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Farukshina, G.G. /Vol. 8 Núm. 21: 276 - 283/ Julio - agosto 2019 276

Artículo de investigación The history of the formation of the population structure of Juniperus communis L. (Cupressaceae) in the Cis-Urals and the Southern Urals

ИСТОРИЯ ФОРМИРОВАНИЯ ПОПУЛЯЦИОННОЙ СТРУКТУРЫ МОЖЖЕВЕЛЬНИКА ОБЫКНОВЕННОГО JUNIPERUS COMMUNIS L. (CUPRESSACEAE) В ПРЕДУРАЛЬЕ И НА ЮЖНОМ УРАЛЕ

Historia de la formación de la estructura de la población de Junger mushroom del juniperus communis l. (Cupressaceae) en la Ural y en los Urales del sur

Recibido: 4 de junio del 2019 Aceptado: 6 de julio del 2019

Written by: G. G. Farukshina84 SPIN-код 6236-4644 Anthor ID 89582

Abstract Аннотация

The possible ways of the historical development Рассмотрены возможные пути исторического of juniper (Juniperus communis L.) in the Cis- развития можжевельника обыкновенного Urals and in the South Urals are considered. The (Juniperus communis L.) в Предуралье и на main habitats and populations of the species are Южном Урале. Указаны основные районы indicated. It is shown that the history of the обитания и популяции вида. Показано, что settlement of Juniperus communis is consistent история расселения можжевельника with the history of the development of pine обыкновенного согласуется с историей forests in the region. The South Ural mountain развития сосновых лесов в регионе. Наиболее population has the earliest origin (over 12 раннее происхождение (старше 12 тысяч лет) thousand years), the Cis-Ural forest-steppe имеет южноуральская горная популяция, population has the most recent (about 3-4 наиболее позднее (около 3-4 тысяч лет) – thousand years), the Cis-Ural forest population предуральская лесостепная популяция, has an average formation time (about 10-12 среднее по времени формирования (около 10- thousand years). 12 тысяч лет) – предуральская лесная популяция. Key words: Juniperus communis, Southern Urals, Cis-Urals, history of settlement, Ключевые слова: Juniperus communis, population. Южный Урал, Предуралье, история расселения, популяции.

Resumen

Se consideran las posibles vías del desarrollo histórico del enebro (Juniperus communis L.) en los Cis- Urales y en los Urales del Sur. Se indican los principales hábitats y poblaciones de la especie. Se muestra que la historia del asentamiento de Juniperus communis es consistente con la historia del desarrollo de los bosques de pinos en la región. La población de las montañas del sur de los Urales tiene su origen más temprano (más de 12 mil años), la población de estepas del bosque Cis-Ural tiene la más reciente (alrededor de 3-4 mil años), la población del bosque Cis-Ural tiene un tiempo de formación promedio (aproximadamente 10-12 mil años).

84 South-Ural Botanical Garden-Institute UFIC RAS Russia, 450080, Ufa, Mendeleev Street., 195, bld. 3. Email: [email protected]

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Palabras clave: Juniperus communis, Urales del sur, Cis-Urales, historia del asentamiento, población.

