The article presents the results of the study of fifty populations of common heather (Calluna vulgaris (L.) Hull) collected throughout its range. A phased comparative analysis (genetic, biochemical, anatomical, morphological, and ecological) was carried out with the estimation of indicators that included two key populations—Zavodouspenskoe (Pritobolye, Western Siberia) and Luga (Baltic, Eastern Europe). It was concluded that heather growing in Western Siberia should be identified as a separate taxonomic group, giving it the status of a subspecies. The gene pool of Pritobolye populations (including Zavodouspenskoe) is represented by the completely dominant (100%) monohaplotype S, which is not found anywhere else. The heather plant growing in Zavodouspenskoe has a longer lifespan. It is distinguished by larger linear leaf dimensions (length 2.06 ± 0.09 mm), thicker cuticle (4.77 ± 0.33 μm), increased number of trichomes (18.98 ± 0.56), and a reduced number of stomata (13.60 ± 0.63) than that growing in Luga. The new subspecies differs in biochemical composition: twice less content of epicatechin (average 1.992 ± 0.005 mg g−1), three times more myricetin (average 2.975 ± 0.005 mg g−1), twice as much chlorogenic acid (average 2.763 ± 0.004 mg g−1). An ecological feature is that C. vulgaris does not grow in the swamps of Western Siberia and has a small population. This species has a high horticultural potential and requires protection as its population in Western Siberia continues to decline rapidly.

Abstract. To preserve biological diversity, it is necessary to determine the genetic structure of species populations, because this gives the most complete picture of the manifestation of polymorphism and makes it possible to detect genetically and phenotypically distinct subspecies. The information obtained will help determine the optimal habitat and protection conditions for endangered and farmed species. The purpose of research is to conduct an allozyme analysis of 10 populations of P. sylvestris L. 2 of the physiographic countries of Yakutia and the Amur region. Results. An allozyme analysis revealed less polymorphism and the most drastic differentiation of the Amur region populations (at the level of geographic race) from Yakutia populations, subdivided from each other at the level of geographical populations. Significant differences are shown between the populations of Yakutia and Amur Region according to the distances of Mahalanobis, calculated according to 19 signs of cones. A positive linear relationship was obtained between the distances of Mahalanobis (D2) and the genetic distances of Ney (DN78) (R2 = 0.4159). Scientific novelty. A dominant role in the origin of the Yakut populations of P. sylvestris of three Siberian group-gradual fellings (GGF) - Tynda, Romanovka, Irkutsk and a decrease in the genetic similarity of the GGF as they move away from Yakutia were found.

Одной из главных проблем лесной экологии является анализ закономерностей динамики численности природных ценопопуляций древесных растений на этапе их возобновления, влияющем на последующую структуру и развитие биогеоценоза. Процесс возобновления определяется как динамикой семеношения, так и особенностями распространения семян лесообразующих древесных видов. Возобновление пятихвойных сосен подсекции Cembrae с бескрылыми семенами обусловлено тесными трофическими связями с птицами рода Nucifraga sp., которые, создавая почвенные кормовые запасы, разносят семена. Цель исследования – выявление географических особенностей связи между семеношением и возобновлением сосны кедровой сибирской (Pinus sibirica Du Tour) при посредническом участии тонкоклювой кедровки (Nucifraga caryocatactes macrorhynchos Brehm C.L.). Исследование выполнено в 3 географических регионах произрастания кедровых лесов: в лесах Алданского нагорья в Южной Якутии (59º40´ с. ш. 125º24´ в. д.), Северного Урала в районе горного массива Денежкин Камень (60º25´ с. ш. 59º32´ в. д.) и горного хребта Хамар-Дабан в Юго-Восточном Прибайкалье (51º32´ с. ш. 103º32´ в. д.). Показано, что каждый регион ареала произрастания P. sibirica отличается динамикой и семеношения, и возобновления. Установлено, что в лесах Алданского нагорья обилие всходов P. sibirica положительно связано (R2 = 0,46) с урожаем шишек прошлого года. Количество всходов на Денежкином Камне и Хамар-Дабане отрицательно связано с урожаем прошлого года, но положительно (R2 = 0,48–0,49) – с урожаем позапрошлого года. Мы считаем, что всходы в любом случае появляются из семян урожая прошлого года, а видимая связь с урожаем позапрошлого года обусловлена деятельностью кедровки. В результате проведенных стационарных исследований на Северном Урале выявлена постоянная тесная (R2 = 0,81) положительная связь количества всходов P. sibirica с численностью кедровки в период запасания и разноса семян в августе–сентябре предыдущего года. Сделано предположение, что характер динамики урожая семян в регионе влияет на изменение численности кедровки в период заготовки ею семян, которая, в свою очередь, определяет обилие всходов P. sibirica следующего года. Это обуславливает выявленные различия в динамике возобновления P. sibirica и ее зависимости от урожаев шишек в рассматриваемых регионах. Для цитирования: Танцырев Н.В., Санников С.Н., Усольцев В.А. Географические особенности возобновления Pinus sibirica Du Tour // Изв. вузов. Лесн. журн. 2023. № 6. С. 44–56. https://doi.org/10.37482/0536-1036-2023-6-44-56

