Petr P. Popov, Chief Researcher, Dr. Sci. in the Institute of the Problems of Northern Development, Federal Research Center Tyumen Scientific Center of the Siberian Branch of the RAS, Russia (625003, Tyumen, str. Malygina, 86, PO Box 2774); e-mail:
Stanislav P. Arefyev, Chief Researcher, Head of the sector of biodiversity and dynamics of natural complexes, Dr.Sci. in the Institute of the Problems of Northern Development, Federal Research Center Tyumen Scientific Center of the Siberian Branch of the RAS, Russia (625003, Tyumen, str. Malygina, 86, PO Box 2774); Professor of the Department of Botany, Biotechnology and Landscape Architecture in the Institute of Biology of the Tyumen State University; e-mail:
Maria N. Kazantseva, PhD, Leading Researcher in the Institute of the Problems of Northern Development, Federal Research Center Tyumen Scientific Center of the Siberian Branch of the RAS, Russia (625003, Tyumen, str. Malygina, 86, PO Box 2774); Associate Professor of the Department of Ecology and Genetics in the Institute of Biology of the Tyumen State University; e-mail:

Reference to article

Popov P.P., Arefyev S.P., Kazantseva M.N. 2019. Phenotypic diversity of spruce populations in some Protected Areas in Eastern Europe and Siberia. Nature Conservation Research 4(4): 26–33.

Section Research articles

In many Protected Areas of Eastern Europe and Siberia, the European spruce (Picea abies) or Siberian spruce (Picea obovata) plays a significant role in plant communities. Difficulties in determining the specific and intraspecific diversity of spruce in Protected Areas occur in the zone of natural (introgressive) hybridisation of spruce. Since the time of Ledebour's studies, the main diagnostic feature of these spruce species is the shape of the upper part of seed (cone) scales. The European spruce has an angularly acuminate and noticeably elongated cone scale, while the cone scale of the Siberian spruce has a close to rounded form and it is less elongated. In East-European spruce populations, formed under introgression influence, this trait has an intermediate character. The shape of the upper part of spruce seed scales remains constant throughout the whole tree life. And it is characterised by a high variability within populations and between different populations. Nowadays, the study of the phenotypic diversity of the spruce populations is performed using visual descriptive assessment, i.e. subjectively. Consequently, disparate results are often obtained even for the same study areas. The applying of metric assessment of a trait and the consequent distinguishing of the phenotypes of individuals allow to perform mathematical processing of the results and to obtain objective data about the population structure. We aimed to study the phenotypic diversity of spruce in Protected Areas in Eastern Europe and Siberia on the basis of metric assessment primarily of the shape of seed scales. The determination of the coefficient of narrowing (Cn) and coefficient of projection (Cp) of the upper part of seed scale can be considered as the most objective method to study this diagnostic trait. The difference between these coefficients (Cn‒Cp) is a more informative indicator to characterise properties of individuals, populations and their groups. In Eastern Europe and Siberia, we distinguished nine areas of distribution of different population groups (or phenotypes): P.e., P.eem., P.em., P.emm., P.m., P.mms.,, P.mss., P.s. They are characterised by an average value of the Cn‒Cp indicator: -50%, -40%, -30%, -20%, -10%, 0%, 10%, 20%, 30%, respectively. On the basis of this gradation in populations, we adopted a similar gradation of individuals' phenotypes: e, eem, em, emm, m, mms, ms, mss, s. On the basis of study of the spruce population diversity in Protected Areas in Eastern Europe and Siberia, considering phenotypes identified by the metric traits of seed scales, we found that all of them significantly differed in terms of all analysed traits. In the National Park «Belovezhskaya Pushcha», Carpathian Biosphere Reserve, Central Forest State Natural Biosphere Reserve, and Oksky State Natural Biosphere Reserve (European range of spruce), we found the highest cone length, highest phenotype frequency of European spruce individuals (60–100%). In these Protected Areas, there are no or very few intermediate phenotypes, and no Siberian spruce phenotypes. The Lapland State Natural Biosphere Reserve, Visimskiy State Natural Biosphere Reserve, Yuganskiy State Nature Reserve, Sokhondinskiy State Natural Biosphere Reserve, and Vitimskiy State Nature Reserve formed the opposite group according to all indicators studied. These Protected Areas are located within the range of the Siberian spruce. They are characterised by predomination (63–100%) of phenotypes of Siberian spruce individuals, while there are no European spruce phenotypes. We established the predominance of intermediate spruce phenotypes in the State Nature Reserve «Kologrivskiy Les», National Park «Chavash Varmane», Volzhsko-Kamskiy State Nature Biosphere Reserve, and Pinega State Nature Reserve. Of them, the first two Protected Areas are a bit closer to European spruce phenotypes, while the second two Protected Areas are characterised by similarity to the Siberian spruce phenotypes. The objective, sufficiently tested method to study the spruce population phenotypic diversity can also be used inother Protected Areas.


