ISSN 2079 0864, Biology Bulletin Reviews, 2013, Vol. 3, No. 6, pp. 493–504. © Pleiades Publishing, Ltd., 2013. Original Russian Text © O.V. Smirnova, D.V. Lugovaya, T.S. Prokazina, 2013, published in Uspekhi Sovremennoi Biologii, 2013, Vol. 133, No. 2, pp. 164–177.
Model Reconstruction of Restored Taiga Forest Cover O. V. Smirnova, D. V. Lugovaya, and T. S. Prokazina Center for Problems and Ecology and Productivity of Forests, Russian Academy of Sciences, Moscow, Russia e mail: [email protected]
Abstract—A model reconstruction of the potential forest cover of dark coniferous taiga on the Russian Plain, the Urals, and some territories of Western and Central Siberia is presented. The dot area of tallherb dark coniferous forests has been constructed based on 850 geobotanical releves (from the Database of the Center for Problems and Ecology and Productivity of Forests, Russian Academy of Sciences). The dot area of tall herb dark coniferous forests has been compared to that of Tilia cordata and contemporary zonality of the studied area. The ecotopes and ecological characteristics of the tallherb forests have been determined. Con stant species of the tallherb forests have been revealed. Based on the literature data and the created database of herbaria labels, refined ranges of tallherb species were constructed. The importance of studying tallherb dark coniferous forests has been assessed to determine the origin of the dark coniferous taiga and select virgin forests for evaluating the main functions of the ecosystem.
Keywords: potential cover, ecosystem functions, dark coniferous taiga, tallherb forests, ranges of constant tall herb species DOI: 10.1134/S207908641306008X
INTRODUCTION along with the paleobotanical and florogenetic meth ods. The analysis of structural and taxonomic variety A vital task of modern natural resource manage of refugium vegetation allows us to define the natural ment is its reorientation towards preservation and res characteristics of vegetation cover and form an opin toration of ecological functions of ecosystems, ion as to the genesis of this vegetation type, while com namely, climate regulation, water and soil protection, paring the obtained results with the paleobotanical productional function, and maintaining the biovariety. and florogenetic data. It was proved earlier that the fullest realization of eco logical functions can be accomplished in a restored So far, plenty of data has been assembled on the forest cover, i.e., in this cover, which can form as a composition and structure of taiga forests communi result of a long spontaneous development (without ties in refugia on the Russian Plain and the Urals, as anthropogenic disturbances and natural disasters) and well as in some regions of Western and Central Siberia. is represented by all species preserved so far According to the studies performed, they are passing (Smirnova, 2004; Smirnova et al., 2006a, 2006b). Nat through the late successional or quasiclimax stage and are ural releves and/or model reconstructions of such characterized by the following features (Turubanova, cover will allow us to assess the ability of the forest 1999, Yaroshenko, 1999; Bobrovskii and Khanina, 2000; cover to fulfill its ecological functions under different Smirnova et al., 2001a, 2011; Yaroshenko et al., 2001; climate conditions. According to the today’s beliefs, Vostochnoevropeiskie lesa…, 2004; Smirnova, 2004; the taiga forests (boreal and hemiboreal) of the North Zaugolnova et al., 2009; Bobrovskii, 2010): ern Hemisphere have preserved their natural composi (1) the presence of dark conifer trees (Picea, abies, tion and ability to fulfill the main ecosystem functions P. obovata, Abies sibirica, and Pinus sibirica), broad to the fullest extent possible (Monitoring…, 2008). leaved trees (Tila cordata, Ulmus laevis, and U. scabra Therefore, the thorough study of these forests is a mat et al.), and small leaved trees (species of the genus ter of theoretical and practical importance. Moreover, Betula and Alnus, Populus tremula, and Padus avium a model reconstruction of the restored taiga forest et al.) in the forest stand of the studied area; cover represents another step towards the solution of problem of origin and development of dark coniferous (2) unevenly aged cenopopulations of late succes taiga. sional tree species; As was rightly mentioned by V.B. Sochava (1946), (3) the development of the entire set of micromo the studies of refugia, i.e., areas in which vegetation saic structural elements: intercrown and undertree has barely been touched by natural disasters and pow spaces of different tree species, elements of windfall erful anthropogenic impacts, play a considerable role soil complexes (treefall soil pits, knobs, debris, and in dealing with the problem of vegetation genesis, stumps) at different stages of development;
