model reconstruction of restored taiga forest cover · the taiga forests (boreal and hemiboreal) of...

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.

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INTRODUCTION

A vital task of modern natural resource manage�ment is its reorientation towards preservation and res�toration of ecological functions of ecosystems,namely, climate regulation, water and soil protection,productional function, and maintaining the biovariety.It was proved earlier that the fullest realization of eco�logical functions can be accomplished in a restoredforest cover, i.e., in this cover, which can form as aresult of a long spontaneous development (withoutanthropogenic disturbances and natural disasters) andis represented by all species preserved so far(Smirnova, 2004; Smirnova et al., 2006a, 2006b). Nat�ural releves and/or model reconstructions of suchcover will allow us to assess the ability of the forestcover to fulfill its ecological functions under differentclimate conditions. According to the today’s beliefs,the taiga forests (boreal and hemiboreal) of the North�ern Hemisphere have preserved their natural composi�tion and ability to fulfill the main ecosystem functionsto the fullest extent possible (Monitoring…, 2008).Therefore, the thorough study of these forests is a mat�ter of theoretical and practical importance. Moreover,a model reconstruction of the restored taiga forestcover represents another step towards the solution ofproblem of origin and development of dark coniferoustaiga.

As was rightly mentioned by V.B. Sochava (1946),the studies of refugia, i.e., areas in which vegetationhas barely been touched by natural disasters and pow�erful anthropogenic impacts, play a considerable rolein dealing with the problem of vegetation genesis,

along with the paleobotanical and florogenetic meth�ods. The analysis of structural and taxonomic varietyof refugium vegetation allows us to define the naturalcharacteristics of vegetation cover and form an opin�ion as to the genesis of this vegetation type, while com�paring the obtained results with the paleobotanicaland florogenetic data.

So far, plenty of data has been assembled on thecomposition and structure of taiga forests communi�ties in refugia on the Russian Plain and the Urals, aswell as in some regions of Western and Central Siberia.According to the studies performed, they are passingthrough the late successional or quasiclimax stage and arecharacterized by the following features (Turubanova,1999, Yaroshenko, 1999; Bobrovskii and Khanina, 2000;Smirnova et al., 2001a, 2011; Yaroshenko et al., 2001;Vostochnoevropeiskie lesa…, 2004; Smirnova, 2004;Zaugolnova et al., 2009; Bobrovskii, 2010):

(1) the presence of dark conifer trees (Picea, abies,P. obovata, Abies sibirica, and Pinus sibirica), broad�leaved trees (Tila cordata, Ulmus laevis, and U. scabraet al.), and small�leaved trees (species of the genusBetula and Alnus, Populus tremula, and Padus aviumet al.) in the forest stand of the studied area;

(2) unevenly aged cenopopulations of late�succes�sional tree species;

(3) the development of the entire set of micromo�saic structural elements: intercrown and undertreespaces of different tree species, elements of windfall�soil complexes (treefall soil pits, knobs, debris, andstumps) at different stages of development;

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, Russiae�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 darkconiferous forests has been constructed based on 850 geobotanical releves (from the Database of the Centerfor 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 thestudied 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 databaseof herbaria labels, refined ranges of tallherb species were constructed. The importance of studying tallherbdark coniferous forests has been assessed to determine the origin of the dark coniferous taiga and select virginforests 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

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(4) presence in ecosystem of both vascular plantsspecies and mosses of all eco�coenotic groups capableto inhabit the studied forests;

(5) the dominance of boreal and nitrophilous tall�herb in the soil cover in terms of spreading and phyto�mass;

(6) the absence of traces of fire and cutting in vege�tation cover, and coal in soil;

(7) undifferentiated soil profile represented byhumus horizon developed as a result of recurrent soiltrenching due to the late�successional tree fall.