One of the main problems of forest ecology is to identify the patterns of the dynamics of natural populations of woody plants at the stage of their renewal, which programs the entire subsequent structure and development of plant ecosystems. The process of reforestation is determined by both the dynamics of seed production and the peculiarities of the seed distribution of forest-forming tree species. The renewal of five-needled pines of the Cembrae subsection with wingless seeds is due to their close trophic relationships with birds of the genus Nucifraga sp., which spread seeds, creating soil forage stocks. The purpose of this study is to reveal geographical features of the relationship between seed production and renewal of Siberian stone pine (Pinus sibirica Du Tour) with the intermediary participation of the thin-billed nutcracker (Nucifraga caryocatactes macrorhynchos Brehm C.L.). The research was carried out in 3 geographical regions of Pinus sibirica forests: in the forests of the Aldan Highlands in Southern Yakutia (59º40´ N., 125º24´ E.), the Northern Urals in the area of the Denezhkin Kamen mountain (60º25´ N., 59º32´ E.), and the Khamar-Daban Mountain range in the South-Eastern Baikal region (51º32´ N., 103º32´ E.). It is shown that each region of the Pinus sibirica growth differs in the dynamics of both seed production and renewal. It was found that the abundance of Pinus sibirica seedlings in the forests of the Aldan Highlands is positively associated (R2 = 0.46) with the yield of cones the previous year. The number of seedlings in the Northern Urals and Khamar-Daban is negatively related to the yield of cones of last year but is positively (R2 = 0.48–0.49) associated with the yield of cones of the year before last. We believe that seedlings in any case appear from the seeds of last year's yield. But the visible connection is due to the activity of the nutcracker. As a result of the stationary studies conducted in the Northern Urals, we identified a constant close (R2 = 0.81) positive relationship between the number of Pinus sibirica seedlings and the number of nutcrackers during the period of seed storage and dispersal in August-September of the previous year. It is assumed that the nature of the dynamics of seed production in the region affects the change in the number of nutcrackers during the seed harvesting period, which in turn determines the abundance of Pinus sibirica seedlings next year. This determines the revealed differences in the dynamics of Pinus sibirica renewal and its dependence on the yields of tree cones in the regions under consideration. For citation: Tantsyrev N.V., Sannikov S.N., Usoltsev V.А. Geographical Features of Pinus sibirica Du Tour Renewal. Lesnoy Zhurnal = Russian Forestry Journal, 2023, no. 6, pp. 44–56. (In Russ.). https://doi.org/10.37482/0536-1036-2023-6-44-56

A comparative allozyme analysis ofPinus sylvestrisL. populations distributed over the entire species range has been performed to estimate Nei's minimum genetic distances among eight phylogeographic population groups (PPGs) from the "glacial" zone and 26 populations from the southern "extraglacial" zone. On this basis, the locations of nine main and eight second-order hypothetical Pleistocene refugia (HPRs) for the species have been determined. Using the conditional "refugium investment index," relative contributions from the gene pools of HPRs to those of PPGs from the glacial zone has been estimated. It is shown that populations of this zone have originated from several refugia. A classification of Pleistocene refugia forP. sylvestrisis proposed, which is based on their location within the range and role in the formation of the current gene pool of the species.

