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Folia Cryptog. Estonica, Fasc. 45: 13–22 (2009)
INTRODUCTION
Bryophytes, including mosses, liverworts, and
hornworts, form a conspicuous and important
component in many ecosystems throughout
the world. Their biological, ecological, and phy-
logenetic signicance is well documented (e.g.,
Pócs, 1980; Frahm, 1990; Coxson et al., 1992;
Hallingbäck & Hodgetts, 2000; O’Neill, 2000;
Gignac, 2001; Gradstein et al., 2001; Wellman
et al., 2003). Yet, compared to vascular plants,
liverworts remain a signicantly understudied
group, possibly due to their generally small
plant size coupled with difculties of identica-
tion. This concerns both the knowledge of their
ecological and biological features and their taxo-
nomic diversity. The importance of the present
paper is in summarizing the results of long-term
research in compiling an inventory of the spe-
cies of liverworts in the Vologda Region, one
of the largest areas of the Northern European
Russia (Fig. 1).
The Vologda Region is situated within the
East-European Plain and divided into 26 ad-
ministrative districts. The Region has an area
of 145 700 km2 and is of hilly relief, with alter-
nating depressions, ridges and elevations, with
the Northern Ridge at the east of the area. The
climate is moderately continental with a cold
winter (mean temperature of January is -14 C°)
and warm summer (mean temperature of July is
+18 C°) (Skupinova, 2007). The combination of
the relatively high mean precipitation, 500 mm
per year, and low evaporation, contributes to
the region abounding in rivers, lakes and bogs.
Vegetation is typical for middle and southern
taiga. The forests are mostly spruce woods and
occupy two-thirds of the area. Soils are podzolic
and soddy-podzolic, with boggy soils in some
areas (Vorob’ev, 2007).
Mosses of the Vologda Region were rst
mentioned in 1780 (Lepyokhin, 1780). Studies
devoted to liverworts of the Vologda Region have
been fragmentary and conducted in various
times in the 1890’s, 1950’s and 1960’s, the late
1980’s, as well as more recently in the present
decade. Signicant works in the last 200 years
include: Zickendrath (1900); Warnstorf (1913);
Perl’ev (1914, 1915), Abramova & Abramov,
1961; Akhminova & Zhukova, 1971; Volkova
et al., 1994; Andreeva, 2002. There are also
scattered papers that include notes on the liv-
Current state of knowledge of the liverwort and hornwort flora
of the Vologda Region, Russia
Michail V. Dulin1, Dmitriy A. Philippov2 & Elena V. Karmazina3
1Institute of Biology of the Komi SC UB RAS, Kommunisticheskaya st., 28, Syktyvkar, 167982, Komi Republic, Russia
E-mail: dulin@ib.komisc.ru
2I.D. Papanin Institute for Biology of Inland Waters of the RAS, Borok, 152742, Yaroslavl Region, Nekouz district, Russia
E-mail: philippov_d@mail.ru
3Vologda State Pedagogical University, Pobedy pr., 37, Vologda, 160035, Vologda Region, Russia
E-mail: ekarmazina@yandex.ru
Abstract: An annotated checklist of liverworts and hornworts is presented for the Volodga Region, Russia, based on the
authors’ collections and those of their colleagues, as well as the literature. e paper presents the first published checklist for
the region and includes 84 species from 42 genera and 22 families. ree species are reported for the first time for the region:
Conocephalum salebrosum Szweykowski et al., Scapania subalpina (Nees ex Lindenb.) Dumort., and Trichocolea tomentella
(Ehrh.) Dumort.
Kokkuvõte: Helvik- ja kõdersamblad Vologda regioonis (Venemaa)
Autorite kollekstsioonide ja kirjanduse põhjal on koostatud Venemaa Vologda oblasti helvik- ja kõdersammalde kommen-
teeritud nimestik. See on esimene selle piirkonna trükitud liiginimestik ning sisaldab kokku 84 liiki 42 perekonnast ja 22
sugukonnast. Esmakordselt registreeriti piirkonnast järgmised liigid: Conocephalum salebrosum Szweykowski et al., Scapania
subalpina (Nees ex Lindenb.) Dumort. ja Trichocolea tomentella (Ehrh.) Dumort.
14 Folia Cryptog. Estonica
erwort ora of Vologda Region (e.g., Karmazina
& Andreeva, 2004; Bakalin, 2005; Bobrov &
Chemeris, 2004, 2005; Bobrov, 2005; Ahti &
Boychuk, 2006; Philippov & Dulin, 2008). From
these references, 80 liverwort and 1 hornwort
species have been reliably reported for the region
up to the present time.
MATERIALS AND METHODS
Collections made by E.V. Karmazina and D.A.
Philippov during eld seasons from 2001 to
2008 throughout 10 municipal districts of the
Vologda Region formed the primary basis for
the present list. The studies were conducted
using a traditional route-based method, with
subsequent processing and identication of the
collections mainly by M.V. Dulin. Altogether ca.
