved to be species rich. Areas with wooded
meadows (and wooded pastures) have decreased
dramatically both in Sweden and Estonia and
are now restricted to very small areas. The
remaining wooded meadows and pastures
consequently have high conservation values.
The trend away from traditional management
practices of such habitats, resulting in denser
forest structure, has become a threat even in
protected areas in Sweden. Practices such as
pollarding is apparently positive to some lichens, e.g. Caloplaca chrysophthalma Degel., a
red-listed species in Sweden (Thor, 1996). Pol-
Fig. 1. Localities visited.
larded trees, however, are rare in the Estonian
wooded meadows investigated compared with
those of Gotland, Sweden, also investigated by
the authors (together with R. Sundin; Nordin
et al., 1990, 1991, 1992, 1996). A few species
found in the Estonian wooded meadows were
not found in Gotland, e. g. Physcia semipinnata
(J. F. Gmelin) Moberg (AN & GT 164, locality 4,
on Fraxinus excelsior).
RESULTS
Altogether 12 species new to Estonia were
found in the wooded meadows, six exclusively
in Laelatu, two exclusively in Tagala, one exclusively in Loode, two in Laelatu and Tagala
124 Folia Cryptog. Estonica
together, and one in the three of them. Four
additional species new to Estonia were found
at other localities specified below. At the same
excursion Cliostomum flavidulum Hafellner &
K. Kalb was collected and published as new to
Estonia by Nordin et al. 1996 (as Lecanora navarrensis Etayo). One more lichen new to Estonia
– Phaeophyscia endophoenicea (Harm.) Moberg
– was collected in Tartumaa Co., Luke park.
This finding has been published already in the
macrolichen flora of Estonia (Trass & Randlane, 1994). Four of the species reported here
are red-listed in Sweden, Buellia violaceofusca
(Care demanding), Cyphelium sessile (Care demanding), Gyalecta truncigena (Vulnerable) and
Opegrapha sorediifera (Vulnerable). The material
will be deposited in UPS. For collections investigated by TLC the results are given together with
the collection numbers. The numbers (1–7) refer
to localities.
List of species new to Estonia
BUELLIA VIOLACEOFUSCA G. Thor & Muhr: 4 (no
substances found by TLC, AN & GT 131).
– On Quercus robur.
CALOPLACA HERBIDELLA (Hue) H. Magn.: 5 (AN & GT
216). – On Quercus robur.
CYPHELIUM SESSILE (Pers.) Trevis.: 4 (AN & GT 69;
AN & GT 150). – On Pertusaria coccodes on
Fraxinus excelsior (no. 150) and Quercus
robur (no. 69).
FUSCIDEA PUSILLA Tønsberg: 1 (divaricatic acid, AN
& GT 55), 5 (divaricatic acid, AN & GT 206).
– On Betula.
GYALECTA TRUNCIGENA (Ach.) Hepp: 7 (AN & GT 280).
– On Quercus robur.
LAUDERLINDSAYA ACROGLYPTA (Norman) R. Sant.: 1
(AN & GT 61), 5 (AN & GT 201). – On Malus
sylvestris (no. 61) and Tilia cordata (no. 201);
no perithecia found
LECANORA STROBILINA (Spreng.) Kieff.: 1 (usnic acid,
zeorin, unidentified triterpen, AN & GT 57),
5 (usnic acid, zeorin, unidentified triterpen,
AN & GT 207), 5 (usnic acid, zeorin, unidentified triterpen, AN & GT 237), 5 (usnic
acid, zeorin, unidentified triterpen, AN & GT
277). – On Betula (no. 57, 207) and Quercus
robur (237, 277).
LEPRARIA EBURNEA J.R. Laundon: 1 (alectorialic,
barbatolic, protocetraric acids, AN & GT 37).
– On Fraxinus excelsior.
LEPRARIA LOBIFICANS Nyl.: 2 (atranorin, stictic acid
complex, zeorin, GT 10283), 4 (atranorin,
stictic acid complex, zeorin, AN & GT 89), 8
(atranorin, stictic acid complex, zeorin, AN
& GT 291). – On Fraxinus excelsior (10283),
Quercus robur (no. 89) and Tilia cordata (no.
291).
LEPTOGIUM TERETIUSCULUM (Wallr.) Arnold: 1 (AN &
GT 38). – On Fraxinus excelsior.
MICAREA PELIOCARPA (Anzi) Coppins & R. Sant.: 1
(AN & GT 53). – On Betula.
OCHROLECHIA SUBVIRIDIS (Hoeg) Erichsen: 3 (GT
10297). – On Quercus robur.
OPEGRAPHA HERBARUM Mont.: 1 (AN & GT 12). – On
Quercus robur.
OPEGRAPHA SOREDIIFERA P. James: 5 (AN & GT 232).
– On Quercus robur.
RINODINA EFFLORESCENS Malme: 1 (pannarin, AN
& GT 31), 4 (pannarin, AN & GT 101), 5
(pannarin, AN & GT 211; pannarin, AN &
GT 247). – On Quercus robur (no. 31, 101,
247) and Betula (211).
R OPALOSPORA VIRIDIS (Tønsberg) Tønsberg: 2
(perlatolic acid, GT 10280b). – On Populus
tremula.
Localities
1. Läänemaa Co., Laelatu wooded meadow,
2 km E of Virtsu, 58°35’N, 23°35’E, 24–26
August 1991. Open, mowed wooded meadow
dominated by Quercus robur.
2. Läänemaa Co., 2 km SSE of Virtsu, at the ornithological field station at Puhtu, 58°34’N,
23°33’E, 26 August 1991. Dense, old broadleaved deciduous forest at the Baltic sea.
3. Saaremaa Co., Ratla pasture, just SE of the
road between Orissaare and Kuressare 30
km NE of Kuressare, 58°28’N, 22°51’E, 27
August 1991. Open pasture with scattered
Quercus robur.
4. Saaremaa Co., Loode wooded meadow, 4 km
W of Kuressare, just S of the road between
Kuressare and Salme, 58°14’N, 22°25’E, 27–
28 August 1991. Moowed wooded meadow
(small parts mowed every year), dominated
by Fraxinus excelsior and Quercus robur.
5. Saaremaa Co., Tagala wooded meadow, just
W of Veere, 58°28’N, 21°59’E, 29–30 August
1991. Open, mowed wooded meadow with
deciduous trees.
125
6. Tartumaa Co., 14 km SSW of Tartu, Unipiha
park, 58°16’N, 26°37’E, 2 September 1991.
Avenue and park with scattered broadleaved deciduous trees.
7. Tartumaa Co., 16 km SSW of Tartu, Luke
park, 58°14’N, 26°33’E, 2 September 1991.
Overgrown park dominated by Tilia cordata.
ACKNOWLEDGEMENTS
We wish to thank H. Trass, T. Randlane, and
A. Saag for their excellent organisation of the
excursion and for assistance in the field. T.
Randlane kindly informed us on which species
were new to Estonia.
REFERENCES
Nordin, A., Sundin, R. & Thor, G. 1990. Lavinventering
i ängen. Rindi 10: 35–36.
Nordin, A., Sundin, R. & Thor, G. 1991. Lecanora strobilina and Polyblastia allobata new to Sweden.
Graphis Scripta 3: 50–51.
Nordin, A., Sundin, R. & Thor, G. 1992. Bacidia caligans and Bacidina chloroticula new to Sweden.
Graphis Scripta 3: 134–137.
Nordin, A., Sundin, R. & Thor, G. 1996 [1995]. Two
sorediate crustose lichens assigned to Lecanora.
Nord. J. Bot. 15: 553–556.
Thor, G. 1996. The biology and distribution of three
red listed lichens in Sweden. Symb. Bot. Upsal.
31: 3, 355–363.
Thor, G. 1998. Red-listed lichens in Sweden: habitats,
threats, protection, and indicator value in boreal
coniferous forests. Biodiversity and Conservation
7: 59–72.
Trass, H. & Randlane, T. (eds) 1994. Eesti suursamblikud. Tartu. 399 pp.
126 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 127–133 (1998)
New and rare calicioid lichens and fungi from relict tertiary forests
of Caucasus and the Crimea
Alexander N. Titov
Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov St., 197376 St. Petersburg, Russia
Abstract: 18 rare species of calicioid lichens and fungi are reported from tertiary forests of Caucasus and the Crimea.
Chaenothecopsis trassii Titov is described as new. One new combination, Chaenothecopsis jaczewskii (Woronich.) Titov, is proposed.
The habitat ecology and distribution of the species are presented.
Kokkuvõte: A. Titov. Uued ja haruldased kalitsioidsed samblikud ja seened Kaukaasia ja Krimmi reliktsetest tertsiaari metsadest.
Kaukaasia ja Krimmi reliktsetest tertsiaari metsadest loetletakse 18 haruldast kalitsioidset samblikku ja seent. Kirjeldatakse
üks uus liik – Chaenothecopsis trassii Titov; esitatakse üks uus kombinatsioon – Chaenothecopsis jaczewskii (Woronich.) Titov.
Käsitletakse liikide levikut ja nende kasvukohtade ökoloogiat.
INTRODUCTION
In accordance with Takhtadjan (1986) floristic
division, Caucasus and the Crimea are situated
on the boreal and tethyan subkingdoms of Holarctis. Although circumboreal elements dominate the flora of Caucasus and the Crimea, many
relict islands of an ancient tertiary flora remain.
There are two main centres of the tertiaty flora
– the dry subtropical Hyrcanian forest and the
wet subtropical forests of Colchis.
Relict Hyrcanian forests (the name originates from the ancient name of the Caspian Sea)
form a separate province of the Irano-Turanion
region, and occupy the northeastern slope of
the Talysh range. The majority of forest-forming
trees, such as Acer vetulinum Boiss., Albizia julibrissin Durazz., Gleditschia caspia Desf., Parrotia persica C.A.M., Quercus castaneifolia C.A.M.,
Zelkova carpinifolia (Pall.) C. Koch are endemics
of the mesophilous tertiary forest.
The flora of the Colchian type (this name
originates from the name of the territory in West
Georgia – the lowlands of the river Rioni) form
the Euxine province, and occurs in the western
Caucasus – Adzharssk, Abkhassk and Krasnodar territories. In contrast to the Hyrcanian
province, where conifers do not occur, the main
forest-forming trees in the Euxine province are
endemic conifers – Abies nordmanniana (Stev.)
Spach and Picea orientalis (L.) Link – together
with Acer trautvetteri Medw., Carpinus betulus
L., Fagus orientalis Lipsky, Pinus kochiana
Klotzsch ex C. Koch, Quercus spp., Tilia begoniifolia Stev.
The forests of the Crimea are mainly of a
broad-leaved Fagus-Quercus type, and were
included by Takhtadjan (1986) in a separate
Crimean-Novorossijsk province. The flora of
these forests is quite similar to that of the Colchian type.
During our field trips in Caucasus and the
Crimea more than 50 species of calicioid lichens
and fungi were found. In this paper only some
rare, threatened, or earlier overlooked species
are included.
All higher plants are named according to
Czerepanov (1995), and lichens according to
Santesson (1993).
MATERIAL AND METHODS
The material was collected from several localities
in Caucasus and the Crimea by the author (Fig.
1) in the period from 1982–1993. The results
are also based on herbarium material from LE,
LEP and KW. For SEM, material was mounted
on specimen stubs, air-dried and coated with
gold. Estimates of the apothecium height,
capitulum width, stalk width, ascus length,
ascus width, spore length and spore width (for
Chaenothecopsis trassii) are given in the following way: (min) a–b (max) – or for combinations of
length and width: (min) a–b × c–d (max)– where
a (and c) represent the arithmetic mean minus 1
128 Folia Cryptog. Estonica
Fig. 1. The map of Caucasus and Crimea.
standard deviation and b (and d) the arithmetic
mean plus 1 standard deviation. The extreme
minimum (min) and maximum (max) values are
given within parentheses.
THE SPECIES
CHAENOTHECA SERVITII Nadv., Fedde, Repert. 36:
308 (1934).
Caucasus. Russia, Krasnodar territory, Adygeisk autonomous region, Guzeripl, plateau
Abago, alt. ca 1500 m, 1993 Titov 4528 (LE).
On lignum of Abies nordmanniana.
C. servitii is known only from the type locality in Carpathians (Nadvornik, 1934).
Here it is reported as a new for Russia and
Caucasus.
CHAENOTHECA SUBROSCIDA (Eitner) Zahlbr., Cat. lich.
univ. 578 (1922) – Cyphelium subroscidum
Eitner, Jahrb. Schlesis. Ges. vaterl. Cult.
88: 53 (1911).
Caucasus. Georgia, Abkhazsk, Gudauta region, lake Ritza, alt. ca 900 m, 1988 Titov
1480, 1555 (LE). On bark and lignum of
Abies nordmanniana.
C. subroscida occurs in Europe and North
America, usually on bark of Picea (Tibell,
1980). In the former USSR it has not previously been found east of Estonia. It is here
reported new to Asia.