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(4) presence in ecosystem of both vascular plants them to be representatives of taiga forests above all species and mosses of all eco coenotic groups capable (Zaugolnova et al., 2009). to inhabit the studied forests; According to the International Code of Phytoso (5) the dominance of boreal and nitrophilous tall ciological Nomenclature (Weber et al., 2005), the herb in the soil cover in terms of spreading and phyto studied tallherb forests have the following syntaxo mass; nomic position: (6) the absence of traces of fire and cutting in vege Class Vaccinio Piceetea Br. Bl. in Br. Bl., Sissingh tation cover, and coal in soil; et Vlieger 1939; (7) undifferentiated soil profile represented by Order Picetalia excelsae Paw lowski in Paw lowski, humus horizon developed as a result of recurrent soil Sokolowski et Wallisch 1928 (=Vaccinio Piceetalia trenching due to the late successional tree fall. Br. Bl. in Br. Bl., Siss. et Vlieger 1939); The above mentioned features are most pro Alliance Vaccinio Piceion Br. Bl., Sissingh et nounced in quasiclimax forests, whereas the late suc Vlieger 1939; cessional forests have not yet developed these charac Suballiance Atrageno Piceenion obovatae Zaugol teristics (The Population Structure …, 1985; The nova et al., 2009; Mosaic Cycle Concept …, 1991). This is most strongly Assoc. Aconito septentrionalis–Piceetum obovatae translated by the floristic incompleteness of commu Zaugolnova et al., 2009; nities caused by the considerable gaps in tree ranges Subassoc. A. s.–P. o. typicum Zaugolnova et al., due to the anthropogenic activities. A good example is 2009; the considerable gaps in Tilia cordata Mill. range in Subassoc. A. s.–P. o. filipenduletosum Zaugolnova the southern taiga subzone documented in the historic et al., 2009. literature (Vostochnoevropeiskie lesa …, 2004; Bakun, Communities of tallherb dark coniferous forests 2006; Bobrovskii, 2010). From a physionomical point are revealed on the watersheds in well drained habitats of view, the late successional and quasiclimax dark and in valleys with small rivers and brooks where there coniferous forests are characterized by the dominance is a flowing hydration. Diagnostic species of the entire of tall boreal herbs of two ecocoenotic groups, i.e., association are as follows: Aconitum septentrionale boreal tallherbs and nitrophilous tallherbs. The herbs Koelle, Veratrum lobelianum Bernh., Chamaenerion of these groups grow quickly and are capable of angustifolium (L.) Scop., Calamagrostis purpurea achieving a height of 1.5–3.0 m during growing (Trin.) Trin., and Thalictrum minus L. The watersheds period; the majority has either large leaves or a big forests belong to the subassociation A. s.–P. o. typi number of shoots with medium sized leaves in large cum, the diagnostic species of the entire association tufts (grasses). The dense canopy of these herbs, as being joined by Diplazium sibiricum (Turcz. ex well as the considerable herbaceous litter prevents the G. Kunze) Kurata; valley forests belong to the subas growth of boreal green mosses (Smirnova et al., 2011). sociation A. s.–P. o. filipenduletosum, within which a The phytomass of the annually falling aboveground group of diagnostic species was defined that included shoots of the vascular plants in tallherb forests is five to Filipendula ulmaria (L.) Maxim., Geum rivale L., Trol ten times higher than the phytomass of vascular plants lius europaeus L., etc. in the most widespread taiga forests, namely, those of Tallherb forests differ from those of green moss and a green moss and sphagnous type. The fall of the tall sphagnous types by the more complicated structure of herb species is highly enriched with mineral elements layers, unevenly aged tree cenopopulations, and the compared to that of the herbs and dwarf shrubs of highest species and ecocoenotic variety. The trees in green moss and sphagnous forests (Lukina and these communities are large and well developed, and Nikonov, 1998; Lukina et al., 2006). It is quickly their deaths are followed by pedoturbations. Conse absorbed by the soil biota, which explains the high quently, it is possible to preserve the mosaic of restora velocity of mineral elements turnover and advanced tion windows that determines the mosaicity of ground forest productivity. Tallherb species are light demand cover light regime, as well as the mosaic of windfall– ing, but at the same time tolerant to a lack of light, i.e., soil complexes that accounts for the mosaicity of they withstand the provisional reduction in light microhabitats with different regimes of temperature, intensity and quickly return to their former status due moisture, soil acidity, etc. to the light window. The herbaceous layer of these forests is dense (70– The tallherb species are different in terms of their 90% spreading), polidominant, and includes at least dependence on ecological factors (Tsyganov, 1983); three sublayers. The first sublayer is formed by the this explains the dominance of different members of boreal and/or nitrophilous tallherbs; the second one is this group under different regimes of moisture, acidity, formed of boreal, nemoral, nitrophilous, and and soil richness. medium sized meadow and edge herbs; the third one The ranges of majority of species of this group is formed of small nitrophilous and boreal herbs and reach the north forest frontier and penetrate even a lit dwarf shrubs. The moss cover is not dense (20–10% tle further to the north, which leads us to consider spreading), but consists of a large number of ecologi
