ISSN 10674136, Russian Journal of Ecology, 2010, Vol. 41, No. 5, pp. 378–385. © Pleiades Publishing, Ltd., 2010.
Original Russian Text © P.P. Popov, 2010, published in Ekologiya, 2010, No. 5, pp. 336–343.
The spruce had spread over the presentday terri
tory of northwestern Russia and neighboring regions
of Finland, Sweden, and Norway no earlier than
10500–9000 thousand years ago, with the source of its
spread apparently lying in more southern regions
(Serebryakov, 1936; Lindquist, 1948; Dagfin, 1970;
Tallantire, 1972; Serebryanyi, 1974; Lang, 1994;
Ravazzi, 2002). As early as in the 19th century, it was
found that spruce stands in Norway, Sweden, Finland,
and on the Kola Peninsula contained some trees that
could not be attributed to either Norway or Siberian
spruce by the shape of seed scales (Keppen, 1885).
In 1863, the corresponding forms of spruce were rec
ognized as varieties and named var.
W. Nylander or var.
by E. Regel (cited from
Pravdin, 1975). Two decades later, Regel (1983) wrote
that “In Finland and the vicinity of St. Petersburg, as
well as in Eastern Russia, the forms of spruce occur
that we have named
P. excelsa fennica.
the shape of seed scales, they should be regarded as
intermediate forms.” Wolf (1925) assigned the name
Picea excelsa fennica
to the whole variety of interme
diate (transitional) forms.
Fedorovich (1876) logically supposed that such
forms appeared due to “cross fertilization,” i.e.,
hybridization between Norway spruce and Siberian
spruce (Danilov, 1943; Bobrov, 1944, 1974; Pravdin,
1972; Koropachinskii and Milyutin, 2006). It is con
sidered that hybrid (intermediate, or transitional)
forms of spruce and their populations originated in the
course of opposite migrations of Norway spruce from
the east and Siberian spruce from the east (Danilov,
1943; Bobrov, 1944; Golubets, 1968; Pravdin, 1975).
According to Korchagin (1968), these species first
came in contact with each other in the Northern
Dvina River basin, whence hybrid spruce has widely
spread westward and eastward. Latitudinal dynamics
of the structure of spruce populations including hybrid
trees have also been studied (Heikinheimo, 1920; Bak
shaeva, 1970; Shcherbakova, 1973; Il’inov et al.,
1998). The majority of relevant studies were based on
visual examination and classification of seed scales by
shape. Although their authors managed to reveal cer
tain trends of changes in population structure, the
level and profundity of such research have changed
only slightly since the 19th century. Therefore, the
purpose of this study was to analyze the form structure
and geographic differentiation of spruce populations
in northwestern Russia using the methods of metrics
and discriminant analysis.
MATERIAL AND METHODS
The study was performed with samples of cones
from spruce populations growing in the vicinities of
Vyborg (Leningrad Region); Sortavala, Ladva, Petroza
vodsk, Pudozh, Reboly, Kem’ (Republic of Karelia);
Konosha, Obozerskoe (Arkhangelsk Region); and Apa
tity (Murmansk Region) (Fig. 1). The sample size
ranged from 124 to 252 trees per population. From each
tree a mediumsized cone was collected, and a “typi
cal” seed scale from its middle part was cut out and
measured to calculate the coefficients of narrowing
) and projection (
) (Fig. 2). Cone length was also
Form Structure and Geographic Differentiation
of Spruce Populations in Northwestern Russia
P. P. Popov
Institute of the Problems of Development of the North, Siberian Branch, Russian Academy of Sciences,
a/ya 2774, Tyumen, 625003 Russia
Received May 12, 2009
—Discriminant analysis has been used to analyze the composition of phenotypic forms and geo
graphic differentiation of spruce populations in northwestern Russia in comparison with those of “standard”
Norway spruce and Siberian spruce populations. The absolute prevalence of trees with an intermediate Nor
way–Siberian spruce phenotype has been revealed in almost all populations, except for the northernmost
ones. Spruce populations in northwestern Russia may be divided into at least three groups, with the southern
group being closer to the Norway spruce, and all other groups, to the Siberian spruce.
: intermediate spruce populations, phenotypes, discriminant analysis, population structure,
squared Mahalanobis distances.