The history of the interaction between the boreal floras of northern Eurasia and those from the mountain areas located at intermediate latitudes remains insufficiently studied. Twenty-four population samples of 4 to 34 trees were analyzed with eight mitochondrial DNA (mtDNA) markers and four chloroplast DNA (cpDNA) microsatellite loci to study the history of Scots pine in the Black Sea region, and infer migration routes and genetic relationships with populations of the northern part of the range. These new data were combined with data previously obtained from Eastern Europe and North Asia. Scots pine past geographic range was also reconstructed using environmental niche modeling. Both mtDNA and cpDNA revealed a common origin of populations in the eastern Black Sea region (EBS), i.e., Crimea, Caucasus, and Asia Minor, and deep divergence from the main range of Scots pine, reminiscent of the isolation of enclaves in Iberian and Apennine Peninsulas. Mitochondrial DNA haplotypes (mitotypes) endemic to Western Caucasus are genetically intermediate between mitotypes from Eastern Europe and other mitotypes from the EBS region, indicating a recombinant origin likely due to migration to the Caucasus from the north and hybridization with local trees. Environmental niche modeling shows the possibility of such migration events during the last glacial maximum.

Simulation of fire impact on forest floor renewal in pine forests was based on predictions for 2100 by the Canadian Climate Centre for increase in temperatures of by 4.5 °C and total precipitation by 14% in the West Siberian pre-forest-steppes and empirical regression relationships between them and the degree of burnt forest floor, as well as amount of Pinus sylvestris L. regeneration. It was predicted that by 2100, pine regeneration on fire prone sites under the canopy of the dominant forest type will increase by 29–54% compared to the 1980s but on adjacent open sites, regeneration will decrease twice as much. This means that the regeneration potential and pine population stability in pre-forest-steppes will become as poor as it is today.

The structure and seed bearing of stands, factors of episoil medium, quantity, vitality and age structure of Scots pine undergrowth 15 years after the disastrous crown and surface fire, which occurred in 2004 in insular pine forest in northern forest-steppe of Western Siberia (Kurgan region), were studied. The studies took place at six sample plots (0.3–0.5 ha) located in two predominant (88 %) types of pine forests – cowberry-short grasses-moss and bilberry-bracken – at four types of burns: 1) «burn under the forest canopy», 2) «thin stand burn», 3) «open burn with peripheral insemination» and 4) «open burn without insemination». The reliable differences in pine regeneration in different types of burns in both types of forests were specified. They were conditioned mainly by differences in their stand density and dissemination levels. The total pine regeneration level in forest-steppe was several times less than in pre-forest-steppe. This was connected with extreme dryness of the studied climate period and low seeding of stands, damaged by the intensive fires. The renewal role of dissemination of 2003 which took place simultaneously with the fire; absence of mouse-like rodents (main seed consumers), as well as sharp change of sex ratio in pine populations (to female dominance) due to intensive surface fire have been shown. On the whole, the total summer precipitation played the main role in pine regeneration of all burn types in forest-steppe zone of Western Siberia.

The aim of the research was to determine the origin of an isolated Atlantic Azorean population of heather (Calluna vulgaris (L.) Hull). The chloroplast DNA analysis showed it possessed genetic affinities with 4 populations of the Dinaric-Pontic group in the extra glacial zone of the Mediterranean area as well as with a population from the Atlantic coast (Coimbra) and a slight similarity to 5 other heather populations in this area. A hypothesis of the ancestral center of the Azorean heather population origin in the Dinaric-Pontic region and the proximate center of its ornithochoric dispersal to the Azores from the west of the Iberian Peninsula (Coimbra) has been proposed.