350 samples were collected and studied, the ma-
jority of which are kept in the herbaria of the of
Department of Botany of the Vologda State Peda-
gogical University and I.D. Papanin Institute
for Biology of Inland Waters of the RAS (IBIW);
duplicates are deposited in the herbarium of the
Institute of Biology of the Komi Science Center
UB RAS (SYKO) and Lomonosov Moscow State
University (MW). In addition, we were also able to
study selected specimens from the Tsitsin Main
Botanical Garden of the RAS (MHA, collections
of M.S. Ignatov and E.A. Ignatova in the basins
of the Sukhona and the Yug rivers, 2001) and
from IBIW (collections of E.V. Chemeris and A.A.
Bobrov, 1996–2007).
The taxa in the checklist are arranged in
alphabetic order. Nomenclature follows Kon-
stantinova, Potemkin & Schlyakov (1992) with
some updates from the recent literature (Kon-
stantinova & Vasiljev, 1994; Konstantinova &
Potemkin, 1996; Grolle & Long, 2000; Damsholt,
2002). Taxonomy of Lophozia (Dumort.) Dumort.
follows Bakalin (2005). We share the ideas of
Schlyakov (1980, 1981), who followed the nar -
row generic concepts suggested by Scandinavian
researchers (Buch, 1933; Arnell, 1956). Modern
analysis of molecular data have provided more
precise denition of the systematic position of a
number of taxa (Vilnet et al., 2007). Synonyms
from regional papers are given in parenthesis
Fig. 1. The main collecting sites of liverworts in the Vologda Region.
15
after the accepted names of species. Because
only limited numbers of liverworts have been col-
lected in the Vologda Region, all known habitats
and localities are listed, by administrative divi-
sion. We include the geographical location of the
collection, collection dates, and the surnames
and initials of the collectors. Further information
on ecotopes and substrate types of the species is
presented in the checklist. Frequency of occur-
rence is estimated according to a 5-point scale:
very rare (1 nd), rare (2–5), sporadic (6–10),
frequent (11–15), very frequent (over 15). The
conservation status (category) is indicated for
rare endangered species registered in the Red
Data Book of the Vologda Region (Konechnaya
& Suslova, 2004). Species included in the Red
Data Book of the Vologda Region are marked
with #. Species recorded in Vologda Region for
the rst time are marked with *.
The main collecting sites of liverworts in
the Vologda Region
(Fig. 1; abbreviations: distr. – district; vill.
– village; coll. – collector)
Babaevo distr.: 1) the Koloshma river near Ko-
loshma vill.; 2) the Nozhema river near Pyazhelka
vill.; 3) the Parans river; 4) the Pyazhelka river
in the vicinity of the Pyazhelka vill., coll. E.V.
Chemeris & A.A. Bobrov in 2000–2008 (station
1–4 by Bobrov & Chemeris, 2004, 2005).
Babushkin distr.: 5) valley of the Zyablikha
river, coll. Mironova in 1957 (Akhminova &
Zhukova, 1971); 6) valley of the Mokrusha river,
coll. Mironova in 1957 (Akhminova & Zhukova,
1971); 7) left bank of the Staraya Tot’ma river
at Bol’shoy Dvor vill., coll. M.S. Ignatov & E.A.
Ignatova in 2001; 8) mire Mikhalyovo, vicinity
of Logduz and Pleshkino vill., coll. D.A. Philip-
pov in 2006.
Belozersk distr.: 9) vicinity of. Mishino vill,
coll. Metel’kova in 1956 (Akhimova & Zhukova,
1971).
Cherepovets distr.: 10) Darwinskiy State Natu-
ral Biospheric Reserve, coll. N.D. Nemtceva in
1987, 1991; A.L. Zhukova & A.D. Potemkin in
1988, 1990 (Volkova et al., 1994) & A.A. Bobrov
in 1996 (Bobrov, 2005).
Kharovsk distr.: 11) slope of riverbank of the
Sit’ river near Kharovsk settlement, coll. 1965
(Akhminova & Zhukova, 1971).
Kirillov distr. (all localities within the National
Park “Russkiy Sever”): 12) near Okulovskoe
Lake, vicinity of Okulovo vill., coll. T.A. Suslova
in 1987, E.N. Andreeva in 2002, E.V. Karmazina
in 2003; 13) near Gora Maura, coll. E.V. Kar-
mazina in 2003; 14) near Tcypina Gora, vicinity
of Oden’evo vill., coll. E.V. Karmazina in 2003;
15) vicinity of Goritcy vill., coll. E.V. Karmazina
in 2003; 16) near the Topornya river, nature
monument «Sokol’skiy Pinewood», coll. E.V. Kar-
mazina in 2003 & 2004; 17) vicinity of Kovarzino
vill., coll. E.V. Karmazina in 2003 & 2004; 18)
vicinity of Rusanovo vill., coll. E.V. Karmazina
in 2004; 19) vicinity of Vognema vill., coll. E.V.
Karmazina in 2004; 20) vicinity of Gorka vill.,
landscape reserve «Shalgo-Bodunovskiy For -
est», coll. E.V. Karmazina in 2003 & 2004, D.A.
Philippov in 2006; 21) along the Kishemskiy
Channel, coll. D.A. Philippov in 2007; 22)
Nikol’skoe Lake, vicinity of Schidorovo vill., coll.
D.A. Philippov in 2008.