BIOLOGY BULLETIN REVIEWS Vol. 3 No. 6 2013 MODEL RECONSTRUCTION OF RESTORED TAIGA FOREST COVER 495 cally different species. The alluvial soils with moder herb dark coniferous forests was constructed based on mull horizon 35–50 cm in depth are formed in tall the geographic coordinates of releves determined by a herb floodplain forests. The brown soils with moder GPS receiver. To compare the obtained dot area of the mull humus horizon reaching a depth of 60 cm are tallherb dark coniferous forests with the modern zonal formed in tallherb watersheds forests (Smirnova et al., division of studied area, we used a map (Karta “Zony i 2006c; Bobrovskii, 2010). tipy ...,” 1999) with some modifications. The maps To date, the accumulated data on ecology and dis were created using ArcMap 9.2 (ESRI Inc. software tribution of the tallherb species, successional status of product). To compare the dot area of the tallherb dark the tallherb dark coniferous forests, their ecotopes, coniferous forests with the modern outlined range of and geographical range allowed us to undertake recon Tilia cordata Mill. and its dot area in the late struction of the restored forest cover of studied area. Holocene, we used data kindly furnished by For this purpose, the following tasks have been car S.A. Turubanova. Geobotanical releves and thematic ried out: maps of quaternary deposits, landscape types, and geological structures of different scale were used to 1. The dot area of the tallherb dark coniferous for analyze the correspondence of the tallherb dark conif ests was constructed and a comparison of their distri erous forests to the relief elements and superficial bution with the modern zonal division of the studied deposits (Atlas…, 1964a, 1964b, 1967, 1968, 1971, area was performed. 1976, 1989, 1999a, 1999b, 2001; Ekologicheskii…, 2. Ecotopes of tallherb forests were identified. 1999). The maps from the published atlases were 3. A comparison of the tallherb forests’ range with anchored to the ArcMap 9.2 environment where the the modern outlined range of Tilia cordata and its dot space analysis of information was conducted. area in late Holocene has been performed. The direct ordination of geobotanical releves by 4. The mean point values of several climatic and environmental indices was performed using ecological ecotypic indices that characterize the communities of scales by Tsyganov (Tsyganov, 1983). The calculations tallherb dark coniferous forests were estimated using of the mean point values by indices of winter severity the ecological scales (Tsyganov, 1983); they were then (Cr), climate warmth (Tm), climate moisture (Om), compared with the range evaluations of the same indi and soil richness (Tr), as well as the analysis of geobo ces for Tilia cordata being the supposed inhabitant of tanical releves were performed using the PcOrd pack these forests. age of SpeDiv 1.3.0 software (developed by 5. The peculiarities of geographical range of boreal V.E. Smirnov). and nitrophilous tallherb species considered the indi To identify the peculiarities of geographical range cators of the late successional and quasiclimax devel of boreal and nitrophilous tallherb species that indi opment stages of dark coniferous forests were revealed. cate late successional and quasi climax dark conifer 6. Based on the data on successional status and dis ous forests, the range maps of these species taken from tribution peculiarities of the tallherb dark coniferous the literature data (Meusel et al., 1978; Hulten and forests, the theories of dark coniferous taiga origin and Fries, 1986) were scanned and digitalized. To apply the development have been analyzed. dots of collecting herbarium specimens, the study of large herbaria of the universities (Moscow State Uni versity, Kazan State University, and Tomsk State Uni MATERIALS AND METHODS versity) and academic institutions (Komarov Botani To carry out the set tasks, 850 releves from the Geo cal Institute, Russian Academy of Sciences; Institute botanical Database of the Center for Problems and of Plant and Animal Ecology, Ural Branch, Russian Ecology and Productivity of Forests, Russian Academy Academy of Sciences; Komi Science Center, Ural of Sciences, performed on the square grounds 100 m2 in Branch, Russian Academy of Sciences; Central Sibe size, were analyzed according to the method published rian Botanical Garden, Russian Academy of Sciences, earlier (Smirnova et al., 2002b) in the Murmansk, et al.) was carried out. The geographic coordinates of Leningrad, Arkhangelsk, Kostroma, Yaroslavl, Kirov, discovery points were established using the labels data Vologda, Sverdlovsk, and Tyumen’ oblasts; Perm and and then added to the GIS. The database preserves the Krasnoyarsk krais; the Republics of Karelia and Komi; information on points of discovery of tallherb species and the Khanty Mansi and Yamalo Nenets autono (7180 records), along with their coordinates. The cor mous okrugs. The analyzed releves were made in rection of meridional groups of these species’ ranges mountainous and elevated areas, as well as rises of was performed using this data. folded or glacial origin, including the Baltic Shield; Andomsk Upland; Belozersko Kirillovka Upland; the Northern Ridge; the Timan Ridge; the Siberian Ridge; RESULTS AND DISCUSSION Kondo Sos’vinsk Priob’e; Western and Eastern Urals To date, tallherb dark coniferous forests have been macroslopes; and on the plain areas, including the revealed in the largest part of the Russian Plain and the Belomorsko Kuloiskoe Plateau, Mezensk Plain, and Urals, as well as in some areas of Western and Central Volkhovo Ilmen Lowlands. The dot area of the tall Siberia (Fig. 1). The irregularity in the distribution of