CHAENOTHECOPSIS BREVIPES Tibell, Symb. Bot. Ups.
27(1): 119 (1987).
Caucasus. Russia, Krasnodar territory,
Adygeisk autonomous region, Mezmaj, alt.
ca 900 m, 1993 Titov 4543 (LE). On bark of
Acer trautvetteri.
C. brevipes seems to be rare species, occuring on the thallus of Arthonia growing on
bark of deciduous trees in shaded situations
in cold temperate, wet forests. Originally described from New Zealand (Tibell, 1987), it
has later been found only in a few localities
129
in eastern North America (Selva, 1988),
Russian Far East (Titov & Tibell, 1993) and
Europe (Tibell & Ryman, 1995). It is here
reported new to Caucasus.
CHAENOTHECOPSIS GOLUBKOVAE Tibell & Titov in Titov
& Tibell, Nord. J. Bot. 13: 320 (1993).
Caucasus. Russia, Krasnodar territory,
Adygeisk autonomous region, 2 km E of
Kamyshanova polyana, alt. ca 1200 m, 1982
Titov 303 (type: LE; isotype: Tibell, Cal. Exs.
211); Guzeripl, plateau Abago, alt. ca 1500
m, 1993 Titov 4527 (LE). Georgia, Borzhomi
region, Tadsrisi, alt. ca 1000 m, 1983 Titov
1460 (LE). On resin of Abies nordmanniana
and Picea orientalis.
C. golubkovae occurs on bark of Abies, Picea
and Pinus in open situations in temperate
forests. It was found mainly in cracks of
the bark where exudate had accumulated.
It was recently included in a revision of
resinicolous calicioid fungi (Tibell & Titov,
1995). C. golubkovae seems to be a rare species. It is known only from a few localities in
Caucasus, Russian Far East (Titov & Tibell,
1993) and China (unpublished).
CHAENOTHECOPSIS HOSPITANS (Th. Fr.) Tibell, Nova
Hedwigia 60: 202 (1995) – Calicium hospitans Th. Fr., Bot. Notiser: 40 (1865).
Caucasus. Georgia, Borzhomi region, Tadsrisi, alt. ca 1000 m, 1983 Titov 1351 (LE).
On thallus of Lecanora carpinea (L.) Vain.
a
C. hospitans occurs as a parasite-parasymbiont on the thallus of Lecanora carpinea
growing on deciduous trees, and also on
Haematomma ochroleucum (Neck.) Laundon on shaded, siliceous rocks. It was previously known only from Central Europe and
Scandinavia. New to Asia (Tibell & Ryman,
1995).
CHAENOTHECOPSIS JACZEVSKII (Woronich.) Titov,
comb. nov. – Protocalicium jaczevskii Woronich., Travaux du Mus. Bot. 21 (1927): 103.
Caucasus. Georgia, Gori region, Licani,
1920, coll. N. N. Voronichin (holotype: LEP).
On mycelium of Hormiscium pinofilum (Nees)
Lind., growing on twigs of Picea orientalis.
Mycelium well developed, consisting of
pale, irregular intervowen hyphae about 3
µm thick, mixed with the host mycelium. Algae not present in the substrate. Apothecia
small, 0.2–0.4 mm high, on pale stalks. Capitulum ellipsoidal, black, 0.06–0.1 mm in
diameter. Hymenium and hypothecium pale.
Excipulum well-developed as a continuation
of the outer part of the stalk, consisting of
4–5 layers of periclinally elongated cells,
with a constricted margin. Stalks straight,
0.02–0.03 mm in diameter, consisting of
loosely interwoven pale hyphae. Mature asci
55–70 × 4–5 µm, cylindrical, always with a
canal in the apical thickening. Spores uni-
b
Fig. 2. Chaenothecopsis jaczevskii. a – apothecium, assotiated with mycelium of Hormiscium
pinophilum; b – spores.
130 Folia Cryptog. Estonica
seriately arranged in the asci, periclinally
orientated, 1-septate, narrowly-ellipsoidal
with rounded ends, pale-brown, 7–9 × 3–4
µm, with a well-developed, areolate ornamentation. All parts of the apothecium K-,
H-, strongly swelling with K. Hymenium
slightly blue-green with iodine (Fig. 2).
C. jaczevskii is characterized by its green in
fresh material (Voronichin, 1927), well developed mycelium, small apothecia on pale
stalks and ellipsoidal capitula, constricted
excipulum margin and one-septate ornamented spores.
C. jaczevskii is known from one collection
only and the material is rather scanty for
suppling a good description, but this is
evidently a distinctive species. In many
respects it is quite different from other
genera of calicioid fungi, and only placed
in Chaenothecopsis tentatively, since the
generic delimitations in Mycocaliciaceae
are in need of reappraisal.
C HAENOTHECOPSIS OCHROLEUCA (Körb.) Tibell &
K. Ryman, Nova Hedwigia 60: 208 (1995)
a
– Calicium ochroleucum Körb., Parerga Lich.:
295 (1865).
Caucasus. Azerbaijan, Lencoran region, Belyasar, 1987 Titov, 1431, 1443, 1458 (LE).
On thallus of Haematomma ochroleucum
(Neck.) Laundon, growing on bark of Fagus
orientalis. C. ochroleuca occurs on the bark
of deciduous trees in humid situations in
temperate forests. A rare species, C. ochroleuca is previously known from some localities of the northern Eurasia. Originally
described from Germany (Koerber, 1865), it
has also been reported from Central Europe
(Nádvorník, 1934, 1942; Tibell & Ryman,
1995) and the Russian Far East (Titov &
Tibell, 1993). New to Caucasus.
CHAENOTHECOPSIS TRASSII Titov sp. nov.
Apothecia parva, 0.23–0.32 mm alta, nigra,
epruinosa. Excipulum brunneum, e hyphis
periclinalibus constans, 4–6 µm crassum.
Asci 28.9–40.7 × 3.4–4.2 µm, cylindrici vel
anguste clavati. Sporae uniseptate, ellipsoideae, pallidae brunneae, laeves, 5.3–6.4
× 2.0–2.6 µm. Partes apotheciorum rubro-
b
Fig. 3. Chaenothecopsis trassii. a – apothecium; b, c – spores.
c
131
pigmentataea K addito intense virescentes,
ceterum H-, K-.
Apothecia associated with Trentepohlia, or
no photobiont present. Apothecia small,
(0.20)0.23–0.33(0.40) mm high, black.
Capitulum subspherical to obconical and
hemispherical, (0.10)0.12–0.20(0.25) mm in
diameter. Epithecium brown, 2–4 µm thick.
Hymenium and hypothecium pale. Excipulum well-developed as a continuation of the
outer part of the stalk, consisting of 3–4 layers of periclinally elongated, 4–6 µm thick,
brown cells. Stalks straight, 0.02–0.05 mm
in diameter, consisting of interwoven periclinally arranged hyphae, which are pale and
more loosely arranged in the central part
and brown in outer part. Hymenium and
more rarely hypothecium and inner part of
stalk containing numerous red crystals. Asci
(24.0)28.9–40.7(45.0) × (3.0)3.4–4.2(4.5) µm,
cylindrical or narrowly clavate, with a thin
canal in the apical thickening. Spores uni- or
rarely biseriately arranged in the asci, periclinally or obliquely orientated, non-septate,
ellipsoidal, pale-brown, (5.0)5.3–6.4(8.0) ×
(2.0)2.0–2.6(3.0) µm, smooth under the light
microscope and weakly ornamented in SEM.
All parts of the apothecium with red pigment
turn aeruginose with K, but are H- (Fig. 3,
a–c).
Caucasus. Azerbaijan, Lencoran region,
Belyasar, 1987 Titov 1766 (Holotype: LE,
isotype: UPS), Vyshnikesh, 1427 (LE). On
bark of Fagus orientalis.
C. trassii is characterized by its small,
short-stalked apothecia, the non-septate
ellipsoidal spores, the ascocarp structure
and by containing red crystals in hymenium,
which yield a green reaction with K. C. trassii
is rather similar to C. rubescens, which also
has short-stalked apothecia which contain a
red pigment, but it differs in having smaller,
smooth spores and a different K reaction. C.
trassii is also somewhat similar to C. ochroleuca, which has a K+ aeruginose reaction
of the capitulum, but fusiform dark spores
and often white pruinose stalks. C. nana
and C. amurensis both have short-stalked
apothecia and non-septate spores and could
be mistaken for C. trassii, but differ in having a negative K reaction.
C. trassii is named in honour of Prof. H.
Trass on the occasion of his 70th birthday
for his invaluable contribution to the exploration of the Russian lichen flora.
CHAENOTHECOPSIS USSURIENSIS Titov, Nord. J. Bot.
13: 326 (1993).
Caucasus. Azerbaijan, Astara region, Sym,
1987 Titov 1435 (LE). On bark of Quercus
castaneifolia.
C. ussuriensis is previously known only
from the southern Far East of Russia and
north-east of China, and seems to be a
common and widely distributed but earlier
overlooked species in relict Pinus-deciduous
forests. The occurrence of this species in the
Gyrcanian forest may indicate a connection
between the tertiary forests of Caucasus and
the Far East.
CHAENOTHECOPSIS VAINIOANA (Nádv.) Tibell, Publ.
Herb. Univ. Upsal. 4: 5 (1979) – Calicium
vainioanum Nádv., Preslia 18–19: 128
(1940).
The Crimea. Alushta region: mount Cziornaya, alt. ca 500 m, 1989 Titov 1900
(Tibell: Cal. exs. 217), 1940, 2000; cordon
“Bukovsky, alt. ca 800 m., 1989 Titov 2101,
2162, 2168 (LE). On bark and lignum of
Acer campestre L., Fraxinus excelsior L. and
Quercus robur L.
C. vainioana is known only from Europe
where it seems to be a rare species, occurring on trunks of old trees, usually Quercus,
in open situations. New for the Crimea and
Ukraine (Tibell, 1993).
CYBEBE GRACILENTA (Ach.) Tibell, Beih. Nova Hedw.
79: 666 (1984) – Calicium gracilentum Ach.,
Lich. Univ.: 243 (1810).
Caucasus. Russia: Krasnodar territory:
Adygeisk autonomous region: 5 km north
of Kamyshanova polyana, alt. ca. 1200 m.,
1982 Titov 315 (Tibell: Cal. exs. 138), 494,
495; Guzeripl, plateau Abago, alt. ca 1500
m., 1993 Titov 4527 (LE). Azerbaijan, Astara
region, Sym, 1987 Titov 1453 (LE), 1471
(Tibell: Cal. exs. 163). Georgia, Abhassk,
Gudauta region, the lake Ritza, alt. ca 900
m., 1988 Titov 1760 (LE). On bark of Acer
trautvetteri, A. velutinum and Tilia begoniifolia.
Cybebe gracilenta occurs on the bark and
lignum of deciduous trees in humid and
shaded situations in temperate forests. It
132 Folia Cryptog. Estonica
is rare in Europe and North America. In
Caucasus it was abundant in the localities mentioned above. New to Asia (Tibell,
1989).
C YPHELIUM KARELICUM (Vain.) Räsänen, Ann.
Bot. Soc. Zool.-Bot. Fenn. 12: 188 (1939)
– Cyphelium lucidum var. karelicum Vain.,
Acta Soc. F. Fl. Fenn. 57,1: 20 (1927).
Caucasus. Georgia, Abhassk, Gudauta region, the lake Ritza, alt. ca 900 m., 1988
Titov 1477, 1547, 1575, 1584 (LE). On bark
of Abies nordmanniana and Picea orientalis.
C. karelicum is a rather rare species occurring in conifer forests of Europe, North
America (Tibell, 1975) and the Siberian
Arctic (Titov, 1984). New to Caucasus.
CYPHELIUM SESSILE (Pers.) Trevis., Flora 45: 3
(1862) – Calicium sessile Pers., Tent. disp.
meth. fung.: 59 (1797).
The Crimea. Alushta region, cordon “Bukovsky”, alt. ca 800 m, 1989 Titov 2155, 2169
(LE). On thallus of Pertusaria sp., growing
on bark of Fagus orientalis.
C. sessile is known only from Europe where
it is a rare species. Usually it occurs on bark
of Quercus as a parasite-parasymbiont of
Pertusaria coccodes (Ach.) Nyl. (Tibell, 1971).
It is here reported new to the Ukraine.
MYCOCALICIUM ALBONIGRUM (Nyl.) Tibell, Lichenologist 14: 238 (1982) – Calicium albonigrum
Nyl., Syn. Lich.: 159 (1860).
The Crimea. Feodosia region, Lesnoe, alt.
ca. 400 m, 1979, coll. Kopachevskaya (KW);
Alushta region, cordon “Vesioly”, alt. ca. 600
m, 1961, coll. Roms (KW), cordon “Bukovsky”, alt. ca. 800 m, 1989 Titov 2801 (LE).