The dates of appearance of a representative portion of Pinus sylvestris L. pollen in palynospectra (≥20%) have been analyzed as a parameter reflecting the spread of leading-edge pine populations from central and eastern Europe to the Lofoten region via probable alternative routes of population immigration. It has been found that they could reach this region not earlier than 6500–7000 years BP, i.e., 2500–3000 years later than the factual palynological date, even at the maximum possible rate of their spread (300–500 km/1000 years). Therefore, the probability of the origin of Lofoten P. sylvestris populations from adventive populations that migrated to Scandinavia from adjacent European regions is excluded. The generalization of the results of studies on the chronopalynology and rate of Holocene expansion of P sylvestris and data on paleoglaciology, paleomarinology, paleobotany, and modern genogeography and ecology of Scots pine has confirmed the hypothesis about the survival of autochthonous refugial populations of this species on the Lofoten Islands during the last glacial phase and their expansion during the Holocene.

As a result of half a century (1970–2017) studies of recovery and age-related dynamics of different types of pine forests in foreforest-steppe and other subzones of Western Siberia and the Urals we have worked out a concept and scheme of their ecologic and genetic classifications based on ecologic dynamic series of biogeocenoses development (EDS BGC) within native autochthonous types of forests. Unlike the monolinear variation of biogeocenoses in course of time within one type of forest site and native forest type in classical genetic Ivashkevich-Kolesnikov typology, we consider the native type of forest as a «bunch» of divergence-convergence of chorologically and ecologically separated types of EDS BGC arising after different forest ecological catastrophes, such as fires, windfalls, pest invasions, tree fellings, stubbing, ploughing up, etc. The main real form of existence and development and syntaxonomic unit of adequate ecological-genetic classification of forest types is the type of EDS BGC as a homogeneous series of their restoration-age-related and century-long plant successions within one native forest type as an ecosystem of higher rank. The general scheme of forest classification based on the EDS BGC as well as dynamic parameters of their structure and diagnostics, are exemplified by predominant forest types relating to moss pine forests in the foreforest-steppe of the Western Siberia. The proposed approaches and methods of chorological ecological-genetic classification of forest types can be applied in forest science and forest management to get a lot of new systematic quantitative information on forest dynamics for its evaluation, development projection and forest management.

The results of molecular-genetic studies on the Scots pine cenopopulations from the isolated gorges of the Central Caucasus are given. The quantitative assessment of genetic differentiation degree for the Scots pine cenopopulations from different altitudes in the area under survey has shown reliable differences between the studied samples of subpopulations and local populations by means of Nei distances.

The distribution of nine chloroplast DNA haplotypes in four insular North-Atlantic and four European coastal Atlantic populations of Calluna vulgaris in the glacial zone of the range has been analyzed in comparison with that in six marginal southern populations in the nonglacial zone of the Atlantic and Mediterranean regions. As a result, two hypothetical Pleistocene refugia (HPRs) for this C. vulgaris population group have been revealed, one in the Cévennes mountain range and the other in the Southern Alps (Trento). Judging from the 1–FST value, it has been found that the group of populations in the glacial Atlantic zone and adjacent European coastal Atlantic region is genetically similar to the HPR in the Cévennes at a highly significant level (p ≥ 0.999) and less similar to the HPR in Trento; however, it differs significantly from other Mediterranean and Atlantic populations. It has been concluded that the most probable hypothetical Pleistocene refugium for the recent C. vulgaris populations of the northeastern Atlantic and European coastal Atlantic regions was in the west of the Mediterranean, in the Cévennes, while the additional refugium was in the Southern Alps. Possible directions of the postglacial dispersal and recolonization of habitats by C. vulgaris populations from the western Mediterranean to the northeast of the Atlantic and to Scandinavia have been revealed.