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1 2 3 4 5 6 0 500 1000 2000 km 7 8
Fig. 1. Dot area of the tallherb dark coniferous forests of the Russian Plain, the Urals, and some areas of Western and Central Siberia compared to the modern outlined range of Tilia cordata and its dot area in late Holocene: 1, location dots of the tallherb forests, 2, location dots of Tilia cordata pollen in Holocene, 3, Tilia cordata outlined range. Natural subzones and zones: 4, forest tundra, 5, northern taiga; 6, middle taiga; 7, southern taiga and subtaiga, 8, mountain taiga forests. tallherb forests within the studied area is primarily nova et al., 2008). The only exception is the taiga caused by the fact that they survived in places least reserves of the Ural slopes where the tallherb forests affected by anthropogenic disturbances. They were area reaches dozens of hectares in the middle and found to the west of the analyzed area in poorly acces upper slopes (Smirnova et al., 2011). The small num sible regions of North Karelia and the Kola Peninsula: ber of locations in Western Siberia is caused by the fee Lapland Biosphere Reserve and Paanajárvi National ble preservation of dark coniferous forests and poor Park and its environs (Smirnova and Korotkov, 2001). knowledge on the subject. One of the most interesting They were also revealed at the heart of the Russian objects is the tallherb spruce fir forests in the Gorno Plain, in the Pinega Nature Reserve, Kologriv Forest, Khadytinskii Reserve of the Southern Yamal (southern Kilemary Reserve of Nihzni Novgorod oblast (Suktse tundra region), which are similar in all characteristics sionnye…, 1999; Shirokov et al., 1999, 2001; Otsenka i to the northern taiga tallherb forests (Smirnova et al., sokhranenie…, 2000; Shirokov, 2002, Lugovaya, 2010). 2008). The tallherb forests were identified in the low hill ter The overview of the tallherb forests locations per rains of nearly all taiga reserves of the Ural western formed using the field survey data and the published macroslopes: Visim, Vishersk, Denezhkin Kamen’, cartographic materials (Atlas…, 1964a, 1964b, 1967, Pechoro Ilych, Malaya Sos’va (Yaroshenko et al., 1968, 1971, 1976, 1989, 1999a, 1999b, 2001; Ekolog 1998; Suktsesionnye…, 1999; Prokazina, 2007, 2011; icheskii …, 1999) allowed us to draw up a list of the Zaprudina and Smirnov, 2010). Surely, the systematic main relief elements, as well as types of quaternary searches of tallherb dark coniferous forests in the areas deposits and parent rocks where the tallherb forests where they have not yet been found may considerably were found. The interfluve and valley complexes were expand this list; however, this presents significant dif analyzed separately (Table 1). In both complexes, the ficulties. It is worth mentioning that the majority of tallherb forests can only be found in the drained areas; the studied tallherb communities have rather small communities of typicum subassociation can be found areas, i.e., from one or less to several hectares (Zaugol in moderately humid areas; and communities of fili
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Table 1. Characteristics of habitats of tallherb dark coniferous forest communities Characteristics of habitats Interfluve complex Valley complex Elements of relief (based on the Leveled interfluve area, interfluvial slope Valley, valley side, first terrace above field survey data) (incl. swamped areas); closed positive and floodplain, high floodplain (ridges, negative relief forms in the interfluve area depressions, subhorizontal surfaces, (incl. karstic ones); lake terrace and natural levees); bottom of the brook valley Quaternary deposits and parent Quartzites, quartzite sand, Alluvial (sands, sand clays, clay loams, rocks based on the literature data clay–limestone deposits; gabbroids and peat interbeds, and lenses; sandy–clay granitoids intrusions; amphibolites and and peat like); Valdai and Moscow gneisses; morainic and fluvioglacial deposits glacial deposits: fluvioglacial (clays, clay loams, boulder clays interlaid (pebblebeds and sands; lake and glacial with inequigranular sands, sands); basic and (sands and clays); glacial (boulder clay ultrabasic rock complexes; red marl beds and loams) sandstones Parent rocks (based on the field Light, middle, and heavy clay loams (incl. Alluvial clay loams, sand clays; crypto survey data) morainic ones); sand; rocky sand podzolic sand clays on clay loams; (with pebble); limestones; quartzites, karsting rocks granites, and diabases penduletosum subassociation can be found in humid ern tallherb taiga forests becomes a matter of consid and over humid areas, provided the existence of flow erable interest with