On lignum of Fagus orientalis.
M. albonigrum occurs on lignum of trees in
open situations. It is known from Australia,
and North and South America (Tibell, 1987).
M. albonigrum is quite similar to M. subtile,
which is a very widely distributed species,
but differs in having an excipulum formed
by isodiametric cells. This species might
have been overlooked since it is rather difficult to identify. It is here reported as new
for Eurasia.
PYRGIDIUM MONTELLICUM (Beltr.) Tibell, Lichenologist
14: 239 (1982) – Acolium montellicum, Beltr.
Lich. Bass.: 285 (1858).
Caucasus. Russia, Krasnodar territory,
Adygeisk autonomous region, Kamyshanova
polyana, alt. ca. 1200 m., 1982 Titov 302
(Tibell: Cal. exs. 195). Azerbaijan, Lencoran
region, Belyasar, 1987 Titov 1433 (LE). On
bark of Acer pseudoplatanus L. and Carpinus
betulus.
P. montellicum occurs on bark and lignum
in warm temperate to tropical forests. P.
montellicum is a rare but widely distributed
species, known only from rather few localities. Recorded from Italy, India, Costa Rica
(Tibell, 1982a), Australia (Tibell, 1987) and
Caucasus (Titov, 1990).
SCLEROPHORA AMABILIS (Tibell), Tibell Beih. Nova
Hedw. 79: 679 (1984) – Coniocybe amabilis Tibell, Publ. Herb. Univ. Upps. 10:6
(1982).
Caucasus. Georgia, Abhassk, Gudauta region, the lake Ritza, alt. ca. 900 m., 1988
Titov 1489 (LE). On decorticated trunks of
Hedera colchica (Koch) Koch.
S. amabilis occurs on lignum and rarely
bark of trees in shaded localities with high
humidity. Originally described from New
Zealand (Tibell, 1982b), S. amabilis was
collected in the Russian Far East (Tibell,
1993), Sweden (Gustavsson, 1995), North
America (Goward et al., 1996), and China (by
the author, unpublished). Here it is reported
as new for the Caucasus.
SCLEROPHORA CONIOPHAEA (Norm.) J. Mattson &
Middelb. in Middelb. & J. Mattson, Sommerfeltia 5: 61 (1987) – Coniocybe coniophaea
Norm., K. Norske Vidensk.-Selsk. Skr. 5:
362 (1868).
Caucasus. Russia, Krasnodar territory, Adygeisk autonomous region, Guzeripl, plateau
Abago, alt. ca 1500 m., 1993 Titov 4529 (LE).
Georgia, Abhassk, Gudauta region, the lake
Ritza, alt. ca. 900 m., 1988 Titov 1540 (LE).
On bark of Abies nordmanniana.
S. coniophaea is known only from Europe
and North America, where it is a rare species, occurring on trunks of old trees. Like
most Sclerophora species, S. coniophaea is
threatened by the disappearance of oldgrowth forests. It is here reported new to
Asia.
SCLEROPHORA PERONELLA (Ach.) Tibell, Nova Hedw.
Beih. 79; 679 (1989) – Lichen peronellus
Ach., Lichenogr. suec. prodr.: 84 (1799).
Caucasus. Georgia, Abhassk, Gudauta region, the lake Ritza, alt. ca. 900 m., 1988
Titov 1494, 1572 (Tibell: Cal. exs. 171), 1762
133
(LE). On bark of Acer trautvetteri, Fraxinus
sp.
S. peronella is known only from Europe,
where it is a rare species, occurring on
trunks of old trees. New for Asia (Tibell,
1989).
ACKNOWLEDGEMENTS
I wish to thank Dr Leif Tibell (Uppsala) for his
help throughout this study, Prof. Nina S. Golubkova (St. Petersburg) for her support of the
investigations of calicioid lichens and fungi in
Russia, and Dr Sergej Krivorotov (Krasnodar) for
organizing field trips in Caucasus.
REFERENCES
Czerepanov, S. K. 1995. Plantae vasculares rossicae et
civitatum collimitanearum. S. Petropolis. 990 pp.
Goward, T., Breuss, O., Ryan, B., McCune, B., Sipman, H. & Scheidegger, C. 1996. Notes on the
Lichens and Allied Fungi of British Columbia. III.
Bryologist 99: 439–449.
Gustavsson, H.-E. 1995. The lichen flora on beech of
Ödegärdet, Småland, SW Sweden. Svensk Bot.
Tidskr. 89: 65–82.
Körber, G.W. 1865. Parerga lichenologica. Breslau.
Nádvorník, J. 1934. Calicieae-Studien aus der Tschechoslowakei. Fedde, Repertorium 36: 307–310.
Nádvorník, J. 1942. Systematische Ubersicht der
mitteleuropäischen Arten der Flechtenfamilie
Caliciaceae. Stud. Bot. Cech. 5: 6–46.
Santesson, R. 1993. The lichens and lichenicolous fungi
of Sweden and Norway. Lund. 240 pp.
Selva, S. 1988. The Caliciales of Northern Maine.
Bryologist 91: 2–17.
Takhtadjan, A. 1986. Floristic regions of the world.
University of California press. Berkley, Los Angeles, London. 522 pp.
Tibell, L. 1971. The genus Cyphelium in Europe.
Svensk Bot. Tidskrift 65: 157–158.
Tibell, L. 1975. The Caliciales of Boreal North America.
Symb. Bot. Upsal. 21, 2: 1–128.
Tibell, L. 1980. The genus Chaenotheca in the northern hemisphere. Symb. Bot. Upsal. 23, 1: 1–65.
Tibell, L. 1982a. Caliciales of Costa Rica. Lichenologist
14, 4: 219–254.
Tibell, L. 1982b. Caliciales exsiccatae 3. Publ. Herb.
Univ. Upsal. 10: 1–10.
Tibell, L. 1987. Australasian Caliciales. Symb. Bot.
Ups. 27(1): 1–279.
Tibell, L. 1989. Caliciales exsiccatae 7. Thunbergia
8: 1–9.
Tibell, L. 1993. Caliciales exsiccatae 9. Thunbergia
18: 1–15.
Tibell, L. & Ryman, K. 1995. Revision of species of
Chaenothecopsis with short stalks. Nov. Hedw.
60: 199–218.
Tibell, L. & Titov A. N. 1995. Species of Chaenothecopsis and Mycocalicium (Caliciales) on exudate. Bryologist 98,4: 550–560.
Titov, A. N. 1984. Lichenes Caliciales in parte boreali-occidentali planitiei Putoran (in Russian).
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Titov, A. N. 1990. Pyrgidium Nyl. (Caliciales) – new
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Titov, A. N. & Tibell L. 1993. Chaenothecopsis in the
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134 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 135–147 (1998)
Die Verbreitung der Gattungen Anaptychia, Heterodermia,
Hyperphyscia und Phaeophyscia (Physciaceae) in Österreich
Von Roman Türk1 und Walter Obermayer2
Universität Salzburg, Institut für Pflanzenphysiologie, Hellbrunnerstraße 34, A-5020 Salzburg, Austria
2
Universität Graz, Institut für Botanik, Holteigasse 6, A-8010 Graz, Austria
1
Summary: The distribution of the genus Anaptychia, Heterodermia, Hyperphyscia and Phaeophyscia (Physciaceae) in Austria is
presented in form of grid maps. The patterns of distribution are explicable by the description of the ecological range of
the treated species.
Kokkuvõte: Perekondade Anaptychia, Heterodermia, Hyperphyscia jaPhaeophyscia (Physciaceae) levik Austrias.
Perekondade Anaptychia, Heterodermia, Hyperphyscia jaPhaeophyscia (Physciaceae) levik Austrias on esitatud kaartidel. Vaadeldud
liikide levikumustreid selgitatakse nende liikide ökoloogiliste nõudluste kirjeldamise abil.
EINLEITUNG
DIE VERBREITUNG IN ÖSTERREICH
Die lobaten Gattungen Anaptychia, Heterodermia,
Hyperphyscia und Phaeophyscia aus der
Familie der Physciaceae sind hinsichtlich ihrer
ökologischen Ansprüche und ihrer Verbreitung
sehr unterschiedlich. Die Abgrenzung einiger
Sippen ist zum Teil schwierig (vgl. Moberg,
1977; Hafellner et al., 1979). Nach Hafellner et
al. (1979) sind z. B. die Gattungen Anaptychia
und Physconia typisch nordhemisphärisch
temperate Genera, Heterodermia tropischsubtropische Breitgürtelelemente und
Hyperphyscia ebenfalls tropisch-subtropisch mit
einigen temperaten Ausläufern. Die Gattungen
Physcia und Phaeophyscia wiederum haben
eine sehr weite Verbreitung, die von der Arktis
und der Antarktis bis in die Tropen reicht, auf
einer Fülle von Substraten, sofern diese einen
bestimmten Grad an Eutrophierung aufweisen.
Die saxicolen Arten haben sehr unterschiedliche
Ansprüche auf das Substrat, was den Säuregrad
betrifft (vgl. Moberg, 1977).
Die Wuchsmöglichkeiten für Flechten sind in
Österreich auf Grund der anstehenden Gesteine,
der Substratvielfalt, der orographischen und
klimatischen reichen Gliederung äußerst
vielfältig und bieten Lebensraum für Flechten
mit den unterschiedlichsten ökologischen
Ansprüchen. Am Beispiel einiger Verterter
aus der Familie der Physciaceae sollen die
Verbreitungsmuster dargestellt und diskutiert
werden.
Die Literaturangaben über Funde in Österreich
zwischen 1860 und 1993 sind aus Türk & Poelt
(1993) zu entnehmen.
ANAPTYCHIA BRYORUM Poelt, syn.: A. fusca (Huds.)
Vain. var. stippea auct. (Abb. 1)
Literatur: Türk & Poelt, 1993; Hafellner,
1993; Hofmann et al., 1995.
Anaptychia bryorum wächst zumeist
an Steilflächen über Moosen und
Pflanzenresten in der alpinen Stufe (Poelt
& Vêzda, 1977). In Österreich bevorzugt sie
sauren bis leicht kalkhaltigen Untergrund
in den Zentralalpen, nur gelegentlich dringt
sie in die Nördlichen Kalkalpen (Göllmassiv,
Steinernes Meer, Hochkönig) vor.
ANAPTYCHIA CILIARIS (L.) Körb. (Abb. 2)
Literatur: Türk & Poelt, 1993; Geyer, 1985;
Neuwirth & Türk, 1993; Pfefferkorn & Türk,
1993; Hafellner & Maurer, 1994; Hofmann
et al., 1995; Baumgartner & Türk, 1996;
Berger, 1996; Boom et al., 1996; Pfefferkorn,
1996; Pfefferkorn & Türk, 1997.
Diese formenreiche Art (vgl. Poelt & Vêzda,
1977) war mit ihren epiphytisch lebenden
Vertretern früher weitaus häufiger als heute.
In ihrem heutigen Hauptverbreitungsgebiet,
im Mittelgebirge der Böhmischen Masse
und im Alpenvorland, tritt sie meist nur
in wenigblättrigen, deformierten Thalli
auf. Normal wachsende Exemplare finden
136 Folia Cryptog. Estonica
Abb. 1. Bisher bekannte Verbreitung von Anaptychia bryorum in Österreich.
Abb. 2. Bisher bekannte Verbreitung von Anaptychia ciliaris in Österreich (Punkte: aktuelle Funde
nach 1960; Kreuze: nachweislich ausgestorben; Kreis: Fund vor 1900; Kreis mit Punkt: Fund
zwischen 1900–1950).
Abb. 3. Bisher bekannte, aktuelle Verbreitung von Heterodermia obscurata in Österreich.
137
sich in den Alpentälern und südlich des
Alpenhauptkammes, wo sie sich auf
einzelstehenden Bäumen in Viehweiden
geradezu üppig entwickeln können. Als
Substrat bevorzugt sie Laubbäume mit
rauher, leicht gedüngter Borke in der
niedermontanen bis hochmontanen Stufe.
HETERODERMIA OBSCURATA (Nyl.) Trevis. (Abb. 3)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; 1994; Pfefferkorn, 1996; Türk, 1996;
Pfefferkorn & Türk, 1997.
Heterodermia obscurata ist eine ozeanische
Flechte, deren Hauptverbreitung in den
ozeanischen Bereichen im Südwesten
Europas (Küsten von Portugal, Spanien,
Frankreich und Großbritannien) liegt (vgl.
Schauer 1963). An niederschlagsreichen
Orten mit gemäßigtem Temperaturregime
findet sie sich auch in den Alpen ein,
vornehmlich in einer Seehöhe zwischen
500 bis 700 Meter. Sie siedelt gern auf
bachbegleitenden Weiden (Salix sp.)
und Esche (Fraxinus excelsior). Der
von Schauer (1963) angegebene Fund
vom Langbathsee am Nordabfall des
Höllengebriges in Oberösterreich konnte
im Zuge neuerer Begehungen zwischen
1975 und 1996 nicht mehr bestätig werden.