Nikol’sk distr: 23) southwards from Nikol’sk
town, left bank of the Yug river near Permas vill.,
coll. M.S. Ignatov & E.A. Ignatova in 2001; 24)
mires on the slope of the Borovoy Kachug river,
near Kachug vill., coll. D.A. Philippov in 2006.
Nyuksenitca distr.: 25) bank of the Sukhona
river near Nyuksenitca settlement, coll. M.S.
Ignatov & E.A. Ignatova in 2001.
Sokol distr.: 26) no exact site specied, prob-
ably near town of Sokol, coll. M.S. Ignatov in
1990 (Bakalin, 2005); 27) mire Sludka, coll. D.A.
Philippov in 2007; 28) mire Alekseevskoe-1, vicin-
ity of Kadnikov town, coll. D.A. Philippov in 2007;
29) mire Durkovskoe northwards from railway st.
Morzhenga, coll. D.A. Philippov in 2007.
Tarnoga distr.: 30) the Ajga river vicinity of Il-
ezskiy Pogost vill., coll. D.A. Philippov in 2007;
Tot’ma distr.: 31) bank of the Kamchuga river,
near Kamchuga vill., coll. A.A. Korchagin in
1969 (Akhminova & Zhukova, 1971); 32) bank
of the Sukhona river, vicinity of Tot’ma town,
coll. 1965, 1969 (Akhminova & Zhukova, 1971);
33) vicinity of Bryukhachikha vill., coll. A.A.
Korchagin & O.F. Hase in 1926 (Akhminova &
Zhukova, 1971); 34) vicinity of Sovetskiy settle-
ment, coll. 1969 (Akhminova & Zhukova, 1971);
35) vicinity of Chernyakovo vill., coll. 1969
(Akhminova & Zhukova, 1971); 36) vicinity of
Ponomaryovo settlement, coll. 1969 (Akhminova
& Zhukova, 1971).
Ust’-Kubenskoe distr.: 37) north-eastern bank
of the Kubenskoe Lake, coll. E. Zickendrath in
1890 (Zickendrath, 1900).
16 Folia Cryptog. Estonica
Velikiy-Ustyug distr.: 38) slope of riverbank of
the river Sukhona near Opoki vill., coll. 1895
(Zickendrath, 1900); coll. 1968 (Akhminova &
Zhukova, 1971); 39) vicinity of Gorka settlement,
coll. E. Zickendrath in 1895 (Zickendrath, 1900);
40) mire near the Sinega settlement, coll. 1895
(Zickendrath, 1900); 41) bank of the Luza river,
coll. A.G. Kolmovskiy in 1895 (Zickendrath,
1900); 42) vicinity of Poldars settlement, on the
slope of river Sukhona, coll. 1968 (Akhminova
& Zhukova, 1971).
Vologda distr.: 43) vicinity of Vologda city, coll.
E. Zickendrath, 1891 (Zickendrath, 1900), A.A.
Snyatkov in 1896 (Zickendrath, 1900), I.A.
Perl’ev, G.I. Shiryaev & A.A. Snyatkov (Perl’ev,
1915), A.A. Korchagin (Akhminova & Zhukova,
1971); 44) vicinity of Nikolaevskoe-Vosimskoe
settlement, coll. E. Zickendrath in 1893 (Zick-
endrath, 1900); 45) vicinity of Severnaya Ferma
vill., coll. A.A. Snyatkov in 1895 (Zickendrath,
1900); 46) western bank of the Kubenskoe Lake,
near Kubenskoe settlement, coll. M.S. Ignatov
& E.A. Ignatova in 2001.
Vozhega distr.: 47) vicinity of railway st. of Kad-
nikovskiy vill., coll. 1958, 1963, 1965, 1966
(Akhminova & Zhukova, 1971); 48) vicinity of
Kuklinskaya vill., coll. D.A. Philippov in 2007.
Vytegra distr.: 49) vicinity of Andomskiy Pogost
vill., near Kivruchey, coll. A.K. Cajander in 1899
(Ahti & Boychuk, 2006); 50) mire Zhabinskoe,
vicinity Oshtinskiy Pogost settlement, coll. L.S.
Korotkevich in 1949 (Konechnaya & Suslova,
2004); 51) mire near the Panskoe Lake, vicinity
of Kedra vill., coll. D.A. Philippov in 2006; 52)
mire Vytegorskoe, vicinity Vytegra town, coll.
D.A. Philippov in 2006; 53) mire Gladkoe near
Vytegra town, coll. D.A. Philippov in 2006; 54)
Tudozero Lake, near Ostrov & Panovo vill., coll.
D.A. Philippov in 2005 & 2007; 55) mire in the
Palaya river valley, coll. D.A Philippov in 2006;
56) mire Shidry, vicinity of Pustosh’ & Nasonova
vill., coll. D.A. Philippov in 2006; 57) mire Soro-
zhskoe-Dol’noe, left bank of the Povreka valley,
near Tudozerskiy Pogost vill., coll. D.A. Philippov
in 2006; 58) slope of Onezhskoe Lake, vicinity
of Ol’kovo vill., coll. D.A. Philippov in 2006; 59)
mire Krestenskoe, vicinity of Ol’kovo vill., coll.
D.A. Philippov in 2006; 60) mire Desyatochnoe,
vicinity of Oktyabr’skiy settlement, coll. D.A.