regard to the model reconstruction ing hydration. of the restored forest cover. The diversity of quaternary deposits and parent The modern continuous area of Tilia cordata over rocks inhabited by the tallherb forests is huge and rep laps the dot area of tallherb dark coniferous forests in resents the largest diversity proportion of the studied the modern middle taiga, primarily in western and area by this parameter. This leads us to assume that central part of the Russian Plain, which indicates a they could enjoy a much wider distribution in the much wider former distribution of south taiga tallherb restored forest cover of the studied area compared to forests in this area. Tilia cordata dot area of late the present. Apparently, to a great extent, the forest Holocene constructed using the spores and pollen outlook was determined by the tallherb forests of typi data also allows us to assume that the tallherb dark cum subassociation on the drained automorphous sur coniferous forests represented by Tilia cordata were faces, while the hydromorphous soils with flowing previously abundant in the territory of the modern hydration were dominated by the tallherb forests of the middle and possibly northern taiga (Fig. 1). The same filipenduletosum subassociation. The other trait of the data was obtained using the historical materials and tallherb forests that distinguishes them from other for herbarium findings of 18th and 19th centuries (Bakun, ests of northern and middle taiga is the stable presence 2006). of nemoral species (Adoxa moschatellina, Lathyrus To evaluate the chance to restore Tilia cordata vernus, Melica nutans, Milium effusum, Paris quadrifo within its late Holocene range, we carried out a com lia, Poa nemoralis, Stellaria holostea, Vicia sylvatica, parison of the mean point values of winter severity and Viola mirabilis), for which the modern broad (Cr), climate warmth (Tm), climate moisture (Om), leaved forests represent ideal ecological conditions and soil richness (Tr) of each geobotanical releve with (Lippmaa, 1938; Tsyganov, 1983). The permanent the range evaluations of Tilia cordata by the same indi presence of nemoral species in the tallherb dark conif ces (Fig. 2). The comparison of the data obtained with erous forests indicates that the local ecological condi regard to Cr and Tm factors showed that the majority tions of these forests are suitable for the stable exist of dots that characterize the mean values by these indi ence of these species. According to G.M. Zozulin ces were shifted toward the bottom borderline of the (Zozulin, 1973), the nemoral species of herbs repre Tilia cordata geographical range (Cr 5–7 and Tm 6–8 sent a group, the genesis of which is associated with points, respectively). These points correspond to the broadleaved forests. boreal and subboreal conditions where the mean tem Among broad leaved tree species, Tilia cordata perature of the coldest month ranges from –16 to penetrates farthest into the North. At the same time, ⎯ 24°C, which is typical of northern and, partly, the the literature data contain numerous descriptions of middle taiga. The bottom borderline of Tilia cordata its universal elimination during the cycles of tradi geographical range is inhabited by the following tall tional natural resource management of late middle herb forests that correspond to these indices: Lapland and late Holocene, up until the last centuries (Martin, Biosphere Reserve; Pinega Nature Reserve; north area 1984; Smirnova et al., 2001a, 2001b, 2002a, 2006a, of Pechoro Ilych Reserve; Paanajárvi National Park; 2006b, 2006c; Kharitonenkov, 2012). Consequently, Gorno Khadytin and Sobtyegan Reserves of Yamalo the assessment of Tilia cordata habitation in the mod Nenets Okrug, Tyumen’ oblast. A comparison of the
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12 (a) 13 (b) 11 12 10 11 9 10 8 9 7 8 6 5 7 Tsyganov scale, points
Tsyganov scale, points 4 6 Climate warmth according to Winter severity according to 3 5 0 200 400 600 800 1000 1200 0 200 400 600 800 1000 1200 Releve number Releve number
12 (c) 10 (d) 11 9 10 8 7 9 6 8 5 7 4 Tsyganov scale, points Tsyganov scale, points 6 Trophicity according to 3 Climate moisture according to 5 2 0 200 400 600 800 1000 1200 0 200 400 600 800 1000 1200 Releve number Releve number
Fig. 2. Correspondence of mean values of geobotanical releves of tallherb forests with ecological indices: (a) winter severity, (b) climate warmth, (c) climate moisture, (d) trophicity (calculated using the Tsyganov scale (1983) within the geographical range of Tilia cordata according to the same indices). data obtained with regard to the climate arid prairies. It is worth mentioning that the number of ity/humidity (Om) and soils poorness/richness (Tr) species of all the above mentioned units (floral com indices showed that the majority of dots that corre plexes) is considerably higher than that of the boreal spond to the mean values of these indices are situated and nitrophilous tallherb species that were revealed in in the middle part of Tilia cordata range (8–9 and 4– the forests, and steadily exist providing the presence of 7 points, respectively) (Fig. 2). Thus, the obtained windows in forest stand