Heterodermia obscurata gehört zu den vom
Aussterben bedrohten Arten in Österreich
(Gefährdungsstufe 1).
HETERODERMIA SPECIOSA (Wulfen) Trevis. (Abb. 4)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; Türk & Breuss, 1994; Rücker &
Wittmann, 1995; Pfefferkorn, 1996; Türk,
1996; Pfefferkorn & Türk, 1997.
Heterodermia speciosa siedelt auf
bemoosten Stämmen und auf Seitenästen
von Laubbäumen (Salix sp., Alnus incana,
Acer pseudoplatanus, Fagus sylvatica
etc.), in extrem luftfeuchten Orten
(enge Talschluchten, schattige Täler,
Nähe von Wasserfällen) kommt sie auch
auf Nadelbäumen vor. Als ozeanische
Flechte bevorzugt sie Gebiete mit hohen
Niederschlagsraten von der montanen (ab
etwa 400 Meter) bis in die hochmontane
(etwa 1400, im Extremfall 1600 Meter)
Stufe. Sie fruchtet selten – und wenn,
dann zumeist an bachnahen Stämmen und
Ästen. Ein geschlossenes Verbreitungsgebiet
weist sie in den Tälern der Nördlichen
Kalkalpen auf, in den Zentralalpen tritt sie
nur an sehr luftfeuchten Standorten auf.
In den südlichen Kalkalpen bevorzugt sie
die niederschlagsreichen Abschnitte der
Karnischen Alpen und der Karawanken.
Außerhalb der Alpen dürfte sie heute in
Österreich ausgestorben sein. Über die
Verbreitung in Europa siehe Schauer
(1965).
HYPERPHYSCIA ADGLUTINATA (Flörke) H. Mayrhofer &
Poelt, syn.: Physciopsis a. (Flörke) M. Choisy
(Abb. 5)
Literatur: Türk & Poelt, 1993; Hafellner &
Maurer, 1994; Türk et al., 1994; Hofmann
et al., 1995; Obermayer, 1996.
Die tropische Gattung Hyperphyscia
ist in Europa lediglich mit einer Art –
Hyperphyscia adglutinata – vertreten. Diese
Flechte ist wohl eine der kleinstwüchsigen
Blattflechten unserer heimischen Flora. Ihr
graugrün bis graubraun gefärbtes Lager liegt
dem Substrat sehr eng an. Sie bevorzugt
subneutrale bis staubimprägnierte Borke
von freistehenden Laubbäumen (Aesculus
hippocastanum, Robinia pseudacacia, Tilia
cordata, Populus sp., Juglans regia u.a.)
in wärmegetönten Lagen außerhalb der
Alpen. In den Alpen folgt sie gelegentlich
tiefer gelegenen Flußtälern. Sie bevorzugt
die kolline (ab 180 Meter) bis niedermontane
(ca. 700 Meter) Stufe. Aus Vorarlberg,
Tirol und Kärnten liegen bisher keine
Fundmeldungen vor. In Europa ist sie
mitteleuropäisch-mediterran verbreitet.
PHAEOPHYSCIA CERNOHORSKYI (Nádv.) Essl., incl.
P. strigosa (Poelt & Buschardt) Golubkova
(Abb. 6)
Literatur: Türk & Poelt, 1993; Hofmann et
al., 1995.
Phaeophyscia cernohorskyi bevorzugt
ähnlich wie Ph. insignis südeponierte,
trockenwarme Standorte. Neben ihrem
gehäuften Auftreten in collinen bis
submontanen Lagen des südöstlichen
Alpenrandes stößt sie inneralpin (unter
geeigneten kleinklimatischen Bedingungen)
auch bis in hochmontane Lagen vor. Sie
wächst überwiegend auf kalkhältigen
Gesteinen oder darüberliegenden Moosen.
P HAEOPHYSCIA CHLOANTHA (Ach.) Moberg, syn.:
Physcia luganensis Mereschk.; Physciella
chloantha (Ach.) Essl. (Abb. 7)
138 Folia Cryptog. Estonica
Abb. 4. Bisher bekannte Verbreitung von Heterodermia speciosa in Österreich (Kreuz: nachweiselich
ausgestorben; Kreis: Fund vor 1900).
Abb. 5. Bisher bekannte Verbreitung von Hyperphyscia adglutinata in Österreich.
Abb. 6. Bisher bekannte Verbreitung von Phaeophyscia cernohorskyi in Österreich.
139
Abb. 7. Bisher bekannte Verbreitung von Phaeophyscia chloantha in Österreich.
Abb. 8. Bisher bekannte Verbreitung von Phaeophyscia ciliata in Österreich (Kreis: Fund vor 1900).
Abb. 9. Bisher bekannte Verbreitung von Phaeophyscia constipata in Österreich.
140 Folia Cryptog. Estonica
Literatur: Türk & Poelt, 1993; Obermayer,
1993; Hafellner & Maurer, 1994; Türk et
al., 1994; Berger, 1996; Boom et al., 1996;
Pfefferkorn, 1996; Pfefferkorn & Türk,
1997.
Phaeophyscia chloantha tritt in Österreich
nur in geringer Abundanz auf, die aktuellen
Funde betreffen zumeist nur einige wenige
Exemplare. Sie wächst in kollinen bis
submontanen, selten in montanen,
wärmegtönten Lagen auf Laubbäumen
mit nährstoffreicher Borke. Ihr höchst
gelegener Fundpunkt liegt in 1000 Meter
Höhe (Gaisberg bei Salzburg).
P HAEOPHYSCIA CILIATA (Hoffm.) Moberg, syn.:
Physcia c. (Hoffm) Du Rietz (Abb. 8)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; Obermayer, 1993; Türk & Breuss,
1994; Hofmann et al., 1995; Rücker
& Wittmann, 1995; Boom et al., 1996;
Pfefferkorn, 1996; Pfefferkorn & Türk,
1997.
Phaeophyscia
ciliata
hat
hohe
Ansprüche an die Feuchte, sie bevorzugt
niederschlagsreiche Lagen von der
montanen bis hochmontanen Stufe, wo
sie auf Laubbäumen wächst. Gelegentlich
wächst sie im Kronentrauf von Bäumen
auch auf Holzzäunen. Ihr Vorkommen
beschränkt sich in Österreich auf den
Alpenraum.
PHAEOPHYSCIA CONSTIPATA (Norrl. & Nyl.) Moberg
(Abb. 9)
Literatur: Türk & Poelt, 1993; Hofmann et
al., 1995.
Phaeophyscia constipata ist eine der
wenigen bodenbewohnenden Vertreter ihrer
Gattung. Sie wächst auf sehr flachgründigen
Kalk- und Dolomitböden, zumeist über
Felsen, mit Moosen vergesellschaftet. Ihre
Wuchsorte zeichnen sich durch einen
hohen Strahlungsgenuß aus, weshalb sie
in Österreich die regenarmen, inneralpinen
Täler in hochmontanen Lagen bevorzugt.
Lediglich in den Nördlichen Kalkalpen
kommt sie auch in niederschlagsreicheren
Regionen vor (Schober bei Thalgau, in 1300
msm). Sie gehört zu den Seltenheiten der
heimischen Flechtenflora.
PHAEOPHYSCIA ENDOCOCCINA (Körb.) Moberg, incl.
var. LITHOTODES Nyl. (Abb. 10)
Literatur: Türk & Poelt, 1993; Obermayer,
1993; Pils & Berger, 1995; Hofmann et al.,
1995; Berger, 1996.
Diese Flechte siedelt bevorzugt auf nicht zu
sauren, silikatischen Gesteinen. Hinsichtlich
ihres Feuchtebedürfnisses scheint sie eine
sehr weite ökologische Potenz zu haben:
sie wächst in der amphibischen Zone
klarerBäche, auf zeitweise überrieselten,
stark geneigten Felsflächen oder auf
überhängenden Felsflächen ohne jede
Zufuhr von tropfbar flüssigem Wasser.
Auch auf Moosen ist sie anzutreffen. In
Österreich ist ihr Hauptverbreitungsgebiet
in den Alpen, wo sie von der montanen bis
in die alpine Stufe vorkommt. Außerhalb der
Alpen ist sie sehr selten.
PHAEOPHYSCIA ENDOPHOENICEA (Harm.) Moberg (Abb.
11)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; 1996; Neuwirth & Türk, 1993;
Obermayer, 1993; Pfefferkorn & Türk,
1993; Berger & Türk, 1994; 1995; Hafellner
& Maurer, 1994; Türk & Breuss, 1994; Türk
et al., 1994; Rücker & Wittmann, 1995;
Baumgartner & Türk, 1996; Boom et al.,
1996; Hafellner et al., 1996; Pfefferkorn
1996.
Phaeophyscia endophoenicea sucht von
vornherein feuchtere Habitate auf, die
durch hohe Niederschläge oder große
Nebelhäufigkeit ausgezeichnet sind.
Zumeist wächst sie zwischen Moosen, was
für ihre Wasserversorgung günstig ist. In
Österreich kommt sie von der kollinenbis
zur hochmontanen (höchster Fundpunkt
in 1600 Meter) Stufe vor, sie meidet kalte
Regionen, weshalb sie im Gebiet der
Böhmischen Masse (Mühl und Waldviertel)
nur an wärmebegünstigten Wuchsorten
gedeiht.
PHAEOPHYSCIA HIRSUTA (Mereschk.) Essl. (Abb. 12)
Literatur: Türk & Poelt, 1993; Hofmann, 1993;
Türk & Breuss, 1994; Boom et al., 1996.
Phaeophyscia hirsuta ist hauptsächlich
südeuropäisch verbreitet, lediglich an
warmen Standorten dringt sie bis in das
südliche Mitteleuropa vor. Sie wächst auf der
subneutralen bis schwach sauren Borke von
Laubbäumen, die gelegentlich auch leicht
eutrophiert sein kann und auf Gestein. Die
Zentralalpen Österreichs meidet sie.
141
PHAEOPHYSCIA HISPIDULA (Ach.) Ess.(Abb. 13)
Literatur: Türk & Poelt, 1993; Berger, 1996;
Hafellner, 1997.
Sie gehört zu den absoluten Seltenheiten
der heimischen Flechtenflora. Diese
of fensichtlich
tropisch-ozeanische
Flechte kommt in den Bergwäldern des
Mittelmeergebietes zerstreut vor (Poelt,
1969) und dringt nur an sehr trockenen,
niederschlagsarmen Standorten in die
Alpen vor. Außerhalb der Alpen wurde sie
im Donautal aufgefunden.
PHAEOPHYSCIA INSIGNIS (Mereschk.) Moberg (Abb.
14)
Literatur: Türk & Poelt, 1993; Hafellner,
1997.
Die Phaeophyscia orbicularis sehr
nahe stehende Sippe mit ihrem
Verbreitungsschwerpunkt in Südeuropa
ist
in
Österreich
großteils
auf
wärmeexponiertere, colline bis montane
Lagen beschränkt. Vereinzelt dringt sie auch
bis in die montane Stufe vor. Phaeophyscia
insignis siedelt zumeist an Allebäumen,
geht aber auch auf (kalkstaubimprägnierte)
Moose über.
PHAEOPHYSCIA KAIRAMOI (Vain.) Moberg (Abb. 15)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; Hofmann et al., 1995.
Phaeophyscia kairamoi wächst sowohl
epiphytisch auf Laubbäumen als auch
auf bemoosten Kalkfelsen. Sie kommt von
der montanen bis in die alpine Stufe vor.
Hinsichtlich der Feuchteanprüche dürfte
sie eine sehr weite Amplitude besitzen. In
Österreich ist ihr Vorkommen auf die Alpen
beschränkt.
PHAEOPHYSCIA LABRATA (Mereschk.) ined., syn.:
Physcia labrata Mereschk. (Abb. 16)
Literatur: Türk & Poelt, 1993; Baumgartner
& Türk, 1996.
Phaeophyscia labrata wächst auf der Borke
von alten Laubbäumen in Bergwäldern.
Ihre hohen Ansprüche auf Feuchtigkeit
beschränkt ihr Vorkommen auf ozeanische
Lagen (vgl. Schauer, 1965) in der montanen
bis hochmontanen Stufe. Sie fehlt nördlich
der Donau und in den kontinental geprägten
östlichen Landschaften Österreichs.
PHAEOPHYSCIA NADVORNIKII (Frey & Poelt) N. S.
Golubkova, syn.: Physcia n. Frey & Poelt
(Abb. 17)
Literatur: Türk & Poelt, 1993; Türk, 1996.