Philippov in 2007; 61) the Oshta river, vicinity
of Kurvoshskiy Pogost vill., coll. E.V. Chemeris &
D.A. Philippov in 2007; 62) the Vodlica river near
Gornruchey vill., coll. D.A. Philippov in 2007; 63)
the Kurzheksa river, near Tikachyovo vill., coll.
E.V. Chemeris & D.A. Philippov in 2007.
CHECK-LIST OF THE LIVERWORTS AND
HORNWORTS OF THE VOLOGDA REGION
In the following list ‘small-leaved forest’ refers
to woodland of birch (Betula), aspen (Populus)
and alder (Alnus) trees.
MARCHANTIOPHYTA
AneurA pinguis (L.) Dumort. (Riccardia pinguis (L.)
Gray). – 10, 24, 38, 47, 48. Among the mosses
on sedge-moss spring bogs, in humid sites in
boggy spruce woods, on silty soil along the
banks of watercourses. Sporadic.
#ApometzgeriA pubescens (Schrank) Kuwah.
(Metzgeria pubescens (Schrank) Raddi)
– 47. Among beds of leafy forest mosses in
a boggy spruce forest. Very rare. Critically
Endangered.
bArbilophoziA bAr bAtA (Schmidel ex Shreb.)
Loeske – 7, 18, 37, 47, 48. On rotten stumps
and spruce trunks mainly in boggy spruce
woods, less frequent in pine forests on sandy
sloping riverbanks. Sporadic.
#bArbilophoziA lycopodioides (Wallr.) Loeske – 12,
47. On forest litter in blueberry and humid
mixed spruce forests. Rare. Least Concern.
blAsiA pusillA L. – 10, 16, 32, 35, 47, 58. On
sandy, less frequent on clay soil, on stream
banks, in various types of pine forests, once
in a swampy forest. Sporadic.
blephArostomA trichophyllum (L.) Dumort. var.
trichophyllum – 10, 16, 20, 21, 25, 34, 35, 40,
43, 45, 47. In small-leaved, mixed and conif-
erous forests of various degrees of humidity,
on rotten wood, stumps and at the foot of
birch and spruce trunks. Very frequent.
cAlypogeiA AzureA Stotler et Crotz (Calypogeia
trichomanis auct.; Kantia trichomanes auct.)
– 32, 43, 47. Boggy spruce forests, on wet
soil and on rotten fallen trees, less frequently
in bogs. Rare.
cAlypogeiA integristipulA Steph. – 10, 12, 17,
48. In boggy coniferous and mixed forests,
on rotten timber, trunk buttresses and less
frequent on soil. Frequent.
cAlypogeiA muelleriAnA (Schiffn.) Müll.Frib. – 10,
18, 20. In spruce and aspen woods, on rotten
wood and fallen trees. Sporadic.
cAlypogeiA neesiAnA (C.Massal. et Carestia) Müll.
Frib. – 6, 10, 12, 14, 27, 32, 47. In spruce
woods (mainly boggy), less frequently pine
forests, at the edges of Sphagnum mires,
17
on rotten wood, at the foot of aspen trunks,
sometimes on soil and litter, and also on
stumps and bare roots, among Dicranum sp.
and Tetraphis pellucida Hedw. Sporadic.
cAlypogeiA sphAgnicolA (Arnell et J.Perss.) Warnst.
et Loeske – 10, 28, 29, 52, 60. In meso- and
oligotrophic pine and dwarf-shrub Sphagnum
mires, on dwarf shrub/moss hummocks and
pine trunk burls, between beds of Sphagnum
ssp. Rare.
cAlypogeiA suecicA (Arnell et J.Perss.) Müll.Frib.
– 10, 16, 20. In a blueberry/monkshood
(Aconitum) pine forest, in a mossy pine for-
est, on rotten wood, stumps, once on peaty
soil. Rare.
cephAloziA Affinis Lindb. ex Steph. – 10. On birch
bark at base of trunks at the edges of mire.
Very rare.
cephAloziA bicuspidAtA (L.) Dumort. var. bicus-
pidAtA – 10, 16, 20, 32, 47. In coniferous
forests (mainly spruce woods) of various
degrees of humidication, in brook valleys,
on rotten wood, fallen trees and bare sandy
soil. Sporadic.
cephAloziA connivens (Dicks.) Lindb. – 10, 27,
32, 47, 59, 60. Between Sphagnum mosses
in boggy forests and on (mainly oligotrophic)
forest mires, on bare peat, at the foot of trees,
less frequent on sandy soil in dry woods and
at sloping riverbanks, sometimes on rotten
wood. Frequent.
cephAloziA loitlesbergeri Schiffn. – 10, 28.
On trunk buttresses on pine/dwarf shrub
Sphagnum bogs and in boggy pine forests.
Rare.
cephAloziA lunulifoliA (Dumort.) Dumort. (Cepha-
lozia media Lindb.) – 10, 16, 18, 20, 21, 32,
43. In small-leaved and coniferous forests
of various types, on rotten timber and for-
est soil, less frequent on peat, fallen trees or
sandy soil. Frequent.
cephAloziA pleniceps (Austin) Lindb. – 12, 17,
21, 40, 41, 43, 48, 53. Mainly on mires, on
hummocks, at the foot of trees and on bark
of pine stumps, among several Sphagnum
species (including Sphagnum centrale C.E.O.