that occasionally form as a data confirm the general idea that winter severity and result of the death of one or several old trees (Zaugol climate warmth constitute a constraining factor for the nova et al., 2009). Tilia cordata distribution in the northern taiga, Analyzing the genus composition of the late whereas it could steadily inhabit the great part of the Pliocene tree layer on the Russian Plain, one can see middle taiga. that, by this time, the forest zone had already been For a long time, representatives of the tallherb represented by the ancestral forms of the modern dark group that determine the outlook of the tallherb dark coniferous, broad leaved, and small leaved species coniferous forests have attracted the attention of (Grichuk, 1989). Clearly, the spontaneous develop researchers of modern forest vegetation genesis of ment of these forests (as any natural forests) was Russia. From the florogenetic perspective, these spe accompanied by the constant creation of restoration cies were formerly attributed to the “Pleistocene floral windows and windfall–soil complexes; therefore, complex,” the birch or alder forest, and the “betula their lighting, humidity, and soil richness conditions cenoelement” (Krasheninnikov, 1937; Gorchakovskii, were suitable for the development of large, light 1969; Zozulin, 1973; Kleopov, 1990). Regarding the demanding species. They were able to survive under light demand as a common trait of the studied species, the trees canopy due to their tolerance to the low light these authors argued that they had formed in open for conditions and ability to pass into a state of secondary ests of Pleistocene, the ecotonic communities of dark dormancy (Vostochnoevropeiskie lesa …, 2004). One coniferous broad leaved forests, and in cryogenic can assume that the species of boreal and nitrophilous
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Table 2. Peculiarities of geographical range of boreal tallherb species Meridional range Number Species groups of species European Aconitum lasiostomum, Angelica archangelica, Crepis paludosa, Trollius europaeus, 6 Valeriana officinalis, Cicerbita alpina Ural Cicerbita uralensis; Knautia tatarica 2 Euro Siberian Actaea spicata, Cirsium heterophyllum 2 Eurasian Aconitium septentrionale, Actaeae erythrocarpa, Angelica sylvestris, Atragene sibirica, 31 Cacalia hastata, Calamagrostis arundinacea, Calamagrostis obtusata, Calamagrostis purpurea, Cardamine macrophylla, Cirsium oleraceum, Crepis sibirica, Cypripedium calceolus, Delphinium elatum, Diplazium sibiricum, Dryopteris expansa, Filipendula ulmaria, Galium boreale, Geranium sylvaticum, Lactuca sibirica, Lactuca tatarica, Lathurus gmelinii, Ligularia sibirica, Lilium martagon, Matteuccia struthiopteris, Paeonia anomala, Pleurospermim uralense, Saussurea alpina, Senecio nemorensis, Thalictrum minus, Veratrum lobelianum, Urtica dioica Holarctic Athyrium filix femina, Athyrium distentifolium, Cinna latifolia, Chamaenerion 4 angustifolium tallherb did not form in the open forests of Pleis leaved coniferous forests of Far East and Caucasus that tocene, but rather much earlier, in broad leaved conif were considered to be refugia of natural forests erous forests of late Pliocene. (Leskov, 1943; Sochava, 1946; Vasil’yev, 1958). The A comparison of the ranges of 45 boreal and nitro idea of the autochthonous origin of taiga forests was philous tallherb species identified so far using 850 geo also voiced by paleobotanists (Baranov, 1954; Doro botanical releves showed that the majority of them feev, 1963, 1964; Grichuk, 1989). According to them, belonged to Eurasian and Holarctic groups (35); a the Arcto Tertiary flora was changing rather slowly considerably minor part belonged to the European during the Pleistocene and the first half of Holocene: group (6), and just a small part fell into Ural (2), Euro throughout the entire cycle, the tree flora was of a Siberian (2), and Meridional groups (Table 2). As is mixed, nemoral–boreal type. clear from the maps of the high constant (5–4 points Taking into account the small distances that can be in releves) and constant (4–3 points in releves) spe covered by the seeds of late successional trees and cies, the use of the previously created arealogical base many herbs (Udra, 1988; Evstigneev, 2004), one must (by T.S. Prokazina) allowed us to considerably aug admit that the permanent existence of a constantly ment the former data on the boundaries of ranges and depleted Arcto Tertiary Geoflora was only possible to identify the areas of highest species concentration due to the significant number of refugia where it could (Figs. 3, 4). The constructed maps of the tallherb spe survive during the unfavorable stages of Pleistocene cies combined with the high resolution pictures form and early Holocene. The existence of refugia and their a necessary basis for further study and description of possible age have frequently been discussed in the lit tallherb dark coniferous forests in Russia. Surely fur erature (Lavrenko, 1930, 1938; Udra, 1988). Recently, ther geobotanical and floristic studies will allow us to the data on the location of refugia in Eastern Europe refine both the ranges of the tallherb species and