Sie ist in Österreich sehr selten. Sie wächst
auf Borke am Stamm und an Seitenästen
von alten Laub- und Nadelbäumen sowie auf
Felsen in sehr feuchten Lagen der montanen
bis hochmontanen Stufe. Im Spritzbereich
von Wasserfällen kann sie Massenbestände
bilden (vgl. Schauer, 1965)
PHAEOPHYSCIA NIGRICANS (Flörke) Moberg (Abb. 18)
Literatur: Türk & Poelt, 1993; Hafellner,
1993; Ober mayer, 1993; Hafellner &
Maurer, 1994; Türk & Breuss, 1994; Türk
et al., 1994; Berger & Türk, 1995; Boom et
al., 1996.
Sie kommt in Österreich von der kollinen
bis in die hochmontane Stufe (1600 msm)
vor. Sie wächst auf eutrophierten, zumeist
kalkhaltigen Gesteinsoberflächen, auf Mörtel,
Beton und auchauf staubimprägnierten
Baumborken am Stammgrund, wo sie
leicht zu übersehen ist. Sie bevorzugt
etwas trockenere, wärmegetönte Lagen. Im
Voralpenland und in den Mittelgebirgen des
Mühl- und Waldviertels tritt sie häufiger auf,
in den Alpen ist sie selten.
PHAEOPHYSCIA ORBICULARIS (Neck.) Moberg, syn.:
Physcia virella (Ach.) Flagey (Abb. 19)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; 1994; Neuwirth & Türk, 1993;
Obermayer, 1993; Pfefferkorn & Türk,
1993; Hafellner & Maurer, 1994; Türk &
Breuss, 1994; Türk et al., 1994; Berger &
Türk, 1995; Hofmann et al., 1995; Rücker
& Wittmann, 1995; Baumgartner & Türk,
1996; Berger, 1996; Boom et al., 1996;
Hafellner et al., 1996; Hafellner & Wittmann,
1996; Pfefferkorn, 1996; Türk, 1996.
Phaeophyscia orbicularis ist der häufigst
auftretende Vertreter ihrer Gattung. Sie
besiedelt alle möglichen Substrate, so sie
genügend eutrophiert sind, seien es nun
Baumborken oder Gesteinsoberflächen.
Auf anthropogenen Substraten kann sie
Massenbestände bilden, in Gebieten mit
hohem Grad an Lufverunreinigungen
ist sie oftmals der einzige Vertreter der
Blattflechten. Sie kommt von der kollinen
bis in die alpine Stufe vor, wo sie an gut
gedüngten Plätzen über Pflanzenresten und
auf Gesteinssubstraten wächst. Die leeren
Grundfelder lassen vermuten, daß sie dort
142 Folia Cryptog. Estonica
Abb. 10. Bisher bekannte Verbreitung von Phaeophyscia endococcina in Österreich.
Abb. 11. Bisher bekannte Verbreitung von Phaeophyscia endophoenicea in Österreich.
Abb. 12. Bisher bekannte Verbreitung von Phaeophyscia hirsuta in Österreich.
143
Abb. 13. Bisher bekannte Verbreitung von Phaeophyscia hispidula in Österreich.
Abb. 14. Bisher bekannte Verbreitung von Phaeophyscia insignis in Österreich.
Abb. 15. Bisher bekannte Verbreitung von Phaeophyscia kairamoi in Österreich.
144 Folia Cryptog. Estonica
Abb. 16. Bisher bekannte Verbreitung von Phaeophyscia labrata in Österreich.
Abb. 17. Bisher bekannte Verbreitung von Phaeophyscia nadvornikii in Österreich.
Abb. 18. Bisher bekannte Verbreitung von Phaeophyscia nigricans in Österreich.
145
Abb. 19. Bisher bekannte Verbreitung von Phaeophyscia orbicularis in Österreich.
Abb. 20. Bisher bekannte Verbreitung von Phaeophyscia pusilloides in Österreich.
Abb. 21. Bisher bekannte Verbreitung von Phaeophyscia sciastra in Österreich.
146 Folia Cryptog. Estonica
Abb. 22. Bisher bekannte Verbreitung von Phaeophyscia stiriaca in Österreich.
übersehen oder bisher noch nicht registriert
wurde.
PHAEOPHYSCIA PUSILLOIDES (Zahlbr.) Essl. (Abb. 20)
Literatur: Türk & Poelt, 1993; Hofmann,
1993; Boom et al., 1996; Pfefferkorn, 1996;
Pfefferkorn & Türk, 1997; Hafellner, 1997.
Phaeophyscia pusilloides ist eine typische
Flechte alter, weitgehend naturnaher
Waldbestände mit dem entsprechenden
Bestandesklima in ozeanischg getönten
Lagen. Zur Hauptsache ist sie in den Alpen
von der montanen bis zur hochmontanen
Stufe verbreitet, wo sie die Borke von
Laubbäumen (Acer pseudoplatanus, Fagus
sylvatica, Fraxinus excelsior, Alnus incana,
Salix sp. etc.) besiedelt. Außerhalb der Alpen
ist sie äußerst selten.
PHAEOPHYSCIA SCIASTRA (Ach.) Moberg (Abb. 21)
Literatur: Türk & Poelt, 1993; Obermayer,
1993; Hafellner & Maurer, 1994; Türk &
Breuss, 1994; Hafellner & Türk, 1995;
Hofmann et al., 1995; Berger, 1996;
Hafellner et al., 1996.
Phaeophyscia sciastra besiedelt eutrophierte
Substrate wie Gesteinsoberflächen (Kalk
und basiche Silikate), Holzschindeln,
Dächer und Holzzäune. Äußerst selten tritt
sie epiphytisch am staubimprägnierten
Stammgrund von Laubbäumen auf. Im
Gebirge wächst sie in höheren Lagen gern
über gesteins- und holzbewohnenden
Moosen. Diese morphologisch sehr
variable Art kann leicht übersehen werden.
Ein geschlossenes Verbreitungsgebiet in
Österreich weist sie im Alpenraum auf, im
übrigen Österreich tritt sie zerstreut auf.
PHAEOPHYSCIA STIRIACA (Poelt) Clauzade & Roux
(Abb. 22)
Literatur: Türk & Poelt, 1993.
Diese Flechte ist submediterran und
mediterran verbreitet, an einigen wenigen
Stellen dringt sie in die Steiermark vor.
Sie wächst auf Boden, Gestein und über
bodenbewohnenden Moosen.
DANK
Wir danken Herrn Professor Dr Josef Hafellner
(Graz) für das Bereitstellen von Funddaten und
Hinweise auf Literatur.
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Biosystematics and Ecology 3: 1–168.
Türk, R., Wittmann, H., Roth, S. & Wögerer, I. 1994.
Die Luftqualität im Stadtgebiet von Linz – Untersuchungen über den epiphytischen Flechtenbewuchs im Bezug zur Schadstoffbelastung. Jahrb.
Stadt Linz 37–39: 457–490.
148 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 32: 149–152 (1998)
Biogeographical data on the lichen flora of Finland
Orvo Vitikainen
Botanical Museum (Mycology), P.O. Box 47, FIN-00014 University of Helsinki, Finland
Abstract: From data of the checklist of lichens and allied fungi of Finland, comprising 1624 taxa (1422 species of lichens,
120 species lichenicolous and 44 species allied non-lichenized fungi), examples of regional species richness and some other
statistical values are presented on the basis of their occurrence in the traditional 21 biogeographical provinces. Ca. 10% of
the taxa have been recorded in all provinces, but 22.6% are recorded only in one. Some 15.6% of the accepted taxa have
their nomenclatural type from Finland.
Kokkuvõte: O. Vitikainen. Biogeograafilised andmed Soome lihhenofloora kohta.
Soome samblike ja neile lähedaste seente nimekirja põhjal , mis sisaldab 1624 taksonit (1422 liiki samblikke, 120 liiki
parasümbiontseid seeni ning 44 liiki mittelihheniseerunud seeni), esitatakse näiteid regionaalsest liigirikkusest ning teisi
statistilisi andmeid, mis põhinevad samblike levikul Soome 21 traditsioonilises biogeograafilises provintsis. Umbes 10%
taksonitest on teada kõigist provintsidest, kuid 22.6% esinevad ainult ühes. 15.6%-l tunnustatud taksonitest pärineb nende
nomenklatuurne tüüpmaterjal Soomest.
INTRODUCTION
One and a half century of lichen floristics in
Finland has accumulated much information,
both records in literature and materials in herbaria (see overview by Vitikainen, 1996). The
spatial and temporal continuum of these studies, however, is rather tenuous when compared
to the situation in countries such as Sweden
and England. The lack of a national checklist
of lichens of Finland for over one hundred years
is one example of evident deficiencies. Even the
recent checklist (Vitikainen et al., 1997) should
rather be regarded as a stimulus for additional
studies than a presentation of well-revised data.
However, the information compiled for the list
may be used for evaluating some aspects of the
lichen flora and the status of its investigation,
even if this treatment is not comparable in accuracy with the information on vascular plant
flora of Finland given by Lahti and al. (1988), nor
does it attempt to analyze the flora elements in
the same degree as Trass (1970), for example.
RESULTS AND DISCUSSION
The Finnish checklist contains 1 624 “accepted”
taxa, of which 1 458 are considered lichenized,
comprising 1422 species as well as additional
15 subspecies and 21 varieties, and 166 either
lichenicolous (120 species and 2 varieties) or
allied (44 species) fungi, both latter categories
being very poorly studied in Finland. In comparison, the checklist of the lichens of Sweden
and Norway (Santesson, 1993) recognizes 2271
accepted species, and in the checklist of Great
Britain and Ireland (Purvis et al., 1993) 1660
species with 61 additional infraspecific taxa
are accepted. 395 of the lichenized species and
9 subspecies plus 11 varieties of the Finnish
checklist are so called macrolichens. In comparison, in the recent macrolichen flora of Estonia
332 species are recognized (Trass & Randlane,
1994), but this area is much smaller and less
variable geographically.
These Finnish figures, even if indicated as
“accepted”, still comprise several imperfectly
known or critical taxa, many of which have been
described from Finnish materials but not subjected to investigation by modern methods, and
considerable changes and amendments of the
list are anticipated with progressing taxonomical
studies and checking of herbarium materials.
For example, there was an increase of 261 species in ten years in the Swedish and Norwegian
checklist, and almost one hundred species were
added in 13 years to the list of the British Isles
(see Santesson, 1993; Purvis et al., 1993).
150 Folia Cryptog. Estonica
Numbers of taxa in the biogeographical
provinces
The numbers of taxa recorded in the 21 biogeographic provinces of Finland (Fig. 1) are presented in Table 1. It should be noted that a low
number of provincial records, especially when
compared with neighbouring provinces, often
indicates less intensive research activity and not
actual adverse conditions for lichen growth.
The provinces South Häme, Varsinais-Suomi
and Uusimaa, with the highest numbers of recorded species – close or over 50 % of the total
– are well-known as centres of activity since the
early periods of lichen floristics in Finland; in
addition, these areas also provide a richer range
of habitats than the adjacent provinces.
The lowest figure, 193 recorded species for
Ladoga Karelia, is explained by the situation
that this area is only represented by a small and
poorly studied corner of the traditional province,
the main part belonging now to Russia. Other
areas of less intensive recording than the average, e.g. South Karelia, South Savo, Middle
and Oulu Ostrobothnia, Kittilä and Sompio
Lapland, with ca. 400 species at most, include
wide areas of rather monotonous and uniform
oligotrophic boreal forest vegetation and nature
with little geological variation and influence of
cultivation but additional research will certainly
increase the number of species. If the anomalous
Ladoga Karelia is omitted, 574 taxa are present
in each province on average, i.e the provinces
Åland and North Savo represent the average
number of taxa.
Patterns of species distribution
Finland, with its length of over 1 100 km from
south to north, provides a gradient of several
boreal vegetation zones, ranging from the hemiboreal zone in the three southernmost provinces
to hemiarctic areas prevailing in Lapland. Some
general observations of the species ranges along
this transection can be made, though the limitations of the material should be remembered.
Not less than 24.5 % of the taxa are recorded
only in one province, and even 11.8 % of the
Fig. 1. The biogeographical provinces of Finland,
their Finnish acronyms and names with their
English equivalents used in the text.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
A
V
U
EK
St
EH
ES
LK
EP
PH
PS
PK
PK
Kn
OP
PeP
Ks
KiL
SoL
EnL
InL
Ahvenanmaa
Varsinais-Suomi
Uusimaa
Etelä-Karjala
Satakunta
Etelä-Häme
Etelä-Savo
Laatokan Karjala
Etelä-Pohjanmaa
Pohjois-Häme
Pohjois-Savo
Pohjois-Karjala
Keski-Pohjanmaa
Kainuu
Oulun Pohjanmaa
Perä-Pohjanmaa
Koillismaa
Kittilän Lappi
Sompion Lappi
Enontekiön Lappi
Inarin Lappi
Åland
South Karelia
South Häme
South Savo
Ladoga Karelia
South Ostrobothnia
North Häme
North Savo
North Karelia
Middle Ostrobothnia
Oulu Ostrobothnia
Outer Ostrobothnia
Kittilä Lapland
Sompio Lapland
Enontekiö Lapland
Inari Lapland
151
species are recorded in two provinces only.