Introduction

The history of the formation and settlement of segments of the geological history. The few Juniperus communis in the Сis-Urals and in the fossil remains of some species of the Oxycedrus southern Urals is inextricably linked with the and Sabina sections are found in various areas of formation and settlement of pine forests in the range, the most ancient of which are found in general. According to the generalized reference Upper Cretaceous sediments (Gorchakovsky, data (Maleev, 1949; Sokolov et al., 1977; 1969; Ismailov, 1974). Mamaev, 1983), the juniper is mainly a companion of pine, pine-larch, and less often - Materials on the history of the settlement of dark coniferous forests in the Urals. The species juniper show that Juniperus communis had an is found in 54 forest types in the mountain forests extensive range covering most of North America of the Middle Urals, and at the same time prefers and Eurasia up to the Tertiary period of P. hyloccmiosa (Kolesnikov et al., 1973; Cenozoic; according to paleobotanical data, the Tishkina, 2009; Kozhevnikov, Tishkina, 2014). species is known from the Tertiary sediments of At the southern tip of the Urals and in the Altai, the Pleistocene sediments of North northern part of Kazakhstan, it is part of the America and the post-Pleistocene sediments of undergrowth of pine-steppe, or mixed with Northern Europe (see: Ismailov, 1974). The broad-leaved species. juniper habitat in glacial and interglacial epochs underwent repeated changes. In the postglacial In the Southern Urals, the species is located at the period, the juniper, moving after the retreating border of its distribution (The determinant…, glacier, restores large areas of the former range 1988), characterized by irregularity and in the north. According to M. I. Ismailov (1974), fragmentation of settlement. In the region under the modern juniper is either a “cold-resistant study, we identified several (Farukshina, mutant” that appeared in the Ice Age and later Putenikhin, 2012; Putenikhin, Farukshina, 2013) spread, or a pre-glacial old look, which “had relatively large habitats of Juniperus communis: hidden features” in the period of 1) in the plain pine forests (including open “cryophilization”. spaces) of the Belaya-Kama-Ufa interfluve in northwest and northern parts of the Bashkir Cis- This report analyzes the historical development Urals and the adjacent part of the Udmurt paths of the Juniperus communis in the Сis-Urals Republic (Nikolo-Berezovskaya, region and in the southern Urals. Dyurtyulinskaya, Amzinskaya, Maksimovskaya and Mazuninskaya cenopopulations); 2) in dark Material and methods coniferous forests in the central highly elevated part of the Southern Urals within the Republic of Within the Southern Urals and the Cis-Urals Bashkortostan and the Chelyabinsk Region (Belaya – Kama interfluve), 10 main (Katav-Ivanovsk cenopopulations); 3) in the cenopopulations were identified (Table 1). central part of the South Urals (Shigaevskaya, Geographically, the juniper research area covers Uzyanskaya, Burzyanskaya, Avzyanskaya the northwestern, northern, and northeastern cenopopulations). parts of the Bashkir Cis-Urals (with the adjacent southeastern part of the Udmurt Cis-Urals), the Paleobotanical materials for junipers are entire mountain belt of the Southern Urals currently extremely inadequate. Based on the (including the northern edge of the Zilair Plateau available data, it is very difficult to determine the in the south) and the territory of the Urals paths of settlement and the importance of adjacent to the east. junipers in the vegetation cover, in some specific

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Table 1. Test areas in juniper cenopopulation in the Urals and in the southern Urals

Cenopopulation Composition of the Geographical position (trial plot) three stand*

Bashkir and Udmurt Сis-Urals (Belaya-Kama-Ufa plain-hilly interfluve)

“Udmurt” Cis-Ural (south-east; right bank of the Mazuninskaya – Kama river)

Bashkir Cis-Ural (north-west; the interfluve of Amzinsky 9P1B+S rarely L the rivers Kama and Buy)

Maksimovskaya Bashkir Cis-Ural (northern forest-steppe part) –

Bashkir Cis-Ural (north-west; between the rivers Nikolo-Berezovskaya 9P1B Kama and Belaya)

Bashkir Cis-Ural (north-west; upper course of Dyurtyulinskaya 10P the Belaya River)

South Ural (mountain forest zone)

3F2S5P Katav-Ivanovskaya North of the central part (Katav River)

Shigaevskaya Central part (North Kraka range, eastern slope) 10P+L rarely B

Uzyanskaya Central part (Middle Kraká ridge, western slope) 10P rarely L

Avzyanskaya Central part (Bashtau ridge, southern slope) 2P8P rarely L

South central part (South Kraká ridge, western Burzyanskaya 10P slope)

Note. * The composition of the stand (on the test plot), to which the cenopopulation of Juniperus To study the population structure, a three-level communis is confined; - “forest-steppe” hierarchical sampling system was implemented cenopopulations growing in open space or edge; (Mamaev, 1973; Putenikhin et al., 2004, 2005): P - pine, B - birch, S - spruce, L- linden, F - fir, 1) sample areas in cenopopulations (one for the L - larch. cenopopulation), 2) randomly selected

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individuals in the test areas (15 female and 15 vegetative organs in the Сis-Urals and the males), 3) individual samples of generative Southern Urals (Farukshina, Putenikhin, 2016b). organs with 15 females and 1-3 vegetative Three phenotypically differing local biological sprouts from all 30 individuals. From each populations were identified (figure 1): Cis-Ural individual sample, 10-20 cones and seeds were forest (Amzinskaya, Nikolo-Berezovskaya and randomly taken (Mamaev, 1973; Putenikhin et Dyurtyulinskaya cenopopulations; all of them are al., 2004, 2005). 10 signs of generative organs cenopopulations growing under a canopy), Сis- and 7 morphological signs of vegetative organs Ural forest-steppe (Maksimovskaya and were studied. Mazuninskaya cenopopulations; confined to open habitats), South Ural Mountain Results and its discussion (cenopopulations of the mountain forest zone of the Southern Urals). Within the last population, Previously, on the basis of multidimensional two subpopulations were distinguished - forest methods, phenotypic differentiation of Juniperus (Avzyanskaya, Burzyanskaya, Uzyanskaya) and communis was assessed using a set of 17 forest edge (Shigaevskaya, Katav-Ivanovskaya morphological features of generative and cenopopulations).