tall during the Late Valdai glacial maximum has been herb dark coniferous forests; however, it is possible obtained using a large amount of palynological mate even now to consider the tallherb species to be an inte rial (Morozova and Kozharinov, 2001). gral component of dark coniferous forests that possess The data on the mixed (nemoral–boreal) nature of well manifested mosaics of restoration windows and the grass–shrub layer of the tallherb dark coniferous windfall–soil complexes. forests inhabited by various boreal species (from dwarf The compilation of the earlier data, as well as the shrubs such as Vaccinium myrtillus to tallherbs, namely, results of studying tallherb dark coniferous forests dis Paeonia, Aconitum, and Delphinium) and nemoral spe tribution, ecotopic diversity, and dependence on the cies (summer vegetative and ephemeroids), may also main ecological factors, allows us to consider the be regarded as an argument in favor of the autochtho problem of dark coniferous taiga origin and the devel nous origin of modern taiga. The following conditions opment from a somewhat different angle. are required for the development of nemoral–boreal Two main hypotheses of taiga origin, namely, the and grasses–shrubs layer: the fall of leafy trees, since autochthonous and allochthonous hypotheses, have the nemoral species often live under their canopy; the long been hotly debated in the literature (Yurtsev, presence of sags usually inhabited by the nitrophilous 1972). The hypothesis of the autochthonous origin of herbs and hemiboreal mosses; the presence of debris at modern taiga zone from the Arcto Tertiary forests that the second or third stage of filling by vegetation popu follow gradual climate change from late Pliocene to lated by small boreal herbs, dwarf shrubs, and green late Pleistocene was based on the studies of the broad boreal mosses; intercrown areas dominated by the tall
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(a)
1 2 3 0 1500 3000 6000 km 4 (b)
1 2 0 1500 3000 6000 km 3 (c)
1 2 0 1500 3000 6000 km 3 Fig. 3. Reconstructed dot area of high constant tallherb species in the studied dark coniferous forests: (a) Aconitum septentrionale Koelle: 1, dot location of a species according to Hulten and Fries (1986), 2, dot location of a species according to the herbarium labels data, 3, boundary of sporadic distribution of a species according to Hulten and Fries (1986), 4, continuous area of a species according to Hulten and Fries (1986). (b) Atragene sibirica L.: 1, dot location of a species according to Meusel et al. (1965, 1978), 2, dot location of a species according to the herbarium labels data, 3, continuous area of a species according to Meusel et al. (1965, 1978). (c) Cacalia hastata L.: 1, dot location of a species according to Hulten and Fries (1986), 2, dot location of a species accord ing to data of herbarium labels, 3, continuous area of a species according to Hulten and Fries (1986).
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(a)
1 2 3 0 1500 3000 6000 km
(b)
1 2 3 4 0 1500 3000 6000 km
Fig. 4. Reconstructed dot area of constant tallherb species in the studied dark coniferous forests: (a) Delphinium elatum L.: 1, dot location of species according to Hulten and Fries (1986), 2, dot location of a species according to the herbarium labels data, 3, continuous area of a species according to Hulten and Fries (1986); (b) Diplazium sibiricum (Turcs. Ex G Kunze) Kurata: 1, dot location of a species according to Hulten and Fries (1986), 2, dot location of species according to the herbarium labels data, 3, boundary of sporadic distribution of a species according to Hulten and Fries (1986), 4, continuous area of a species according to Hulten and Fries (1986). herbs; etc. It is worth mentioning that the presence of chagin and Baryshnikov, 1980), taiga was considered nemoral herbs species in north and middle taiga only the young formation, while the process of taiga land inhabited by the small leaved trees may implicitly sug scape development was seen as the movement of gest the former existence of the broad leaved tree spe already formed complexes from north to south. In cies there. accordance with the hypothesis of mountainous ori gin, taiga was considered the ancient formation that The hypothesis of the modern taiga allochthonous existed, at least, at the same time with the Arcto ter origin was also voiced by different researchers. The tiary flora or even had developed earlier and then starting point was believed to be located in the ancient replaced it in late Pliocene or in Pleistocene as a result land close to the Bering Sea (Beringiya, B.K. Shteg of a climatic deterioration. man, 1931), southern Palearctic mountains (Tolma chev, 1954), and the areas around the North Pole (sub The stages of origin and development of dark conif polar hypothesis), M.G. Popov, 1957). According to erous taiga, its natural outlook, floristic variety, and the hypotheses of subpolar and Bering origin (Veresh other characteristics are described in the monograph