Only some 6.7 % of the total number of species
have been recorded in all provinces, even if not
in the same degree of abundance or frequency
throughout the range. Such species include
Arctoparmelia centrifuga, Cetraria sepincola,
Cladina arbuscula, C. stellaris, Evernia prunastri, Hypogymnia physodes, Parmelia saxatilis
and Peltigera aphthosa, among others. With
exclusion of Ladoga Karelia the percentage of
such widely occurring species is still close to 10
%. This relatively low figure is understandable
bearing in mind those underexplored provinces
discussed above.
Table 1. The numbers of taxa (A) in the biogeographical provinces of Finland, their percentage
of the total (B), and percentage of taxa recorded
in only one province (C)
Province
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
A
V
U
EK
St
EH
ES
LK
EP
PH
PS
PK
KP
Kn
OP
PeP
Ks
KiL
SoL
EnL
InL
A
B%
C%
577
869
788
385
599
907
492
193
460
525
571
592
405
533
389
625
692
424
371
630
638
35.6
53.6
48.6
23.8
37.0
56.0
30.4
11.9
28.4
32.4
35.2
36.5
25.0
32.9
24.0
38.6
42.7
26.2
22.9
38.9
39.4
2.3
7.0
2.9
–
1.0
9.2
0.6
–
0.2
0.8
1.8
2.2
0.3
1.9
0.5
2.9
4.3
2.6
0.8
10.5
6.3
Southern and western elements
Some 9 % of the total number of taxa are recorded
only in the area comprising the three southwesternmost provinces Åland, Varsinais-Suomi and
Uusimaa; for vascular plants the corresponding
percentage is 15 (see Lahti et al., 1988). Species
recorded in all of these three provinces include
Acrocordia gemmata, Caloplaca microthallina,
Cladina ciliata, C. portentosa, Cladonia foliacea
and Physcia leptalea, for instance. South Karelia
provides only five additional taxa to this “southern” or southwestern element.
A few marine species have their area extended
along the coastal provinces of Gulf of Bothnia
until Satakunta (Ramalina siliquosa) and South
Ostrobothnia (so e.g. Anaptychia ciliaris var.
melanosticta and Physcia tenella var. marina)
or even to Outer Ostrobothnia like Caloplaca
scopularis. A western pattern is represented
also by Cladonia strepsilis, Lecanora intumescens, Ramalina baltica and R. subfarinacea,
but these species extend their ranges farther
off inland.
The reverse, an eastern distribution pattern is
less obvious but may be seen in the ranges of
e.g. Hypocenomyce anthracophila, Menegazzia
terebrata and Tuckermannopsis ciliaris.
A major decrease in the number of “southern”
species takes place in the inland province South
Häme with its great number of “endemics”, so
that 18 % of the total number of taxa have not
been recorded north of this province. A notable
decline of 5.1 % after Outer Ostrobothnia and of
7.7 % after Koillismaa, with additional decrease
in Kittilä and Sompio Lapland, results in that
ca. 50 % of the species have not been recorded
in the northernmost provinces Enontekiö and
Inari Lapland.
The province Åland (A) shares some 91 % of its
taxa with Varsinais-Suomi (V), whereas some
39,4 % of the taxa present in V have not been
recorded from A. This may partly be explained by
the different size and digressing natural conditions of these areas but is more likely due to the
less intense of study of Åland lichens, which is
also seen in the lower figure of taxa recorded in
this province and the relatively high number of
taxa recorded in V only.
Comparison of the records of Åland – representing the hemiboreal zone – with some other
areas shows that this province shares 63 % of
its taxa with Koillismaa, in the northern boreal
zone, and 50 % of its species are also found
in Enontekiö Lapland or some 52.5 % in Inari
152 Folia Cryptog. Estonica
Lapland, these two northernmost provinces
representing orohemiarctic and even oroarctic
areas. 46 % of the species found in VarsinaisSuomi are shared with Inari Lapland, respectively; however, the number of species shared
by Varsinais-Suomi and Inari Lapland is higher
than in the case of Åland because of the higher
number of species found in Varsinais-Suomi.
the areas of research by taxonomists like William Nylander, Edvard Vainio and Veli Räsänen:
South Häme leading with 174 described taxa,
of which 104 are “accepted”, then Uusimaa (59
v. 24 taxa, respectively), Varsinais-Suomi (43 v.
17), Inari Lapland (38 v. 22), Outer Ostrobothnia (31 v. 9), Enontekiö Lapland (30 v. 15) and
Koillismaa (29 v. 24).
Northern pattern of range
Not surprisingly, E. Acharius stands as the
describing author of some 200 accepted taxa,
but the most proliferous author proves to be W.
Nylander (some 260 taxa plus about ten taxa
validated by others but credited to Nylander).
Next comes E. Vainio with about 100 taxa, then
C. von Linné, who has named the epithet of 92
taxa, and Th. Fries, with 55 taxa. These five
Nordic authors have described 43.7 % of the
lichens and allied fungi of Finland.
As to the “northern” element, species only
recorded in the two northernmost provinces
Enontekiö and Inari Lapland comprise 8 % of
the total (19 % for vascular plants, see Lahti
et al., 1988). For example, Dactylina ramulosa,
Lecanora frustulosa, Solorina bispora and S. octospora are known only in Enontekiö Lapland,
and Siphula ceratites only in Inari Lapland,
whereas Lobaria linita, Rhizoplaca chrysophthalma, R. melanophthalma and Stereocaulon
spathuliferum have been recorded in both. 14
% of the lichens of Finland have not been found
south of Koillismaa (15 % of vascular plants!).
The respective figure for Middle Ostrobothnia
– in the middle boreal zone – is 18.2 %, for
South Ostrobothnia 23 %, and 1/3 of the taxa
have their total recorded range in or north of
South Häme.
Who described the Finnish lichens?
An attempt was made in the checklist to indicate, if the nomenclatural type of a taxon was
known to be from a Finnish province. At least
257 accepted taxa (15.6 %) and 284 additional
synonymous names have their type from Finland, these listings certainly not being complete.
The highest numbers of types emerge in those
biogeographical provinces, where floristic activity on lichens has been intensive, and display
REFERENCES
Lahti, T., Kurtto, A. & Väisänen, R. A. 1988. Floristic
composition and regional species richness of
vascular plants in Finland. Ann. Bot. Fenn. 25:
281–291.
Purvis, O., Coppins, B. J. & James, P. W. 1993. Checklist of lichens of Great Britain and Ireland. Bull.
British Lichen Soc. (Suppl.) 72: 1–75.
Santesson, R. The lichens and lichenicolous fungi of
Sweden and Norway. SBT-förlaget, Lund. 240
pp.
Trass, H. 1970. The elements and development of the
lichen-flora of Estonia (in Russian). Pap. Bot. 9:
5–233.
Trass, H. & Randlane, T. (eds.) 1994. Eesti suursamblikud. Tartu. 399 pp.
Vitikainen, O. 1996. Lichen floristics in Finland. Mem.
Soc. Fauna Fl. Fenn. 72: 213–218.
Vitikainen, O., Ahti, T., Kuusinen, M., Lommi, S. &
Ulvinen, T. 1997. Checklist of lichens and allied
fungi of Finland. Norrlinia 6: 1–123.
Folia Cryptog. Estonica, Fasc. 32: 153–159 (1998)
Lichens and lichenicolous fungi from the north of Pyasino
lake,Taimyr peninsula, Siberia
Mikhail Zhurbenko
Lab. Systematics & Geography of Fungi, Komarov Botanical Institute of the Russian Academy of Sciences,
2 Prof. Popov St., 197376 St.-Petersburg, Russia
Abstract: A total of 152 species of lichens and 16 species of lichenicolous fungi is reported from the area of southern
tundras north of Pyasino Lake on the Taimyr Peninsula. Bachmanniomyces uncialicola, Bispora christiansenii, Epicladonia sandstedei,
and Taeniolella pertusariicola are new to Russia; Acarospora nitrophila, Lecanora chlarotera, Pilophorus dovrensis, and Protoparmelia
picea are new to the Russian Arctic; Buellia epigaea, Dactylospora attendenda, Lecanora leptacina, Pertusaria carneopallida, Polyblastia
fuscoargillacea, Sagediopsis campsteriana, and Taeniolella beschiana are new to the north of Central Siberia; Amygdalaria elegantior, A.
panaeola, Bacidia herbarum, Lecania cyrtella, Rinodina archaea, and R. lecideoides are new to Taimyr.
Kokkuvõte: Mihhail Zhurbenko. Põhja pool Pjasino järve (Taimõri ps., Siber) kasvavad samblikud ja parasümbiontsed seened.
Taimõri poolsaarel Pjasino järvest põhja pool paikneva lõunatundra liikide nimekiri sisaldab 152 samblikuliiki ja 16 parasümbiontset seeneliiki. Bachmanniomyces uncialicola, Bispora christiansenii, Epicladonia sandstedei jaTaeniolella pertusariicola on uued
liigid Venemaa floristilises nimestikus; Acarospora nitrophila, Lecanora chlarotera, Pilophorus dovrensis ja Protoparmelia picea on uued
liigid Vene Arktikas; Buellia epigaea, Dactylospora attendenda, Lecanora leptacina, Pertusaria carneopallida, Polyblastia fuscoargillacea,
Sagediopsis campsteriana ja Taeniolella beschiana on uued liigid Kesk-Siberi põhjaosas; Amygdalaria elegantior, A. panaeola, Bacidia
herbarum, Lecania cyrtella, Rinodina archaea ja R. lecideoides on uued liigid Taimõri poolsaarel.
INTRODUCTION
Although about 38 local lichen floras have
been studied from the Taimyr Peninsula and
neighbouring islands (Zhurbenko 1996), lichens
of its southern tundras have been heretofore
much neglected. Lichens of southern tundras
of Taimyr have been specially treated only by
T. Piin (1984). There is also some information
about dominant lichens from southern tundras
in a few geobotanical papers (Avramchik, 1937;
Alexandrova, 1937; Matveeva & Zanokha, 1986;
Pospelova, 1974; Tyulina, 1937; Vinogradova,
1937). The present paper is a further contribution to the lichen flora of the area. It is a pleasure
to include it in a festschrift of Prof. Hans Trass,
an outstanding representative of Estonian lichenology, who has contributed much to the
knowledge of Taimyr lichens.
From 16 till 24 July 1983, the author collected lichens near the field research station
"Belyi Yar" of the Norilsk Far North Agricultural
Research Institute (70º05’N, 87º43’E). The study
area is located between the northern extremity of
Pyasino Lake, west of the mouth of the Pyasina
River and Ladanakh and Polovinnoe Lakes (Fig.
1). It is part of the northern Siberian Lowland.
The landscape is hilly plain with numerous lakes
and moraine deposits, like low crests and boulder fields. Altitudes range from 30 to 150 m. The
region is situated within the zone of continuous
permafrost and is influenced by the continental
Siberian climate (Atlas of the Arctic, 1985). Mean
annual temperature is approximately -12ºC with
the lowest mean monthly temperature approximately -30ºC in January and the highest, +12ºC
in July. The daily mean is above freezing from
the beginning of June up to the middle of September. Annual precipitation is about 300-400
mm. The dominant vegetation is classified as
southern or shrub tundra (Chernov & Matveeva,
1979) and is characterized by closed shrub vegetation in upland tundras. Here and there vegetation cover includes areas of forest tundras with
Larix sibirica Ledeb., typical tundras, meadows,
and bogs. Frequently dominating species are Alnus fruticosa Rupr., Betula nana L., Salix lanata
L., S. reptans Rupr., S. pulchra Cham., S. glauca
L., Dryas octopetala L., Vaccinium uliginosum
L., V. vitisidaea L., Cassiope tetragona (L.) D.
Don, Ledum decumbens (Ait.) Lodd. ex Steud.,
154 Folia Cryptog. Estonica
SPITZBERGEN
SEVERNAYA ZEMLYA
TAIMYR P.
NOVAYA ZEMLYA
Fig. 1. Location of study area in Taimyr peninsula, Siberia.
Empetrum subholarcticum V. Vassil., Salix polaris Wahlenb., Rubus chamaemorus L., Arctous
alpina (L.) Niedenzu, Eriophorum vaginatum L.,
Hedysarum hedysaroides (L.) Schinz & Thell.
(Matveeva & Zanokha, 1986).
MATERIAL AND METHODS
Lichens were collected at 19 sample plots, representing the following nine habitat types (their
numbers are cited in the list of taxa).
1 – exposed windswept areas of prominent
convexities; spotty lichen-bryophyte-dwarfshrub communities.
2 – slopes of terraces of Pyasino Lake in places
with open boulder fields; shrubs (Alnus fruticosa, Betula nana), meadow tundras.
3 – slopes of moraine crests and lake terraces in
places with boulder fields; lichen-bryophytedwarf-shrub communities.