Jensen et al.). Sporadic.
cephAloziellA divAricAtA (Sm.) Schiffn. (Cephaloz-
iella byssacea (A. Roth.) Warnst.) – 10, 43.
On sandy soil, in relatively dry pine forests.
Growing mixed with Buxbaumia apophylla
Hedw. Rare.
#cephAloziellA elAchistA (J.B.Jack ex Gottsche et
Rabenh.) Schiffn. – 50. On mesotrophic mire,
among Sphagnum and Hypnum mosses,
frequently mixed with other liverworts. Very
rare. Data Decient.
cephAloziellA hAmpeAnA (Nees) Schiffn. – 10, 32,
44. At sides of forest paths, on soil among
Ceratodon purpureus (Hedw.) Brid., Polytri-
chum sp., on a hummock near tree trunk in
a wet grassy birch forest. Rare.
cephAloziellA rubellA (Nees) Warnst. – 10. On
soil in relatively dry pine forests, at edges of
ditches, on rotten timber. Sporadic.
cephAloziellA spinigerA (Lindb.) Warnst. – 29, 48,
60. On meso- and oligotrophic pine/dwarf
shrub Sphagnum mires, on tussocks, among
Sphagnum. Rare.
chiloscyphus pAllescens (Ehrh. ex Hoffm.) Du-
mort. – 10, 12, 16, 17, 20, 47. In wet forests
(mainly spruce and birch woods) and stream
gullies, on soil, fallen trees, less frequently on
fallen leaves and rotten timber. Sporadic.
chiloscyphus polyAnthos (L.) Corda – 10, 16, 18,
19, 20, 21, 43, 49, 55, 56. In wet and boggy,
less frequent in dry spruce woods and small-
leaved forests, at shrubby and mesotrophic
mires and their edges, on rotten timber, fallen
trees, less frequently on bark and on soil
among fallen leaves. Frequent.
chiloscyphus rivulAris (Schrad.) Hazsl. – 1. On
banks and rapids of a rapid dark-water
stream with sandy-rocky ground. Rare.
clAdopodiellA fluitAns (Nees) H.Buch – 10, 28,
59, 60. In wet hollows and between tussocks
of mesotrophic and oligotrophic mires, and
also at edges of small lakes on bare peat be-
tween and on Sphagnum mosses. Sporadic.
conocephAlum conicum (L.) Dumort. – 9, 11, 12,
17, 20, 25, 30, 34, 48. On shrubby and for-
est-covered banks of brooks and on stones in
beds of small streams, in boggy small-leaved
forests, on wet soil, less frequently on fallen
trees. Frequent.
*conocephAlum sAlebrosum Szweykowski et al.
– 61. On forest-covered river bank on wet
soil. Very rare.
crossocAlyx helleriAnus (Nees ex Lindenb.) Meyl.
(Anastrophyllum hellerianum Schust.) – 34.
On rotten timber in a wood-sorrel (Oxalis)
spruce forest. Very rare.
crossogynA AutumnAlis (DC.) Schljakov (Jameso-
niella autumnalis (DC.) Steph.) – 43. Mire.
Very rare.
#frullAniA bolAnderi Austin – 34. On spruce
and stumps of elms and rowans in mixed
broad-leaved forest. Very rare. Critically
Endangered.
18 Folia Cryptog. Estonica
gymnocoleA inflAtA (Hads.) Dumort. – 10, 44. On
peaty soil and rotten timber at the edges of
boggy ditches and on mires. Rare.
isopAches bicrenAtus (Schmidel ex Hoffm.)
H.Buch (Lophozia bicrenata (Hoffm.) Dum.)
– 10. On sandy, less frequent loamy soil in
dry woods and at the banks of the Rybinsk
reservoir. Sporadic.
leiocoleA heterocolpos (Thed. ex Hartm.) H.Buch
var. heterocolpos – 10. On soil at eroded
sloping riverbank. Very rare.
leiocoleA rutheAnA (Limpr.) Müll.Frib. – 57. Eu-
trophic mire site saturated by groundwater,
in Sphagnum cover. Very rare.
lepidoziA reptAns (L.) Dumort. – 6, 10, 16, 18,
20, 21, 23, 25, 43, 47, 56. In coniferous and
small-leaved forests, on rotten stumps, rot-
ten timber, at foot of birch trunks, on peaty
soil in boggy forests and on eutrophic and
mesotrophic forest mires. Very frequent.
liochlAenA lAnceolAtA Nees – 20, 21, 60. On bare
peat at sedge tussocks and on super-humid
semi-rotten stumps of Salix aurita L. at
shrubby edges of eutrophic and mesotrophic
mires. Rare.
lophocoleA bidentAtA (L.) Dumort. – This species
has been indicated for the Vologda Region
without geographical and ecological data
(Andreeva, 2002). Rare.
lophocoleA cuspidAtA (Nees) Limpr. (Lophocolea
alata Mitt. et Larter) – 36. In alder forest at
the foot of a tree. Very rare.
lophocoleA heterophyllA (Schrad.) Dumort. – 8,
10, 16, 20, 21, 27, 32, 34, 36, 47, 48, 52, 53.