The above�mentioned features are most pro�nounced in quasiclimax forests, whereas the late�suc�cessional forests have not yet developed these charac�teristics (The Population Structure …, 1985; TheMosaic�Cycle Concept …, 1991). This is most stronglytranslated by the floristic incompleteness of commu�nities caused by the considerable gaps in tree rangesdue to the anthropogenic activities. A good example isthe considerable gaps in Tilia cordata Mill. range inthe southern taiga subzone documented in the historicliterature (Vostochnoevropeiskie lesa …, 2004; Bakun,2006; Bobrovskii, 2010). From a physionomical pointof view, the late�successional and quasiclimax darkconiferous forests are characterized by the dominanceof tall boreal herbs of two ecocoenotic groups, i.e.,boreal tallherbs and nitrophilous tallherbs. The herbsof these groups grow quickly and are capable ofachieving a height of 1.5–3.0 m during growingperiod; the majority has either large leaves or a bignumber of shoots with medium sized leaves in largetufts (grasses). The dense canopy of these herbs, aswell as the considerable herbaceous litter prevents thegrowth of boreal green mosses (Smirnova et al., 2011).The phytomass of the annually falling abovegroundshoots of the vascular plants in tallherb forests is five toten times higher than the phytomass of vascular plantsin the most widespread taiga forests, namely, those ofa green moss and sphagnous type. The fall of the tall�herb species is highly enriched with mineral elementscompared to that of the herbs and dwarf shrubs ofgreen moss and sphagnous forests (Lukina andNikonov, 1998; Lukina et al., 2006). It is quicklyabsorbed by the soil biota, which explains the highvelocity of mineral elements turnover and advancedforest productivity. Tallherb species are light�demand�ing, but at the same time tolerant to a lack of light, i.e.,they withstand the provisional reduction in lightintensity and quickly return to their former status dueto the light window.

The tallherb species are different in terms of theirdependence on ecological factors (Tsyganov, 1983);this explains the dominance of different members ofthis group under different regimes of moisture, acidity,and soil richness.

The ranges of majority of species of this groupreach the north forest frontier and penetrate even a lit�tle further to the north, which leads us to consider

them to be representatives of taiga forests above all(Zaugolnova et al., 2009).

According to the International Code of Phytoso�ciological Nomenclature (Weber et al., 2005), thestudied tallherb forests have the following syntaxo�nomic position:

Class Vaccinio�Piceetea Br.�Bl. in Br.�Bl., Sissinghet Vlieger 1939;

Order Picetalia excelsae Paw owski in Paw owski,Soko owski et Wallisch 1928 (=Vaccinio�PiceetaliaBr.�Bl. in Br.�Bl., Siss. et Vlieger 1939);

Alliance Vaccinio�Piceion Br.�Bl., Sissingh etVlieger 1939;

Suballiance Atrageno�Piceenion obovatae Zaugol�nova et al., 2009;

Assoc. Aconito septentrionalis–Piceetum obovataeZaugolnova et al., 2009;

Subassoc. A. s.–P. o. typicum Zaugolnova et al.,2009;

Subassoc. A. s.–P. o. filipenduletosum Zaugolnovaet al., 2009.

Communities of tallherb dark coniferous forestsare revealed on the watersheds in well�drained habitatsand in valleys with small rivers and brooks where thereis a flowing hydration. Diagnostic species of the entireassociation are as follows: Aconitum septentrionaleKoelle, Veratrum lobelianum Bernh., Chamaenerionangustifolium (L.) Scop., Calamagrostis purpurea(Trin.) Trin., and Thalictrum minus L. The watershedsforests belong to the subassociation A. s.–P. o. typi�cum, the diagnostic species of the entire associationbeing joined by Diplazium sibiricum (Turcz. exG. Kunze) Kurata; valley forests belong to the subas�sociation A. s.–P. o. filipenduletosum, within which agroup of diagnostic species was defined that includedFilipendula ulmaria (L.) Maxim., Geum rivale L., Trol�lius europaeus L., etc.