national park, nature reserve, phenotypic diversity, Picea abies, Picea obovata

Artice information

Received: 04.06.2019. Revised: 12.08.2019. Accepted: 13.08.2019.

The full text of the article

Bobrov E.G. 1944. On Specific Features of the Flora of an Erratic Region (One of Pathways in the Development of Forms). Sovetskaya Botanika 2: 3–20. [In Russian]
Bobrov E.G. 1971. History and systematic of genus Picea A. Dietr. Novitates Systematicae Plantarum Vascularium 7: 5‒40. [In Russian]
Bobrov E.G. 1974. Introgressive hybridisation in the genus Picea A. Dietr. Proceedings of Institute of Plant and Animal Ecology of AS USSR 90: 60–66. [In Russian]
Bobrov E.G. 1978. Forest-forming Conifer Species in the Soviet Union. Leningrad: Nauka. 188 p. [In Russian]
Borovikov V.P. 1998. A popular introduction to STATISTICA. Moscow: ComputerPress. 267 p. [In Russian]
Danilov D.N. 1943. Variability of the seed scales Picea excelsa. Botanicheskii Zhurnal 28(5): 191‒202. [In Russian]
Forest Encyclopedia. Vol. 1. Moscow: Sovetskaya Entsiklopedia, 1985. 563 p. [In Russian]
Golubets M.A. 1960. Two subspecies of Picea excelsa Link. and the question of their ranges. Botanicheskii Zhurnal 45(5): 684‒694. [In Russian]
Goncharenko G.G., Padutov V.E. 2011. Population and evolutionary genetics of Palearctic spruce. Gomel: Belarus. 197 p. [In Belarus]
Grahl-Nielsen O., Mjaavatten O., Øvstedal D.O. 1991 A chemometric comparison between Picea abies and P. obovata (Pinaceae) in Norway. Nordic Journal of Botany 11(6): 613–618. DOI: 10.1111/j.1756-1051.1991.tb01271.x
Indigenous dark-coniferous forests of the southern taiga (reserve area «Kologrivsky les»). Moscow: Nauka, 1982. 220 p. [In Russian]
Koropachinskiy I.Yu. 1992. Botanic-geographical aspect natural hybridisation of woody plants. Lesovedenie 2: 3–10. [In Russian]
Koropachinsky I.Yu., Milyutin L.I. 2006. Natural hybridisation of woody plants. Novosibirsk: Geo. 223 p. [In Russian]
Krutovskii K.V., Bergmann F. 1995. Introgressive hybridization and phylogenetic-relationships between Norway, Picea abies (L.) Karst., and Siberian, P. obovata Ledeb., spruce species studied by isozyme loci. Heredity 74(5): 464–480. DOI: 10.1038/hdy.1995.6
Lagercrantz U., Ryman N. 1990. Genetic structure of Norway spruce (Picea abies): concordance of morphological and allozymic variation. Evolution 44(1): 38‒53. DOI: 10.1111/j.1558-5646.1990.tb04278.x
Lundkvist K., Rudin D. 1977. Genetic variation in eleven populations of Picea abies as determined by izozime analisis. Hereditas 85(1): 67‒73. DOI: 10.1111/j.1601-5223.1977.tb00951.x
Minyaev N.A., Konechnaya G.Yu. 1976. Flora of the Central Forest State Reserve. Leningrad: Nauka. 104 p. [In Russian]
Orlova L.V., Egorov A.A. 2010. To systematics and geographical distribution of Finnish spruce (Picea fennica (Rtgel) Kom., Pinaceae). Novitates Systematicae Plantarum Vascularium 42: 5–23. [In Russian]