In forests of all types, streamside shrubs, on
mesotrophic and oligotrophic mires, at the
edges of eutrophic bogs, on rotten timber and
on bark of the lower parts of birch, alder, wil-
low, aspen, spruce, and lime trunks, frequent
on fallen trees, on ne earth between roots of
fallen trees, sometimes on soil and on fallen
spruce trees covered with sandy-silty sedi-
ments. Also found on Sphagnum centrale and
among Plagiothecium sp. Very frequent.
lophocoleA minor Nees – 7, 10, 14, 20, 32. In
aspen forests, on bark, roots and at foot of
aspen trees, less frequent on decomposed
timber and on soil at sloping stream banks.
Sporadic.
lophoziA excisA (Dicks.) Dumort. var. excisA
(Jungermannia socia Nees ab. Es.) – 10, 20,
37. On sandy and podzolic soils in dry leafy
and mixed forests. Rare.
lophoziA longidens (Lindb.) Macoun – 17, 20, 42,
47. On rotten timber, less frequent at the
foot of trees and on tree roots in coniferous
(mainly spruce forests) and small-leaved
forests. Rare.
lophoziA silvicolA H.Buch – 26, 60. On super-hu-
midied semi-rotten stumps of Salix aurita in
shrubby sedge-Menyanthes-Sphagnum bog
edge, mainly on rotten timber in coniferous
forests. Rare.
lophoziA ventricosA (Dicks.) Dumort. var. ven-
tricosA – 18, 39, 40, 47. In spruce woods of
various degrees of humidity, on mires, on
bark and rotten timber. Sporadic.
– var. guttulAtA (Lindb. et H.W. Arnell) Bakalin
(Lophozia longiora (Nees) Schiffn. var. gut-
tulata (Lindb. et H. Arn.) Schljak.) – 10, 20.
In spruce woods and aspen forests, on rotten
timber and decomposed fallen trees.
– var. longiflorA (Nees) Macoun (Lophozia porphy-
roleuca (Nees) Schiffner; L. longiora (Nees)
Schiffn. var. longiora (Nees) Schiffn.) – 10,
43. On soil in forests.
lophoziA wenzelii (Nees) Bakalin var. groenlAn-
dicA (Lophozia groenlandica (Nees) Macoun.;
L. confertifolia Schiffn.) – 10. On soil, trunk
buttress, rotten timber in mixed and conifer-
ous forests. Rare.
mArchAntiA polymorphA L (Marchantia aquatica
(Nees) Burgeff) – 6, 10, 34. On silty soil on
banks of forest brooks and on shallow banks
of the Rybinsk reservoir, less frequent in river
valleys on soil among large sedge clumps.
Rare.
mArchAntiA stellAtA Scop. (Marchantia poly-
morpha auct.) – Reliably known in over 50
sampling sites from Babaevo, Velikiy Ustug,
Verkhovazh’e, Vozhega, Vologda, Vytegra, Ki-
rillov, Sokol, Syamzha, Tot’ma, Cherepovets
districts on bare wet soils in grassy forests,
on stream banks, brooks, sites of old res, on
eutrophic grassy bogs, on soil between bricks
of walls and paths of Kirillovo-Belozerskiy
Monastery and Spaso-Prilutskiy Monastery.
Very frequent.
mArsupellA AquAticA (Lindenb.) Schiffn. – 4. In
rapid poorly-mineralized streams on siliceous
boulders, usually together with Scapania
undulata and Fontinalis dalecarlica Bruch
et al. Rare.
myliA AnomAlA (Hook.) Gray – 10, 12, 27, 28, 39,
43, 51, 52, 59. On mesotrophic and oligo-
trophic mires on hummocks and in hollows,
less frequent on hummocks around trunks,
on bare peat, sometimes among Sphagnum
mosses. Frequent.
19
nArdiA geoscyphus (De Not.) Lindb. – 10. On soil
in a wet coniferous forest. Very rare.
obtusifolium obtusum (Lindb.) S.W.Arnell (Lo-
phozia obtusa Evans) – 32. On soil at sloping
riverbank. Very rare.
odontoschismA denudAtum (Mart.) Dumort. – 10.
On peaty soil of a path at the edges of raised
bogs. Very rare.
#orthocAulis floerkei (F.Weber et D.Mohr)
H.Buch (Barbilophozia oerkei (F. Weber et D.
Mohr) Loeske) – 50. On a mesotrophic mire,
among leafy mosses and on trunk buttress.
Very rare. Endangered.
orthocAulis kunzeAnus (Huebener) H.Buch (Bar-
bilophozia kunzeana (Hübener) Buch) – 44,
53. On meso-olygotrophic pine/dwarf shrub
Sphagnum mire, on bark and burls of pine,
and also on Sphagnum centrale. Rare.
pelliA endiviifoliA (Dicks.) Dumort. – 11, 30, 32.
On wet soil at sloping stream bed, especially
near the water (at bank-water interface).
Together with other species of Pellia and
Marchantia. Rare.
pelliA epiphyllA (L.) Corda. – 1, 2, 3, 4, 10, 15.