Tallherb forests differ from those of green moss andsphagnous types by the more complicated structure oflayers, unevenly aged tree cenopopulations, and thehighest species and ecocoenotic variety. The trees inthese communities are large and well developed, andtheir deaths are followed by pedoturbations. Conse�quently, it is possible to preserve the mosaic of restora�tion windows that determines the mosaicity of groundcover light regime, as well as the mosaic of windfall–soil complexes that accounts for the mosaicity ofmicrohabitats with different regimes of temperature,moisture, soil acidity, etc.

The herbaceous layer of these forests is dense (70–90% spreading), polidominant, and includes at leastthree sublayers. The first sublayer is formed by theboreal and/or nitrophilous tallherbs; the second one isformed of boreal, nemoral, nitrophilous, andmedium�sized meadow and edge herbs; the third oneis formed of small nitrophilous and boreal herbs anddwarf shrubs. The moss cover is not dense (20–10%spreading), but consists of a large number of ecologi�

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MODEL RECONSTRUCTION OF RESTORED TAIGA FOREST COVER 495

cally different species. The alluvial soils with moder�mull horizon 35–50 cm in depth are formed in tall�herb floodplain forests. The brown soils with moder�mull�humus horizon reaching a depth of 60 cm areformed in tallherb watersheds forests (Smirnova et al.,2006c; Bobrovskii, 2010).

To date, the accumulated data on ecology and dis�tribution of the tallherb species, successional status ofthe tallherb dark coniferous forests, their ecotopes,and geographical range allowed us to undertake recon�struction of the restored forest cover of studied area.

For this purpose, the following tasks have been car�ried out:

1. The dot area of the tallherb dark coniferous for�ests was constructed and a comparison of their distri�bution with the modern zonal division of the studiedarea was performed.

2. Ecotopes of tallherb forests were identified.3. A comparison of the tallherb forests’ range with

the modern outlined range of Tilia cordata and its dotarea in late Holocene has been performed.

4. The mean point values of several climatic andecotypic indices that characterize the communities oftallherb dark coniferous forests were estimated usingthe ecological scales (Tsyganov, 1983); they were thencompared with the range evaluations of the same indi�ces for Tilia cordata being the supposed inhabitant ofthese forests.

5. The peculiarities of geographical range of borealand nitrophilous tallherb species considered the indi�cators of the late�successional and quasiclimax devel�opment stages of dark coniferous forests were revealed.

6. Based on the data on successional status and dis�tribution peculiarities of the tallherb dark coniferousforests, the theories of dark coniferous taiga origin anddevelopment have been analyzed.

MATERIALS AND METHODS

To carry out the set tasks, 850 releves from the Geo�botanical Database of the Center for Problems andEcology and Productivity of Forests, Russian Academyof Sciences, performed on the square grounds 100 m2 insize, were analyzed according to the method publishedearlier (Smirnova et al., 2002b) in the Murmansk,Leningrad, Arkhangelsk, Kostroma, Yaroslavl, Kirov,Vologda, Sverdlovsk, and Tyumen’ oblasts; Perm andKrasnoyarsk krais; the Republics of Karelia and Komi;and the Khanty�Mansi and Yamalo�Nenets autono�mous okrugs. The analyzed releves were made inmountainous and elevated areas, as well as rises offolded or glacial origin, including the Baltic Shield;Andomsk Upland; Belozersko�Kirillovka Upland; theNorthern Ridge; the Timan Ridge; the Siberian Ridge;Kondo�Sos’vinsk Priob’e; Western and Eastern Uralsmacroslopes; and on the plain areas, including theBelomorsko�Kuloiskoe Plateau, Mezensk Plain, andVolkhovo�Ilmen Lowlands. The dot area of the tall�

herb dark coniferous forests was constructed based onthe geographic coordinates of releves determined by aGPS receiver. To compare the obtained dot area of thetallherb dark coniferous forests with the modern zonaldivision of studied area, we used a map (Karta “Zony itipy ...,” 1999) with some modifications. The mapswere created using ArcMap 9.2 (ESRI Inc. softwareproduct). To compare the dot area of the tallherb darkconiferous forests with the modern outlined range ofTilia cordata Mill. and its dot area in the lateHolocene, we used data kindly furnished byS.A. Turubanova. Geobotanical releves and thematicmaps of quaternary deposits, landscape types, andgeological structures of different scale were used toanalyze the correspondence of the tallherb dark conif�erous forests to the relief elements and superficialdeposits (Atlas…, 1964a, 1964b, 1967, 1968, 1971,1976, 1989, 1999a, 1999b, 2001; Ekologicheskii…,1999). The maps from the published atlases wereanchored to the ArcMap 9.2 environment where thespace analysis of information was conducted.

The direct ordination of geobotanical releves byenvironmental indices was performed using ecologicalscales by Tsyganov (Tsyganov, 1983). The calculationsof the mean point values by indices of winter severity(Cr), climate warmth (Tm), climate moisture (Om),and soil richness (Tr), as well as the analysis of geobo�tanical releves were performed using the PcOrd pack�age of SpeDiv 1.3.0 software (developed byV.E. Smirnov).

To identify the peculiarities of geographical rangeof boreal and nitrophilous tallherb species that indi�cate late�successional and quasi�climax dark conifer�ous forests, the range maps of these species taken fromthe literature data (Meusel et al., 1978; Hulten andFries, 1986) were scanned and digitalized. To apply thedots of collecting herbarium specimens, the study oflarge herbaria of the universities (Moscow State Uni�versity, Kazan State University, and Tomsk State Uni�versity) and academic institutions (Komarov Botani�cal Institute, Russian Academy of Sciences; Instituteof Plant and Animal Ecology, Ural Branch, RussianAcademy of Sciences; Komi Science Center, UralBranch, Russian Academy of Sciences; Central Sibe�rian Botanical Garden, Russian Academy of Sciences,et al.) was carried out. The geographic coordinates ofdiscovery points were established using the labels dataand then added to the GIS. The database preserves theinformation on points of discovery of tallherb species(7180 records), along with their coordinates. The cor�rection of meridional groups of these species’ rangeswas performed using this data.

RESULTS AND DISCUSSION

To date, tallherb dark coniferous forests have beenrevealed in the largest part of the Russian Plain and theUrals, as well as in some areas of Western and CentralSiberia (Fig. 1). The irregularity in the distribution of

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tallherb forests within the studied area is primarilycaused by the fact that they survived in places leastaffected by anthropogenic disturbances. They werefound to the west of the analyzed area in poorly acces�sible regions of North Karelia and the Kola Peninsula:Lapland Biosphere Reserve and Paanajárvi NationalPark and its environs (Smirnova and Korotkov, 2001).They were also revealed at the heart of the RussianPlain, in the Pinega Nature Reserve, Kologriv Forest,Kilemary Reserve of Nihzni Novgorod oblast (Suktse�sionnye…, 1999; Shirokov et al., 1999, 2001; Otsenka isokhranenie…, 2000; Shirokov, 2002, Lugovaya, 2010).The tallherb forests were identified in the low�hill ter�rains of nearly all taiga reserves of the Ural westernmacroslopes: Visim, Vishersk, Denezhkin Kamen’,Pechoro�Ilych, Malaya Sos’va (Yaroshenko et al.,1998; Suktsesionnye…, 1999; Prokazina, 2007, 2011;Zaprudina and Smirnov, 2010). Surely, the systematicsearches of tallherb dark coniferous forests in the areaswhere they have not yet been found may considerablyexpand this list; however, this presents significant dif�ficulties. It is worth mentioning that the majority ofthe studied tallherb communities have rather smallareas, i.e., from one or less to several hectares (Zaugol�

nova et al., 2008). The only exception is the taigareserves of the Ural slopes where the tallherb forestsarea reaches dozens of hectares in the middle andupper slopes (Smirnova et al., 2011). The small num�ber of locations in Western Siberia is caused by the fee�ble preservation of dark coniferous forests and poorknowledge on the subject. One of the most interestingobjects is the tallherb spruce�fir forests in the Gorno�Khadytinskii Reserve of the Southern Yamal (southerntundra region), which are similar in all characteristicsto the northern taiga tallherb forests (Smirnova et al.,2008).