4 – shore of Pyasino Lake; shrubs (Alnus fruticosa, Betula nana ).
5 – plain and gentle slopes; tussock and spotty
tundras.
6 – swampy plain with peat hillocks; lichendwarf-shrub-bryophyte communities.
7 – shores of lakes and streams with open sand
and pebbles.
8 – lower slopes of lake depressions; willow
shrubs.
9 – boulder beds on lower slopes of lake depressions.
Standard TLC methods were used for identification of some specimens of Cladonia and
Peltigera. Voucher specimens are deposited in
LE and H.
RESULTS AND DISCUSSION
The study resulted in a list of 152 species of
lichens and 15 species of lichenicolous fungi
which are representative for a typical local flora
of southern tundras of Taimyr. All lichen species
fall into wide-ranging circumpolar arctic-alpine,
arcto-boreal, or boreal distributional categories.
155
The list is no doubt incomplete, for instance a
nearby local flora from the vicinities of settlement Kresty Taimyrskie numbers 178 species
just of epigaeic lichens (Piin, 1984).
Recently a few publications summarizing
knowledge on lichens and lichenicolous fungi
of the Russian Arctic have been published. Andreev et al. (1996) report 1069 species of lichens
and nine species of lichenicolous fungi for the
Russian Arctic as a whole. Zhurbenko & Santesson (1996) report 74 species of lichenicolous
fungi for the Russian Arctic, and Zhurbenko
(1996) reports 660 species of lichens and 61
species of lichenicolous fungi for the north
of Central Siberia. To these lists we can now
add 20 species. Bachmanniomyces uncialicola,
Bispora christiansenii, Epicladonia sandstedei,
and Taeniolella pertusariicola are new to Russia; Acarospora nitrophila, Lecanora chlarotera,
Pilophorus dovrensis, and Protoparmelia picea
are new to the Russian Arctic; Buellia epigaea,
Dactylospora attendenda, Lecanora leptacina,
Pertusaria carneopallida, Polyblastia fuscoargillacea, Sagediopsis campsteriana, and Taeniolella
beschiana are new to the north of Central Siberia; Amygdalaria elegantior, A. panaeola,
Bacidia herbarum, Lecania cyrtella, Rinodina
archaea, and R. lecideoides are new to Taimyr.
Dactylospora purpurascens is reported for the
second time for Russia.
Annotated list of taxa
Annotations for the taxa include infor mation on: substrate; incidental notes; habitat
numbers, 1-9, in which the taxon occured; the
number of different plots, at 19 sample plots, in
which the taxon was noted (in parentheses). Lichenicolous fungi are marked with asterisks.
ACAROSPORA NITROPHILA H. Magn. – on pebbles:
7 (1).
ALECTORIA NIGRICANS (Ach.) Nyl. – over soil; common: 1, 2, 3 (4).
A. OCHROLEUCA (Hoffm.) A. Massal. – over soil, also
on driftwood; common: 1, 2, 3, 7 (6).
AMYGDALARIA ELEGANTIOR (H. Magn.) Hertel & Brodo
– on rock: 1 (1).
A. PANAEOLA (Ach.) Hertel – on rock: 1 (1).
ARCTOCETRARIA NIGRICASCENS (Nyl.) Kärnefelt & A.
Thell – on twigs of Betula nana: 4 (1).
ARCTOMIA DELICATULA Th. Fr. – on bryophytes: 9
(1).
ASAHINEA CHRYSANTHA (Tuck.) W. L. Culb. & C. F.
Culb. – on soil, also on rock: 1, 2, 3 (4).
*BACHMANNIOMYCES UNCIALICOLA (Zopf) D. Hawksw.
– on Cladonia pyxidata thallus: 5 (1).
BACIDIA BAGLIETTOANA (A. Massal. & De Not.) Jatta
– on decaying bark of Salix glauca: 7, 8
(2).
B. HERBARUM (Stizenb.) Arnold – on degenerating
thallus of Peltigera sp.: 5 (1).
BAEOMYCES PLACOPHYLLUS Ach. – on soil: 1, 3 (2).
B. RUFUS (Huds.) Rebent. – on soil: 1, 2, 9 (4).
BIATORA SUBDUPLEX (Nyl.) Printzen – on bryophytes
and plant remnants, on dead bark and naked lignum of Salix glauca; fairly common:
1, 3, 4, 7, 8, 9 (9).
*BISPORA CHRISTIANSENII D. Hawksw. – in apothecia
of Caloplaca cf. cerina, and Lecania cyrtella:
4 (1). Conidia 0–1, septate, 5–10 × 5–6 µm.
BRYOCAULON DIVERGENS (Ach.) Kärnefelt – over soil;
common: 1, 2, 3, 6 (6).
BRYONORA CASTANEA (Hepp) Poelt – on bryophytes,
soil and plant remnants; : 3, 9 (3).
BRYORIA NITIDULA (Th. Fr.) Brodo & D. Hawksw.
– over soil: 1, 3 (3).
BUELLIA EPIGAEA (Pers.) Tuck. – on bryophyte: 9
(1).
B. INSIGNIS (Hepp) Th. Fr. – on bryophyte: 9 (1).
CALOPLACA AMMIOSPILA (Ach.) H. Olivier – on bryophytes and bare driftwood: 1, 7 (2).
C. CERINA (Ehrh. ex Hedw.) Th. Fr. – on bryophytes
and plant remnants, including
degenerating thallus of Peltigera sp., on dead bark of
Alnus fruticosa; common: 1, 3, 4, 9 (5).
C. JUNGERMANNIAE (Vahl) Th. Fr. – on bryophytes
with earth: 7 (1).
C. TETRASPORA (Nyl.) H. Olivier – on bryophytes
and plant remnants, including
degenerating squamules of Cladonia pyxidata;
common: 3, 7, 9 (4).
C. TIROLIENSIS Zahlbr. – on dead bark of Alnus
fruticosa: 4 (1).
C. TORNOËNSIS H. Magn. – on saxicolous bryophytes: 3, 9 (2). Some specimens are
distinguished for there large spore size: 20–23
× 7-8 µm, septum – 1–2 µm.
CANDELARIELLA PLACODIZANS (Nyl.) H. Magn. – on
sandy soil: 1, 3 (2).
C. VITELLINA (Hoffm.) Müll. Arg. – on bryophytes,
naked driftwood and rocks: 1, 3, 7, 9 (4).
*CARBONEA VITELLINARIA (Nyl.) Hertel – on Candelariella cf. vitellina thallus: 1 (1).
CATAPYRENIUM CINEREUM (Pers.) Körb. – on bryophyte: 7 (1).
156 Folia Cryptog. Estonica
CETRARIA ACULEATA (Schreb.) Fr. – on soil: 1, 7 (3).
C. INERMIS (Nyl.) Krog – on Betula nana twigs:
8 (1).
C. ISLANDICA (L.) Ach. – on soil; abundant throughout the region: 1, 2, 3, 5, 6, 7, 8 (11).
C. LAEVIGATA Rass. – on soil; very common: 1, 2,
3, 5, 7, 8 (9).
C. ODONTELLA (Ach.) Ach. – on soil; common: 1,
2, 3, 7 (4).
CETRARIELLA DELISEI (Schaer.) Kärnefelt & A. Thell
– on soil; common and locally abundant in
humid sites: 3, 7, 9 (3).
CLADINA ARBUSCULA (Wallr.) Hale & W. L. Culb.
s. lat. – on soil; very common: 1, 2, 3, 6,
7 (7).
C. RANGIFERINA (L.) Nyl. – on soil; common: 1, 2,
3, 6 (6).
C. STELLARIS (Opiz) Brodo – on soil: 1, 3 (2).
C. STYGIA (Fr.) Ahti – on soil: 3 (Zhurbenko 8311)
(1); rev. T. Ahti, 1992.
CLADONIA ACUMINATA (Ach.) Norrl. – on degenerating
bryophytes: 7, 9 (2).
C. ALASKANA A. Evans – on soil: 1 ( Zhurbenko
835), 3, 7 (Zhurbenko 8316) (3); rev. T. Ahti,
1993 (H, LE). The species previously thought
to be restricted to Beringia, occurs sporadically at Taimyr and extends eastwards up to
the Polar Ural (Ahti & Zhurbenko, 1994).
C. AMAUROCRAEA (Flörke) Schaer. – on soil, also on
driftwood; very common: 1, 2, 3, 6, 7 (7).
C. BACILLIFORMIS (Nyl.) Glück – on soil: 5, 6 (2).
C. BOREALIS S. Stenroos – on soil: 9 (Zhurbenko
837) (1); rev. T. Ahti, 1992.
C. CARIOSA (Ach.) Spreng. – on soil: 7 (1).
C. CERVICORNIS (Ach.) Flot. ssp. VERTICILLATA (Hoffm.)
Ahti – on soil: 1, 3 (Zhurbenko 8317) (2); rev.
T. Ahti, 1992.
C. CHLOROPHAEA (Flörke ex Sommerf.) Spreng.
– on rotting lignum of Alnus fruticosa: 5
(Zhurbenko 831) (1); rev. T. Ahti, 1992.
C. COCCIFERA (L.) Willd. s. lat. – on soil; common:
1, 2, 3, 6, 9 (8).
C. CORNUTA (L.) Hoffm. – on soil and driftwood:
1, 7 (2).
C. CYANIPES (Sommerf.) Nyl. – on soil: 5 (Zhurbenko
8313) (1); rev. T. Ahti 1992.
C. cf. ECMOCYNA Leight. – on soil: 1 (1).
C. FIMBRIATA (L.) Fr. – on soil: 7 (Zhurbenko 832)
(1); rev. T. Ahti, 1992.
C. GRACILIS ssp. ELONGATA (Wulfen) Vain. – on soil:
3 (Zhurbenko 8315) (1); rev. T. Ahti, 1992.
Common, but formerly misunderstood by
the author.
C.
(Delise) Hav. – on soil: 7
(Zhurbenko 8312), 8 (Zhurbenko 8310) (2);
rev. T. Ahti, 1992. common, but formerly
misunderstood by the author.
C. MACROPHYLLA (Schaer.) Stenh. – on sandy soil
with silt: 9 (1).
C. PLEUROTA (Flörke) Schaer. – on soil: 6 (1).
C. POCILLUM (Ach.) Grognot – on soil; very common: 1, 2, 3, 5, 7, 8, 9 (10).
C. PYXIDATA (L.) Hoffm. – on soil and bryophytes;
very common: 1, 2, 3, 4, 5, 7, 8, 9 (11).
C. SQUAMOSA Hoffm. – on soil: 6 (1).
C. STRICTA (Nyl.) Nyl. – on soil: 9 (1).
C. SYMPHYCARPA (Flörke) Fr. – on soil: 7 (Zhurbenko
834) (1); rev. T. Ahti, 1992.
C. UNCIALIS (L.) Weber ex Wigg. – on soil: 1, 2,
3 (4).
DACTYLINA ARCTICA (M. J. Richardson) Nyl. – on
soil; common: 1, 2, 5, 7, 8 (7).
D. RAMULOSA (Hook.) Tuck. – on soil: 2 (1). Studied
specimens belong to P- chemical race.
*DACTYLOSPORA ATTENDENDA (Nyl.) Arnold – on Pilophorus dovrensis thallus: 5 (Zhurbenko
8395) (1); confirm. D. Triebel, 1997. Spores
brown, (1–3-septate, 12–15.5 × 5–6 µm.
*D. DEMINUTA – on Biatora subduplex, Lecanora
epibryon, Ochrolechia frigida, and Rinodina turfacea thalli: 1, 2, 5 (3); confirm. D.
Triebel, 1997. Spores brown with olive tinge,
3–7-septate, 15–22.5 × 5–6 µm.
*D. PURPURASCENS Triebel – on Amygdalaria panaeola thallus: 3 (Zhurbenko 83138) (1). Spores
8–15 × 5-6 µm, epithecium K+ purpur.
*D. sp. – on Cladonia pyxidata thallus: 7
(Zhurbenko 8381) (1). Spores brown, 1septate, 10.5–15 × 5–7 µm.
*ENDOCOCCUS RUGULOSUS Nyl. – on Aspicilia sp.
thallus: 3 (1).
*EPICLADONIA SANDSTEDEI (Zopf) D. Hawksw. – forms
galls on primary squamules of Cladonia
pyxidata: 5 (Zhurbenko 8350) (1). Conidia
hyaline, 1-septate, 10–12 × 3–4 µm.
*EPILICHEN SCABROSUS (Ach.) Clem. – on Baeomyces
placophyllus thallus: 1 (1).
FLAVOCETRARIA CUCULLATA (Bellardi) Kärnefelt & A.
Thell – on soil, also on driftwood; very common: 1, 2, 5, 6, 7, 8 (9).
F. NIVALIS (L.) Kärnefelt & A. Thell – on soil; very
common: 1, 2, 3, 5, 7, 8 (7). Specimen
Zhurbenko 8374 has round soralia, representing a sorediate form of the species.