On soil in humid forests of various types, on
steep banks of streams, forming Association
Pellio-Conocephaletum Maas 1959 var. Pellia
epiphylla. Sporadic.
pelliA neesiAnA (Gottsche) Limpr. – 10, 16, 20. On
wet soil in spruce forest and in pine wood, on
sandy soil of a re-protection ditch. Rare.
plAgiochilA Asplenioides (L. emend. Taylor) Du-
mort. (Plagiochila major (Nees) S. W. Arnell)
– 5, 20, 25, 31, 32, 43, 44, 45. In wood-
sorrel (Oxalis) and wood-sorrel/blueberry
spruce and spruce/aspen forests, and also
at eroded banks of ravines and brooks, on
soil. Sporadic.
plAgiochilA porelloides (Torr. ex Nees) Lindenb.
– 10, 18, 19, 20, 30, 37, 43, 48. In boggy
spruce woods, at the edges of eutrophic
mires, in stream gullies, on wet almost bare
soil, less frequently on rotten timber, fallen
leaves and twigs, trunk buttress and at foot
of aspens. Very frequent.
plectocoleA hyAlinA (Lyell) Mitt. (Jungermannia
hyalina Lyell.) – 10. Sandy stream bank, on
soil. Very rare.
preissiA quAdrAtA (Scop.) Nees – 11. On soil on
sloping riverbank. Very rare.
ptilidium ciliAre (L.) Hampe – 10, 18, 45. On soil in
dry moss-lichen pine forests, on bark and at
foot of trees in boggy spruce woods. Rare.
ptilidium pulcherrimum (Weber) Vain. –Widespread
all over the region. It is reliably known
(over 100 specimens in recent years) from
Verkhovazh’e, Vozhega, Vytegra, Kirillov,
Sokol, Syamzha, Tarnoga, Tot’ma, Chere-
povets districts. In small-leaved, mixed and
coniferous forests of various humidity levels,
on bark, lower limbs, tree burls, at the foot of
trees (frequent on birch, aspen, less frequent
on alder, rowan tree, spruce, juniper, lime),
rotten timber and fallen trees, seldom on
rocks. Very frequent.
rAdulA complAnAtA (L.) Dumort. – 12, 13, 14, 18,
20, 21, 38, 46, 47, 60. In mixed forests and
at the edges of mires, on bark of deciduous
trees (aspen, rowan, less frequently alder),
sporadic on fallen leaves of Populus suaveo-
lens Fisch. Very frequent.
riccArdiA lAtifrons (Lindb.) Lindb. – 20, 32, 40,
47. On rotten timber in spruce and birch
woods, on forest bogs. Sporadic.
riccArdiA pAlmAtA (Hedw.) Carruth. – 16, 43,
45. On rotten timber in mossy pine wood.
Rare.
#ricciA cAnAliculAtA Hoffm. – 10. On the banks
of water bodies on wet silty soil among
grassy vegetation. Very rare. Critically En-
dangered.
ricciA cAvernosA Hoffm. – 10. On silty soil of
banks in the ooded zone of Rybinsk reser-
voir. Rare.
ricciA fluitAns L. – 10, 46. On lake banks, wet
silt, or on exposed soil near water line of
water bodies, and in open water of ooded
sedge meadows and black alder wood around
beaver ponds. Sporadic.
ricciA huebeneriAnA Lindenb. – 4. In a very rapid
stony stream, on siliceous boulders. Rare.
ricciA sorocArpA Bisch. – 14. In a track rut on a
path. Very rare.
ricciocArpos nAtAn s (L.) Corda – 10, 22, 54.
In shallow slow-moving streams, bays of
streams and lakes, at the edges of oodplain
lakes, between sedge-reed beds. Rare.
scApAniA curtA (Mart.) Dumort. var. curtA – 10,
32, 33, 47. On sandy soil on sloping stream
banks, on forest edges and along paths in
small-leaved forests. Sporadic.
scApAniA irriguA (Nees) Nees – 5, 16, 32, 47, 56,
60. On soil, less frequent on rotten wood in
boggy birch and mixed forests, at the edges
of Sphagnum mires, on wet soil of sloping
stream banks and ditch edges, less frequent
on tussock sides in bogs. Frequent.
scApAniA pAludicolA Loeske et Müll.Frib. – 53.
Mesotrophic margin of a raised bog, sedge-
20 Folia Cryptog. Estonica
Hypnum-Sphagnum willow wood, on trunk
buttress of Salix cinerea L. and on sedge-Hyp-
num-Sphagnum carpet by shrubs. Rare.
*scApAniA subAlpinA (Nees ex Lindenb.) Dumort.
– 62, 63. On a semi-submerged log and rocks
near the stream bank. In pure patches or
mixed with Chiloscyphus polyanthos. Rare.
scApAniA undulAtA (L.) Dumort. – 1, 3, 4, 30. On
soil at wet meadows, in beds of small streams,
on stones, less frequently on wet timber. On
siliceous boulders in rapid streams together
with Marsupella aquatica (Association.
Scapanietum undulatae Schwick. 1944; Bo-
brov & Chemeris, 2004). Sporadic.
schistochilopsis incisA (Schrad.) Konstant. (Lo-
phozia incisa (Schrad.) Dum.) – 18, 20, 32,
40, 43, 47. In spruce woods (mainly boggy),
on rotten wood, decomposed fallen trees and
on soil. Sporadic.
solenostomA cAespiticium (Lindenb.) Steph.