В итоге сравнительного анализа распространения 9 гаплотипов хлоропластной ДНК в четырех островных северо-атлантических и четырех европейских приатлантических популяциях Calluna vulgaris ледниковой зоны ареала, с одной стороны, а также в шести маргинальных южных популяциях внеледниковой зоны Атлантики и Средиземноморья, с другой, выявлены два гипотетичных плейстоценовых рефугиума (ГПР) этого вида. По параметру 1-FST установлено, что изучавшаяся группа популяций ледниковой зоны Атлантики и смежной европейской Приатлантики на высоко достоверном уровне (р 0.999) генетически сходна с ГПР в Севеннах, в меньшей мере с ГПР в Тренто, но резко отличается от других тестированных популяций Средиземноморья и Атлантики. Сделан вывод о том, что наиболее вероятный гипотетический плейстоценовый рефугиум для современных популяций C. vulgaris северо-восточной Атлантики и Приатлантики располагался на западе Средиземноморья, в горном массиве Севенны, а аддитивный - в Южных Альпах (Тренто). Выявлены вероятные направления пост- гляциальной реколонизация популяций C. vulgaris из западного Средиземноморья на северо-запад Атлантики и в Скандинавию.

A new method for determining the upper forest border (UFB) as the border of the “root-closed” stand and regrowth as its potential vanguard is proposed. The mass dispersal of Pinus sibirica populations by the nutcracker ( Nucifraga caryocatactes ) in the UFB ecotone in the Northern Urals (Pavdinskii Kamen’ and Tretii Bugor Mountain) from the middle-mountain taiga to the mountain forest tundra and tundra at a distance reaching 1 km is found. It is done based on the original reconstruction method of the dynamics of population number in P. sibirica annual seedlings and on an analysis of their relations with the dynamics of the seed bearing and population number of the nutcracker. The trustworthy consortive relations of the number of Pinus sibirica generations in the mountain forested tundra and tundra zones, with its seed bearing and nutcracker number in the previous year in the middle-mountain taiga zone, are found. A significant increase, 3.7 times, in the number of seedling generations during the last 49 years is found in connection with the increase in the summer air temperature in the subarctic to 2.0°C. A hypothesis of the formation of the united genetic metapopulation of P. sibirica in the forest–tundra mountain ecotone zone as a consequence of its seed ornitochory from the highly different populations is formulated. A forecast of the mosaic formation of the vanguard “thin forests” in 20–25 years and the root-closed curtains of the forest in 40–50 years in the mountain tundra of the Northern Urals at the current rate of the rise in temperature is made.

As a result of allozyme analysis, Nei’s genetic distances were determined between the phylogeographic group of seven populations of Pinus sylvestris L. in the “glacial” zone of the range in Central Yakutia and 25 populations of its hypothetical Pleistocene refugia of the southern nonglacial zone within the entire range of the species in Northern Eurasia. The location of the most likely “avant-garde” refugium of the modern yakutian populations of the Scots pine is Northern Priamur’e (Tynda), as well as less likely Siberian (Romanovka, Irkutsk), South Ural (Kryktytau), and Central European (the Czech Republic) refugia.

The microevolution theory can serve as the theoretical foundation of forest seed zoning and of the population introduction, the subdivision of Pinus sylvestris range in Russia into 10 phylogenogeographical regions – as their genetic-geographical base and climatologic-geographical regularities of the acclimatization of the introduced crops – as the ecological base. As a result of regressive analysis of Pinus sylvestris provenance crops parameters in Russian Plain forest-steppe is was determined the authentic decrease of the parameters of their survival and stands production, according to the increase of gradients of climate limiting factors – duration of vegetative period and photoperiod – between sites of origin and introduction of populations both along of the submeridional transect and of the longitudinal ones. Thus, the hypothesis about the greater adaptivity of the local plant populations by comparison with introduced ones and the correctness of the principle of «climatic topoanalogues» in the acclimatization were confirmed on the quantitative factorial-climatological level. The total net of geographical coordinates and of corresponding elementary forest seed regions measuring 1° in latitude and 5° in longitude formed by it was suggested as the ordination matrix of forest seed zoning within the P. sylvestris areal. The parameters of seed transfer reglamentation in latitude, longtitude and altitude were suggested on the connections base of survival and stands production of introduced Scots pine crops with climatic gradients. These principles can serve as a genetic-climatologic-geographical base for the development of the theory of plants acclimatization and of regional and federal seeds zoning of Scots pine forests for their crops, selection and hybridization.