Popov P.P. 1999. Geographic variation in the form of spruce seed scales in Eastern Europe and Western Siberia. Lesovedenie 1: 68‒73. [In Russian]
Popov P.P. 2012. Reference Populations for Discriminant Analysis in the Continuous Range of Norway and Siberian Spruces. Russian Journal of Ecology 43(1): 13–18. DOI: 10.1134/S1067413612010092
Popov P.P. 2013. The phenotypic structure of the populations Picea abies and P. obovata (Pinaceae) in the eastern of Europe. Botanicheskii Zhurnal 98(11): 1384–1402. [In Russian]
Popov P.P. 2017. Population Structure in European and Siberian Spruces According to Their Phenotypes. Russian Journal of Ecology 48(5): 403‒408. DOI: 10.1134/S1067413617050101
Popov P.P. 2017. The distribution of individuals of intermediate form in the populations of Norway and Siberian spruces. Siberian Journal of Forest Science 4: 13‒19. DOI: 10.15372/SJFS20180402 [In Russian]
Popov P.P., Arefyev S.P., Kazantseva M.N. 2017. Phenotypic structure of spruce populations in the reserves «Tsentralno-lesnoy», «Kologrivskiy les» and «Basegi». Tyumen State University Herald. Natural Resource Use and Ecology 3(3): 50‒60. DOI: 10.21684/2411-7927-2017-3-3-50-60 [In Russian]
Pravdin L.F. 1975. Norway Spruce and Siberian Spruce in the Soviet Union. Moscow: Nauka. 200 p. [In Russian]
Shcherbakova M.A. 1973. The Spruce Genetics and Ecology of Picea abies (L.) Karst. in different forest areas. PhD Thesis Abstract. Krasnoyarsk. 26 p. [In Russian]
Schmidt-Vogt H. 1972. Studien zur morphologischen Variabilitдt der Fichte (Picea abies (L.) Karst.). 3. Der gegenwдrtige Stand der Forschung zur morphologischen Variabilitдt der Fichte – gesetzmдssigkeiten und Theorien. Allgemeine Forst und Jagdzeitung 143(11): 221‒240.
Sokolov S.Ya., Svyazeva O.S., Kubli V.A. 1977. The Ranges of trees and shrubs of the USSR. Leningrad: Nauka. 163 p. [In Russian]
Tatarinov V.V. 1987. Comparative analysis of the phenotypic variability populations of spruce in the communities of spruce forest in the central part of Russian Plains. Botanicheskii Zhurnal 72(2): 229‒238. [In Russian]
Tatarinov V.V. 1989. The structure of spruce populations in the forests of the Middle Urals. In: Population studies of plants in nature reserves. Moscow: Nauka. P. 130‒142. [In Russian]
Teplouchoff T. 1868. Ein Beitrag zur Kenntniss der sibirischen Fichte – Picea obovata Ledeb. Bulletin de la Societe imperial des naturalistes de Moscou 41(3): 244–252.
Teploukhov F.A. 1872. News of the activities of the Forest Society. Lesnoy Zhurnal 6: 86‒91. [In Russian]
Zhivotovskiy L.A. 1982. Parameters of population variability by polymorphic features. In: Phenetics of populations. Moscow: Nauka. P. 38‒45. [In Russian]