ISSN 1067-4136, Russian Journal of Ecology, 2009, Vol. 40, No. 3, pp. 180–186. © Pleiades Publishing, Ltd., 2009.
Original Russian Text © E.Z. Baisheva, S.N. Zhigunova, V.B. Martynenko, P.S. Shirokikh, 2009, published in Ekologiya, 2009, No. 3, pp. 195–201.
Studies on the ﬂoristic composition and cenotic role
of bryophytes in forest communities have been per-
formed since the ﬁrst half of the 20th century (Grebe,
1917; Sukachev, 1927). Conditions of illumination,
moisture supply, and mineral nutrition, as well as
parameters of tree stand and the composition and depth
of forest litter are regarded as the main ecological fac-
tors determining the composition and abundance of
bryophytes in forest ecosystems (Akhminova, 1970,
1983; Cherepanova and Ipatov, 2003; Rambo and Muir,
1998; and others). Epigeic moss synusiae have a con-
siderable effect on regeneration of conifers and on the
course of soil-forming processes in boreal forests.
Hence, bryophytes have been widely used for classiﬁ-
cation of plant communities and for interpretation and
indication of many complex phenomena and processes
in forests of the taiga zone, including natural regenera-
tion, succession, paludiﬁcation, and soil formation
(Sukachev, 1927; Akhminova, 1970, 1983; Økland,
1994; Rambo and Muir, 1998; Vellak et al., 2003).
At the same time, it has been noted that the develop-
ment of the moss layer depends on the structure of other
layers of the forest phytocenoses: the higher the com-
plexity of the community and population density in its
lower layers, the less developed is the moss layer.
Bryophytes in ecosystems of boreal spruce forests
have been studied in fair detail, whereas such studies on
conifer and mixed nemoral forests are rare (Hokkanen,
2004; Shestakova, 2004). The results of ordination
analysis of mixed forests with a rich herbaceous layer
in Finland (Hokkanen, 2004) show that the abundance
of bryophytes in these communities is low, but their
species diversity is high and reﬂects those habitat con-
ditions that barely affect the composition of vascular
plants (air humidity, ecological conditions on the soil
surface, etc.). Therefore, for a detailed classiﬁcation of
such forests, it is necessary to take into account avail-
able data on both vascular plants and mosses.
The Ufa Plateau is of interest for studies on the
effects of ecological factors on the structure of bryo-
phytes in forest communities. This area is characterized
by a high diversity of site conditions, tree species, and
plant communities. There is a unique complex of nem-
oral herbaceous broadleaf–dark conifer, dark conifer,
and light conifer–broadleaf forests as well as typical
boreal light and dark conifer forests of green moss type
on poorly developed cryogenic humus–calcareous soils
(Kulagin, 1978; Mart’yanov et al., 2002; Zhigunova,
The purposes of this study were as follows: to ana-
lyze the diversity of bryophytes in forest communities
on the Ufa Plateau, to reveal trends in the distribution
of bryophyte species in the space of higher syntaxa of
forest vegetation (at the level of unions), and to esti-
mate the effect of phytocenotic and abiotic environ-
mental factors on the ground moss layer.
A promising approach to the study of factors inﬂu-
encing the ﬂoristic composition and synusial structure
of the moss layer is to compile and analyze the check-
lists of bryocenoﬂoras of forest associations. Such
checklists include parameters indicating phytocenotic
signiﬁcance of individual species and allow a fairly
detailed assessment of factors that have an effect on the
species diversity and synusial structure of the moss
layer (Akhminova, 1983).
Ecological and Phytocenotic Features of the Bryophyte
Component of Water Protection Forests on the Ufa Plateau
E. Z. Baisheva, S. N. Zhigunova, V. B. Martynenko, and P. S. Shirokikh
Institute of Biology, Ufa Scientiﬁc Center, Russian Academy of Sciences,
prosp. Oktyabrya 69, Ufa, 450054 Bashkortostan, Russia;
Received March 28, 2007
—The composition of bryophytes in forests of the Ufa Plateau has been studied. The results obtained
by ordination methods show that the coverage of epigeic mosses decreases with an increase in tree stand density
and proportion of broadleaf tree species as well as in the coverage and average height of herbaceous layer. The
main factors determining the distribution of epigeic mosses are illumination level, soil fertility, and the degree
of soil development.
: bryophytes, ordination, conifer–broadleaf forests, ecological factors.