HAFELLNERA PARASEMELLA (Nyl.) Houmeau & Cl.
Roux – on Betula nana twigs: 1, 2 (2).
MACROCERAS
157
HYPOGYMNIA AUSTERODES (Nyl.) Räsänen – on soil,
also on rock: 1, 2 (2).
H. PHYSODES (L.) Nyl. – on soil: 1, 2, 3 (3).
JAPEWIA TORNOËNSIS (Nyl.) Tønsberg – on plant
remnants: 1, 2 (2).
LECANIA CYRTELLA (Ach.) Th. Fr. – on dead bark
of Alnus fruticosa: 4 (1). Apothecia 0.2–0.4
mm, hymenium K+ (violet), spores 7.5–13 ×
2.5–3.5 µm, 0–1-septate.
LECANORA BOLIGERA (Norman ex Th. Fr.) Hedl. – on
dead bark of Alnus fruticosa: 4 (1).
L. CHLAROTERA Nyl. – on dead bark of Alnus fruticosa: 4 (1).
L. EPIBRYON (Ach.) Ach. – on bryophytes and plant
remnants, including dead parts of dwarf
shrubs; common: 1, 2, 3, 5, 9 (6).
L. HAGENII (Ach.) Ach. var. SAXIFRAGAE (Anzi) R.
Sant. – on degenerating bryophytes and
peat: 1, 6 (2).
L. LEPTACINA Sommerf. – on degenerating bryophytes and peat: 1, 6 (2).
L. POLYTROPA (Ehrh. ex Hoffm.) Rabenh. – on pebbles and rocks: 3, 7 (2).
LECIDEA RAMULOSA Th. Fr. – on bryophytes: 7 (1).
LECIDELLA EUPHOREA (Flörke) Hertel – on bryophytes, plant remnants, dead bark of Alnus fruticosa and Salix glauca, and naked
driftwood; common: 1, 2, 3, 4, 7, 8, 9 (7).
LECIOPHYSMA FINMARKICUM Th. Fr. – on degenerating
bryophytes: 1 (1).
LEPRARIA NEGLECTA (Nyl.) Lettau – on saxicolous
bryophytes: 3, 9 (2).
LOPADIUM CORALLOIDEUM (Nyl.) Lynge – on bryophytes and plant remnants: 2, 9 (2).
L. PEZIZOIDEUM (Ach.) Körb. – on bryophytes and
plant remnants; common: 1, 2, 3, 6 (5).
MEGASPORA VERRUCOSA (Ach.) Hafellner & V. Wirth
– on dead bryophytes: 3 (1).
MELANELIA DISJUNCTA (Erichsen) Essl. – on rock:
3 (1).
M. HEPATIZON (Ach.) A. Thell – on rock, also on
peat: 1 (1).
M. PANNIFORMIS (Nyl.) Essl. – on rock and soil:
1, 3 (3).
M. SEPTENTRIONALIS (Lynge) Essl. – on bark of Alnus
fruticosa: 4 (1).
M. SOREDIATA (Ach.) Goward & Ahti – on rocks:
3, 9 (2).
MYCOBILIMBIA BERENGERIANA (A. Massal.) Hafellner
& V. Wirth – on sandy soil with plant remnants: 3, 9 (2).
M. CARNEOPALLIDA (Müll. Arg.) Printzen – on bryophyte; common: 5 (1).
M. HYPNORUM (Lib.) Kalb & Hafellner – on bryophytes: 5 (1).
M. LOBULATA (Sommerf.) Hafellner – on degenerating bryophytes with sandy soil: 5, 7 (2).
NEPHROMA EXPALLIDUM (Nyl.) Nyl. – on soil: 1 (1).
OCHROLECHIA ANDROGYNA (Hoffm.) Arnold – on bryophytes, plant remnants, also on bark of
Alnus fruticosa; common: 1, 2, 5, 6, 9 (8).
O. FRIGIDA (Sw.) Lynge – on bryophytes and plant
remnants; common: 1, 2, 3, 9 (6).
O. GRIMMIAE Lynge – on bryophytes and degenerating cushion of graminoid: 2 (1).
O. cf. INAEQUATULA (Nyl.) Zahlbr. – on bryophytes
and lichens: 3 (2).
O. UPSALIENSIS (L.) A. Massal. – on bryophytes and
dead parts of dwarf shrubs: 1, 2, 3 (3).
PANNARIA PEZIZOIDES (Weber) Trevis. – on soil: 8, 9 (2).
P. cf. PRAETERMISSA Nyl. – on soil: 2 (1).
PARMELIA FRAUDANS (Nyl.) Nyl. – on rock and soil:
3 (2).
P. OMPHALODES (L.) Ach. – on soil: 1, 2, 3 (4).
PARMELIOPSIS AMBIGUA (Wulfen) Nyl. – on bark of
Alnus fruticosa: 4 (1).
P. HYPEROPTA (Ach.) Arnold. – on bark of Alnus
fruticosa and Salix sp.: 4, 8 (2).
PELTIGERA APHTHOSA (L.) Willd. – on soil; common:
1, 5, 8, 9 (5).
P. CANINA (L.) Willd. – on soil: 1, 5, 7 (3).
P. DIDACTYLA (With.) J. R. Laundon – on soil: 1, 5 (2).
P. LEPIDOPHORA (Nyl.) Bitter – on soil: 5 (1).
P. LEUCOPHLEBIA (Nyl.) Gyeln. – on soil: 5, 8 (2).
P. NECKERI Hepp ex Müll. Arg. – on soil: 1 (Zhurbenko
8333), 7 (Zhurbenko 8328), 8 (Zhurbenko
8326) (3); rev. O. Vitikainen, 1993.
P. RUFESCENS (Weiss) Humb. – on soil; common:
1, 3, 5, 7 (6).
PERTUSARIA BRYONTHA (Ach.) Nyl. – on bryophytes;
common: 1, 2, 3, 5 (6).
P. CARNEOPALLIDA – on bark of Alnus fruticosa and Salix glauca: 4, 8 (2); rev. I. I. Makarova, 1997.
P. CORIACEA (Th. Fr.) Th. Fr. – on bryophytes and
dead parts of dwarf shrubs: 1, 2, 3 (3).
P. DACTYLINA (Ach.) Nyl. – on bryophytes; rather
common: 1, 2, 3 (4).
P. GEMINIPARA (Th. Fr.) C. Knight – on bryophytes,
also on plant remnants with sand above
rock: 3, 9 (2).
P. GLOMERATA (Ach.) Schaer. – on degenerating
bryophytes with sandy soil: 1 (1).
P. PANYRGA (Ach.) A. Massal. – on bryophytes,
158 Folia Cryptog. Estonica
partly with sandy soil, also on dead parts of
dwarf shrubs; common: 1, 2, 3 (4).
PHAEOCALICIUM COMPRESSULUM (Nyl. ex Szatala) A.
F. W. Schmidt – on bark of Alnus fruticosa;
sparse: 4 (1).
*PHAEOSPOROBOLUS ALPINUS R. Sant., Alstrup & D.
Hawksw. – on Ochrolechia androgyna, O.
frigida, and Pertusaria coriacea thalli; frequently: 1, 2, 3 (3). One of the most common and noticeable lichenicolous fungi in
the Siberian Arctic.
PILOPHORUS DOVRENSIS (Nyl.) Timdal, Hertel & Rambold – on sandy soil: 5 (1).
PLACYNTHIELLA ULIGINOSA (Schrad.) Coppins & P.
James – on bryophytes, also on dead bark
of Alnus fruticosa: 1, 4 (2).
POLYBLASTIA FUSCOARGILLACEA Anzi – on rock: 7
(1).
P. SENDTNERI Kremp. – on plant remnants with
sandy soil: 7 (1).
POLYCHIDIUM MUSCICOLA (Sw.) Gray – on plant remnants with sandy soil: 3 (1).
PORPIDIA FLAVOCAERULESCENS (Hornem.) Hertel & A.
J. Schwab in Hertel, Syn. P. flavicunda (Ach.)
Gowan – on rock: 3 (1).
PROTOPARMELIA PICEA auct. – on rock: 1 (1).
PSOROMA HYPNORUM (Vahl) Gray – on bryophytes
and plant remnants: 1, 5 (3).
RHIZOCARPON HOCHSTETTERI (Körb.) Vain. – on pebbles: 7 (1).
RINODINA ARCHAEA (Ach.) Arnold – on naked driftwood: 7 (1).
R. LECIDEOIDES (Nyl.) Kernst. –on dead bark of
Salix glauca: 8 (1).
R. MNIARAEA (Ach.) Körb. var. MNIARAEA – on bryophytes and plant remnants: 2, 9 (2).
R. MNIARAEA var. CINNAMOMEA Th. Fr. – on bryophytes and plant remnants: 3, 9 (2).
R. OLIVACEOBRUNNEA C. W. Dodge & G. E. Baker –
on bryophytes and plant remnants
(dead
parts of dwarf shrubs etc. ); common: 1, 2,
5, 7 (5).
R. TURFACEA (Wahlenb.) Körb. – on bryophytes and
plant remnants; common: 1, 2, 3, 5 (6).
PSOROMA HYPNORUM (Vahl) Gray – on soil: 1, 5
(4).
*SAGEDIOPSIS CAMPSTERIANA (Linds.) D. Hawksw.
& R. Sant – on Ochrolechia cf. androgyna
thallus: 3 (1).
SOLORINA SACCATA (L.) Ach. – on sandy soil with
plant remnants: 1, 5 (2).
S. SPONGIOSA (Ach.) Anzi – on sandy soil with plant
remnants: 1, 5, 7, 8 (4).
*SPHAERELLOTHECIUM ARANEOSUM (Rehm ex Arnold)
Zopf – on Cladonia pocillum and Ochrolechia
frigida thalli: 2, 9 (2).
*S. MINUTUM Hafellner – on Sphaerophorus fragilis
and S. globosus thalli; common: 1, 2 (2).
SPHAEROPHORUS FRAGILIS (L.) Pers. – on soil; 1 (1).
S. GLOBOSUS (Huds.) Vain. – on soil; common: 1,
2, 3, 6 (6).
STEREOCAULON ALPINUM Laurer – on soil; abundant
throughout the region: 1, 2, 3, 5, 7, 8, 9
(13).
S. BOTRYOSUM Ach. – on rock: 3 (2).
S. GLAREOSUM (Savicz) H. Magn. – on sandy soil,
often with silt: 7, 9 (2).
S. cf. PASCHALE (L.) Hoffm. – on soil: 2 (1).
S. RIVULORUM H. Magn. – on sandy soil; common:
2, 3, 7, 9 (7).
*STIGMIDIUM PELTIDEAE (Vain.) R. Sant. – on Peltigera leucophlebia thallus: 1 (1).
*T AENIOLELLA BESCHIANA Diederich – on Cladonia sp. primary squamules: 3 (Zhurbenko
83143) (1).
*T. PERTUSARIICOLA D. Hawksw. & H. Mayrhofer
– on Pertusaria carneopallida, mainly on
apothecia, also on thallus: 2 (Zhurbenko
83127) (1).
THAMNOLIA VERMICULARIS (Sw.) Schaer. var. VERMICULARIS – on soil; common: 1, 2, 5, 7, 8 (6).
T. VERMICULARIS var. SUBULIFORMIS (Ehrh.) Schaer.
– on soil; common: 1, 2, 3, 5, 8 (6).
TONINIA SQUALIDA (Ach.) A. Massal. – on bryophytes
with sandy soil: 3 (1).
TUCKERMANNOPSIS SEPINCOLA (Ehrh.) Hale – on bark
of Alnus fruticosa, Betula nana, and Salix
spp.: 2, 4, 8 (3).
UMBILICARIA DEUSTA (L.) Baumg. – on rocks: 3,
9 (2).
U. PROBOSCIDEA (L.) Schrad. – on rocks: 1, 3 (2).
U. TORREFACTA (Lightf.) Schrad. – on rock: 1 (1).
VULPICIDA PINASTRI (Scop.) J.-E. Mattsson & M. J.
Lai. – on bark of Alnus fruticosa and Salix
spp., also on soil and bryophytes: 1, 3, 4.
5, 8 (5).
ACKNOWLEDGEMENTS
Thanks are due to Prof. Dr Boris N. Norin and
Mr. Lembit Kitsing, the leaders of the Polar
Expedition of the Komarov Botanical Institute,
for the opportunity to collect lichens at Taimyr.
The help of Prof. Dr Teuvo Ahti, Dr Orvo Vitikainen, Dr Dagmar Triebel and Dr Irina I.
Makarova in the revision of some taxa is gratefully acknowledged. The preparation of this
159
paper has been financially supported by the
Russian Fund for Basic Researches (RFBR
grant no 97-04-48358). The study is part of the
project (NTP of Russia) "Complex studies of the
oceans and seas of the Arctic and Antarctic".
English has been kindly revised by Dr Barbara
Murray.
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160 Folia Cryptog. Estonica