1067-4136/05/3605- © 2005 Pleiades Publishing, Inc.
Russian Journal of Ecology, Vol. 36, No. 5, 2005, pp. 312–319. Translated from Ekologiya, No. 5, 2005, pp. 344–352.
Original Russian Text Copyright © 2005 by Shvartsman, Bolotov.
The distribution of biomes on land is characterized
by altitudinal zonality, which is explained by the fact
that the amount of incident solar radiation decreases
from the equator to the poles (Chernov, 1975). How-
ever, there are many other factors that inﬂuence the for-
mation of terrestrial biocenoses and that are responsible
for their marked heterogeneity, such as speciﬁc combi-
nations of heat and moisture gradients on continents,
the orientation of large mountain ranges, and some his-
torical factors (Chernov and Penev, 1993). Analysis of
the causes of the spatial heterogeneity of biocenoses is
among the most complex tasks in modern ecology.
Islands are convenient model objects for a broad
spectrum of studies in different ﬁelds, from synecolog-
ical and biogeographic to evolutionary. Insular bio-
cenoses of high northern latitudes have been studied
fragmentarily: on the one hand, access to northern
islands is difﬁcult; on the other hand, researchers have
always been more interested in investigating tropical
This is also true of the Solovetskiye Islands. This
unique archipelago has been studied mainly from the
historical and cultural aspects. It has the status of a
national park, which is famous for its historical–archi-
tectural museum: its architectural ensemble is on the
United Nations World Heritage List. Nevertheless, data
on components of the natural environment of these
islands are still fragmentary.
In this study, we made an attempt to answer the con-
troversial question as to why biocenoses characteristic
of several natural zones have developed within the
small area of the Solovetskiye Islands.
MATERIAL AND METHODS
Studies of vegetation.
The distribution of bio-
cenoses over the area of the archipelago was studied
using images made by the Landsat 7 satellite in differ-
ent regions of the spectrum (channels 3, 4, and 5) at a
resolution of 30 m on June 21, 2000. They were synthe-
sized into an integrated image, which was deciphered
by means of cluster analysis in the i.cluster module of
the GRASS 5.0.3 GIS program package.
The results of deciphering were interpreted on the
basis of materials obtained in the course of ﬁeld studies
on the islands (Fig. 1). Expeditions launched in 2001
and 2003 made concise geobotanical descriptions of
biocenoses along their routes, including the identiﬁca-
tion of dominant plant species. In 2003, an additional
39 descriptions of forest phytocenoses in 10
plots were made on the Bol’shoi Solovetskii Island.
Geographic coordinates of each plot were determined
using a Garmin 3 Plus GPS receiver. A cluster analysis
of the geobotanical descriptions for similarity in the
species composition of the herb–dwarf shrub layer (the
Czekanowski–Sørensen index) and interpretation of the
resulting dendrogram allowed us to distinguish three
groups of forest biocenoses that were close to the mid-
dle taiga, northern taiga, and subtundra biocenoses.
These data were used to make a raster map of the
plant cover of the Solovetskiye Islands, which was sub-
Mechanisms of the Formation of Extrazonal Biocenoses
on the Solovetskiye Islands
Yu. G. Shvartsman* and I. N. Bolotov**
*Lomonosov State University,
pr. Lomonosova 4, Arkhangelsk, 163006 Russia
**Institute of Ecological Problems of the North, Ural Division, Russian Academy of Sciences,
nab. Severnoi Dviny 23, Arkhangelsk, 163061 Russia
Received November 24, 2004
—On the Solovetskiye Islands, subtundra forests and forest–tundra crooked forests are considered to
be zonal biocenoses, as this region is intersected by the 12
C isotherm for July, which marks the boundary
between the northern taiga and forest–tundra zones in northern Europe. The formation of extrazonal tundras on
the islands is attributed to the cooling effect of the sea and the prevalence of cold northeasterly winds in the
growing period. Herbaceous and herb–bilberry forests similar to their middle-taiga counterparts develop in this
region due to a convective heat ﬂow anomaly of high intensity (up to 40–50 W/m
). The phenomenon provides
evidence for the existence of an active heat source in the crust under the archipelago. This source is probably a
diapir fold involving the upward movement of the mantle.
: biocenoses, the Solovetskiye Islands, natural zonality, convective heat ﬂow.