(Jungermannia caespiticia Lindenb.) – 10. On
soil at the edge and bottom of a dry ditch in
a dry pine wood. Very rare.
#solen ostomA sphAerocArpum (Hook.) Steph.
(Jungermannia sphaercarpa Hook.) – 32, 40.
On bare soil at sloping stream banks and on
bare peat in bogs. Rare. Near threathened.
*trichocoleA tomentellA (Ehrh.) Dumort. – 48.
In humid meadowsweet (Filipendula)/mossy
spruce forest, on wet soil. Very rare.
ANTHOCEROTOPHYTA
phAeoceros lAevis (L.) Prosk. – 10. On a channel
bank, on sandy soil. Very rare.
RESULTS AND DISCUSSION
We report a total of 84 species and two varieties
of liverworts from 42 genera and 22 families for
the Vologda Region. Most of the liverwort spe-
cies belong to subclasses Jungermanniidae and
Marchantiidae of division Marchantiophyta and
only one species Phaeoceros laevis to division
Anthocerotophyta.
Three liverwort species, Conocephalum sale-
brosum, Scapania subalpina and Trichocolea
tomentella are listed for the rst time for the
Vologda Region.
Seven liverwort species are included in the
Red Data Book of the Vologda Region (Konech-
naya & Suslova, 2004) (Table 1).
All red-listed species are known only from
rare occurrences, except for Barbilophozia lyco-
podioides and Solenostoma sphaerocarpum, both
found at two geographical sites. Three montane
species, Apometzgeria pubescens, Orthocaulis
oerkei, and Frullania bolanderi, can be con-
sidered objectively rare owing to the absence or
extreme scarcity of available habitats. The lat-
ter species is a relic. In Europe it is known only
from several localities in Norway and Sweden
(Damsholt, 2002). In Russia it was found in the
Caucasus, the Northern and the Southern Urals,
the Altay, West Siberia, Yakutiya, and the Far
East including Sakhalin and Kamchatka (Kon-
stantinova et al., 1992). Cephaloziella elachista,
a boreal Atlantic liverwort, is found further to
the north in the Komi Republic and Karelia
(Bakalin, 1999; Dulin, 2008) and most likely it
Table 1. Rare liverwort species in the ora of the Vologda Region. Localities in the list of collecting
sites and Fig. 1. Each locality represents only one nding.
Taxa reat category No of localities Habitat Substrate
Apometzgeria pubescens Critically Endangered 47 Boggy spruce forest Among forest mosses
Barbilophozia lycopodioides Least Concern 12, 47 Blueberry and humid
small-leaved/spruce
forests
On forest litter
Cephaloziella elachista Data Deficient 50 Mesotrophic mire Among Sphagnum and
Hypnum mosses
Frullania bolanderi Critically Endangered 34 Coniferous/broad-
leaved forest
On stumps of elm and
rowan
Orthocaulis floerkei Endangered 50 Mesotrophic mire Among leafy mosses
Riccia canaliculata Critically Endangered 10 Banks of a water body On wet silty soil
Solenostoma sphaerocarpum Near threathened 32, 40 Peat bog and sloping
stream bank
On bare soil
21
is overlooked during collecting owing to its tiny
size. The nemoral amphiatlantic liverwort Riccia
canaliculata obviously has the north-eastern
limit of its distribution in the region. The rar-
ity of arctic-boreal circumpolar Barbilophozia
lycopodioides and Solenostoma sphaerocarpum
is presumably caused by insufcient study of
regional bryoora, and without any doubt future
study will reveal new localities for these spe-
cies. We recommend excluding Barbilophozia
lycopodioides and Solenostoma sphaerocarpum
from the threatened species list of the Red Data
Book and propose including some rare species
to it (Crossocalyx hellerianus, Lophocolea cus-
pidata, Marsupella aquatica, Odontoschisma
denudatum, Phaeoceros laevis, Riccia cavernosa,
Trichocolea tomentella).
Among the new discoveries, Conocepha-
lum salebrosum, a recently described taxon
(Szweykowski et al., 2005) is of interest. An
insufcient knowledge coupled with consider-
able morphological similarity to the closely
allied species, C. conicum, made it difcult to
identify C. salebrosum both in the eld and
during processing of collections. However, a
careful revision of our collections enabled us
to detect C. salebrosum among samples previ-
ously identied by us as C. conicum. Although
C. salebrosum is cited only from one locality,
we predict that it has a wider distribution than
currently reported.
ACKNOWLEDGEMENTS
Authors express their gratitude to N.A. Kon-
stantinova (PABSI RAS, Kirovsk) for checking
the authenticity of some samples and also ac-
knowledge E.V. Chemeris & A.A. Bobrov (IBIW
RAS, Borok), M.S. Ignatov (SBG RAS) & E.A.
Ignatova (MSU) for providing the opportunity to
work with herbarium samples. They also express
sincere gratitude to S.V. Balina for assistance
in translation of the article and V.V. Elsakov for
help in creating of the map. The research of M.V.
Dulin was performed under the nancial sup-
port of Russian Foundation for Basic Research
(project No. 09-04-00281).
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