Figure 1. Dendrogram of differences and similarities of coenopopulations of Juniperus communis based on signs of vegetative and generative organs

The cis-Ural forest population is completely the central part of the Southern Urals and is confined to the flat pine forests of the mainly associated with mountain pine and dark northwestern part of the Bashkir Cis-Urals. The coniferous forests. The forest subpopulation is Cis-Ural forest-steppe population is localized in represented by typically sub-forested locations, the north-west of the Bashkir Cis-Urals and in the and the forest edges subpopulation is represented south-eastern part of the Udmurt Cis-Urals. The by areas on open steppe slopes or forest edges. mountainous South Ural population is located in

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It was found that the mountainous South Ural forests in the region, especially in the Сis-Urals, population with a relatively increased has decreased significantly. Juniper squares intrapopulation phenotypic (genetic) diversity decreased in the wake of pine. In the western part according to hereditarily determined signs of the of the Republic of Bashkortostan, the juniper is a generative organs is at the same time less relic of the former boron vegetation (Putenikhin, variable, i.e. phenotypically more homogeneous Farukshina, 2013). The distribution of Juniperus in vegetative indicators. Cis-Ural populations are communis in the two parts of the habitat under characterized by an inverse pattern (Farukshina, consideration (in the Cis-Ural and in the southern Putenikhin, 2016b). Urals), apparently, was closely related to the Pleistocene-Holocene settlement of Scotch pine. It is possible to determine the possible ways of settlement in the Urals based on the history of the Given the above, the differentiation of the juniper settlement of pine forests (see above), of which it populations in the study area can be represented is a satellite. For the Scots pine during the as follows. The mountainous South Ural repeated Pleistocene glaciations, the Southern population was probably formed in the pre- Urals region became one of its refuge in Pleistocene and Pleistocene time as a result of the landscapes of larch-pine-birch forest-steppe settling of Scots pine from Siberia to the (Blagoveshchensky, 1943; Krasheninnikov, Southern Urals, as well as in the process of 1939; Panova, 1982; Filippova et al., 2006; Holocene expansion of the South Ural pine Frenzel, 1960). Akchagyl marine ingression had refuge. In the Holocene, as the climate is a significant influence on the change in the xerophytic, the juniper partly leaves the pine vegetation cover of the Cis-Urals at the end of the forests under the canopy to the edges and open Pliocene and the beginning of the Pleistocene, spaces, and, on the other hand, is introduced into when the region of the Belaya-Kama depression the composition of dark coniferous forests. The was flooded (Gorchakovsky, 1969; Fedorova, initial stages of differentiation of the South Ural 1970). Since that time, the pine forests isolated in population with the formation of an edge - forest the Cis-Urals have their own history subpopulation may be associated with these (Krasheninnikov, 1939; Popov, 1980). processes. Thus, the age of the mountainous southern Ural population of juniper is more than In general, the historical age of the South Ural 12 thousand years. Microevolutionary processes pine forests is significantly greater than that of aimed at the divergence of the mountain the pine in the main, more northern part of the population itself into subpopulations probably range (Frenzel, 1960). The forest vegetation of began no earlier than 1-2 thousand years ago and the plain Belaya-Kama interfluve (Сis-Ural), in so far have not led to the formation of particular, pine and spruce forests have an age independent local populations. Therefore, the not older than the end of the Pleistocene, and only local biological population represents possibly Holocene (Popov, 1980; Shalandina, Juniperus communis in the mountains of the 1998). If flat pine forests spread to this territory Southern Urals. from the northwest, mountain pine forests penetrated into the Southern Urals from Eastern As it was shown earlier with the phenotypic Siberia through the Kazakh low mountain characteristics of populations (Farukshina, (Gorchakovsky, 1969; Popov, 1980), therefore, Putenikhin, 2016b), according to some signs of these two groups of forests have not only vegetative organs, as well as the density of different ages, but also origin. individuals in cenopopulations, the mountainous South Urals population deviates towards the In the xerothermic phase of the Holocene (4.5- Siberian and northern populations of common 2.5 thousand years ago), warming and drying of juniper. According to the parameters of the the climate led to a shift of vegetation zones; generative organs, together with the сis-Ural Steppe vegetation, especially in the Cis-Ural populations, the mountainous South Ural region, strongly advanced to the north, increasing population is intermediate between Eastern the spatial isolation of the cis-Ural pine forests European and Siberian habitats. This may be a from the South Urals (Krasheninnikov, 1939; consequence of the processes of historical Igoshina, 1963; Gorchakovsky, 1969). At the end development of the species in the region, which of the Holocene, the range of pine trees in the we have mentioned above. region underwent more significant disjunctions not only due to the expansion of deciduous The cis-Ural populations of common juniper, forests and forest-steppe vegetation, but also due coenotic associated with relatively young pine to intensive anthropogenic activities (Popov, forests that penetrated into the Cis-Urals from the 1980). In the past two centuries, the area of pine northwest as the glacier receded, are probably of