BIOLOGY BULLETIN REVIEWS Vol. 3 No. 6 2013 502 SMIRNOVA et al. by A.I. Tolmachev (1954). His ideas about the natural Atlas Karel’skoi ASSR (Atlas of Karelia ASSR), Moscow: outlook of dark coniferous taiga were based on the Glav. Uprav. Geodez. Kartogr., 1989. geobotanical studies of the abundant mono or oligo Atlas Kirovskoi oblasti (Atlas of Kirov Oblast), Moscow: dominant floristically quite poor forests of the Russian Glav. Uprav. Geodez. Kartogr., 1968. Plain, Urals, and Siberia dominated by the evergreen Atlas Komi ASSR (Atlas of Komi ASSR), Moscow: Glav. vascular plants and boreal mosses (Rastitel’nyi …, Uprav. Geodez. Kartogr., 1964. 1956; Rastitel’nost’ …, 1980). These studies led him to Atlas Leningradskoi oblasti (Atlas of Leningrad Oblast), the conclusion that “the typical dark coniferous taiga Moscow: Glav. Uprav. Geodez. Kartogr., 1967. is characterized by not only a limited number of for Atlas Permskoi oblasti (Atlas of Perm Oblast), Moscow: est forming species, but also by a generally poor flora DIK, 1999. composition. The limited composition of species also Atlas Respubliki Komi (Atlas of the Komi Republic), Mos agrees with the scarcity of vegetation cover…. The cow: DIK, 2001. plants that inhabit the canopy of the dark coniferous Atlas Tyumenskoi oblasti (Atlas of Tyumen Oblast), Mos forest represent only a negligible portion….” (Tolma cow: Glav. Uprav. Geodez. Kartogr., 1971. chev, 1954, p. 11). However, the studies of refugia of Atlas Yaroslavskoi oblasti (Atlas of Yaroslavl Oblast), Mos taiga forests carried out during the last decades and cow: Glav. Uprav. Geodez. Kartogr., 1964. based on the modern concepts of synecology and his Atlas Yaroslavskoi oblasti (Atlas of Yaroslavl Oblast), Mos torical ecology (Vera, 2000; Bobrovskii, 2010) lead us cow: DIK, 1999. to the conclusion that the natural outlook of dark Bakun, E.Yu., Reconstruction of tree species areas in the coniferous taiga was mostly determined by the tallherb Russian plain, Lesovedenie, 2006, no. 2, pp. 64–70. forests. Baranov, V.I., Etapy razvitiya flory i rastitel’nosti SSSR v treti chnom periode (Stages of Development of Flora and Thus, the results of previous studies of successional Vegetation of USSR in Tertiary Period), Kazan: Kazan. processes in taiga forests, along with the data on the Gos. Univ., 1954. ranges of tallherb species and the tallherb forests dis Bobrovskii, M.V., Lesnye pochvy Evropeiskoi Rossii: bioti tribution, support the autochthonous hypothesis of cheskie i antropogennye factory formirovaniya (Forest dark coniferous taiga origin. Soils of European Russia: Biotic and Anthropogenic Factors), Moscow: KMK, 2010. Bobrovskii, M.V. and Khanina, L.G., Kaluzhskie Zaseki CONCLUSIONS Nature Reserve, in Otsenka i sokhranenie bioraznoob The distribution of tallherb forests on the large area raziya lesnogo pokrova v zapovednikakh Evropeiskoi from the northern boundaries of forest belt to the Rossii (Assessment and Preservation of Biodiversity of southern taiga and subtaiga, their considerable Forests in Reserves of European Russia), Moscow: ecotypic variety, and the ability to survive under severe Nauchnyi Mir, 2000, pp. 104–122. climate conditions while preserving the nemoral and Dorofeev, P.I., New data on Pleistocene floras of Belorussia and Smolensk oblast, in Materialy po istorii flory i rasti boreal flora species allows us to assume that these par tel’nosti SSSR (The Data on History of Flora and Vege ticular forests represented in large part the prehistori tation of USSR), Moscow: Akad. Nauk SSSR, 1963, cal outlook of taiga forests in the studied area. no. 4, pp. 5–180. This suggestion may be regarded as an argument in Dorofeev, P.I., Razvitie tretichnoi flory SSSR po dannym favor of the autochthonous origin of forests. On the paleokarpichekikh issledovanii (Development of Ter other hand, the earlier data on the highest degree of tiary Flora of USSR Assessed on the Data of Paleocarp biological variety, productivity, and intensity of biolog Studies), Leningrad: Nauka, 1963. ical turnover among the so far analyzed dark conifer Ekologicheskii atlas Murmanskoi oblasti (Ecological Atlas of ous taiga forests of the studied area allows us to con Murmansk Oblast), Moscow: Inst. Probl. Prom. Ekol. sider tallherb forests to be the most important objects Severa, 1999. in estimating the ability of the forest cover to fulfill its Evstegneev, O.I., Population strategies of wood species, in ecosystem functions. Vostochnoevropeiskie lesa: istoriya v golotsene i sovre mennost’ (East European Forests: the Holocene His tory and the Current State), Moscow: Nauka, 2004, ACKNOWLEDGMENTS book 1, pp. 176–205. Gorchakovskii, P.L., Osnovnye problemy istoricheskoi fito This work was supported by the Russian Founda geografii Urala (General Problems of Historical Phyto tion for Basic Research, project no. 10 04 00355a, geography of Ural), Sverdlovsk, 1969. and the Fundamental Research Program, Russian Grichuk, V.P., Istoriya flory i rastitel’nosti Russkoi ravniny v Academy of Sciences Presidium “Wildlife: Current pleistotsene (History of Flora and Vegetation of Russian Status and Problems of Development.” Plain in Pleistocene), Moscow: Nauka, 1989. Hulten, E. and Fries, M., Atlas of north European vascular plants north of the Tropic of Cancer Konigstein: Koeltz REFERENCES Scientific Books, 1986, 3 vols. 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