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late Pleistocene or Holocene age (not older than determined morphological features of the 10–12 thousand years). Consequently, the generative organs of the mountain populations, formation of juniper populations in the Сis-Urals and in this respect, they are similar to each other. took place during a shorter time (in the post- It can be assumed that the pre-Ural forest-steppe glacial), moreover, at a distance from the South population in the xerothermic epoch was Ural part of the range. In the cis-Ural part of the distinguished from the originally single Early area, during the Holocene (as the climate Holocene population of the Cis-Urals. The warmed), intense microevolutionary processes juniper ordinary, still confined mainly to the pine took place, which led to the division of the forests of Kama and Belaya, began originally single cis-Ural population into two simultaneously to master the steppe areas that independent local populations — the forest and had advanced from the south. Under the action of the forest-steppe. Based on the time of intensive disruptive selection, a differentiation of the initial advance of the steppe and forest-steppe to the population occurred with the formation of the north (middle Holocene) (Gorchakovsky, 1969), forest-steppe cis-Ural population that was the age of the forest-steppe of the Cis-Ural phenotypically (and genetically) different from population can be approximately determined in the forest cis-Ural population. Due to this about 3-4 thousand years. separation, intrapopulation “genetic” variability could be reduced to a certain extent. Some indicators of cis-Ural populations — juniper density, signs of vegetative organs In the forest-steppe population in the process of (Farukshina, Putenikhin, 2012, 2016b) indicate a its development, the predominant role was certain phenotypic similarity with Eastern acquired by the driving selection, thanks to European populations of juniper (as already which it acquired strong phenotypic differences mentioned, the populations are intermediate from the original cis-Ural population. position between European and Siberian Interestingly, the cis-Ural forest-steppe populations). Perhaps this confirms the origin of population (Farukshina, Putenikhin, 2016b) is the cis-Ural populations from the Eastern distinguished by the highest level of variability European populations that settled along with the due to the labile traits of vegetative organs, as common pine after the retreating glacier. The well as by high shaped diversity (Farukshina, established nature of the population structure of Putenikhin, 2016a). In our opinion, this confirms common juniper, which corresponds to the the active action of a moving selection in it, “colonial type” (Grant, 1991), is consistent not leading to a shift in the phenotypic status of the only with the history of the development of pine population and the “chipping” of various forms forests, but, to a certain extent, with the according to the habit of the plants. phenotypic differentiation of the pine tree populations in the region (Putenikhin, 2000). In general, the colonial nature of the population structure of a species in the region (including the Estimation of intrapopulation variability showed disjunction of populations into more or less large (Farukshina, Putenikhin, 2016b) that the isolates) does not exclude the effect of gene drift mountainous South Ural population is in populations, especially сis-Ural, occurring characterized by a higher phenotypic (and against the background of other genetic) diversity compared to the cis-Ural microevolutionary factors. populations. This is consistent with the larger range of the mountain population. At the same Conclusion time, this indicates active processes of natural selection (probably stabilizing), maintaining a The South Ural mountain population has the high level of intrapopulational diversity and earliest origin (older than 12 thousand years), the providing a certain degree of stability of the most cis-Ural forest-steppe population is the species in mountain conditions (Schmalgauzen, latest (about 3-4 thousand years), and the cis- 1946; Timofeev-Resovsky, etc., 1969; Grant, Ural forest population is about the average 1991). However, the initial stages of divergence formation time (about 10-12 thousand years). of this population into separate subpopulations Populations differ in the level of intrapopulation indicate the intensification of other forms of diversity and the directions of microevolutionary selection (for example, moving and / or processes occurring in them. These differences, disruptive) in conditions of significant climate as well as the reduction of the ranges of warming in the modern era. populations throughout the second half of the Holocene, the formation of a colonial-type Сis-Ural populations are inferior in terms of the population structure determine the need to phenotypic variability of genotypically preserve the gene pool and the selective use of

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