The overview of the tallherb forests locations per�formed using the field survey data and the publishedcartographic materials (Atlas…, 1964a, 1964b, 1967,1968, 1971, 1976, 1989, 1999a, 1999b, 2001; Ekolog�icheskii …, 1999) allowed us to draw up a list of themain relief elements, as well as types of quaternarydeposits and parent rocks where the tallherb forestswere found. The interfluve and valley complexes wereanalyzed separately (Table 1). In both complexes, thetallherb forests can only be found in the drained areas;communities of typicum subassociation can be foundin moderately humid areas; and communities of fili�

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Fig. 1. Dot area of the tallherb dark coniferous forests of the Russian Plain, the Urals, and some areas of Western and CentralSiberia compared to the modern outlined range of Tilia cordata and its dot area in late Holocene: 1, location dots of the tallherbforests, 2, location dots of Tilia cordata pollen in Holocene, 3, Tilia cordata outlined range. Natural subzones and zones: 4, foresttundra, 5, northern taiga; 6, middle taiga; 7, southern taiga and subtaiga, 8, mountain taiga forests.



penduletosum subassociation can be found in humidand over humid areas, provided the existence of flow�ing hydration.

The diversity of quaternary deposits and parentrocks inhabited by the tallherb forests is huge and rep�resents the largest diversity proportion of the studiedarea by this parameter. This leads us to assume thatthey could enjoy a much wider distribution in therestored forest cover of the studied area compared tothe present. Apparently, to a great extent, the forestoutlook was determined by the tallherb forests of typi�cum subassociation on the drained automorphous sur�faces, while the hydromorphous soils with flowinghydration were dominated by the tallherb forests of thefilipenduletosum subassociation. The other trait of thetallherb forests that distinguishes them from other for�ests of northern and middle taiga is the stable presenceof nemoral species (Adoxa moschatellina, Lathyrusvernus, Melica nutans, Milium effusum, Paris quadrifo�lia, Poa nemoralis, Stellaria holostea, Vicia sylvatica,and Viola mirabilis), for which the modern broad�leaved forests represent ideal ecological conditions(Lippmaa, 1938; Tsyganov, 1983). The permanentpresence of nemoral species in the tallherb dark conif�erous forests indicates that the local ecological condi�tions of these forests are suitable for the stable exist�ence of these species. According to G.M. Zozulin(Zozulin, 1973), the nemoral species of herbs repre�sent a group, the genesis of which is associated withbroadleaved forests.

Among broad�leaved tree species, Tilia cordatapenetrates farthest into the North. At the same time,the literature data contain numerous descriptions ofits universal elimination during the cycles of tradi�tional natural resource management of late middleand late Holocene, up until the last centuries (Martin,1984; Smirnova et al., 2001a, 2001b, 2002a, 2006a,2006b, 2006c; Kharitonenkov, 2012). Consequently,the assessment of Tilia cordata habitation in the mod�

ern tallherb taiga forests becomes a matter of consid�erable interest with regard to the model reconstructionof the restored forest cover.

The modern continuous area of Tilia cordata over�laps the dot area of tallherb dark coniferous forests inthe modern middle taiga, primarily in western andcentral part of the Russian Plain, which indicates amuch wider former distribution of south taiga tallherbforests in this area. Tilia cordata dot area of lateHolocene constructed using the spores and pollendata also allows us to assume that the tallherb darkconiferous forests represented by Tilia cordata werepreviously abundant in the territory of the modernmiddle and possibly northern taiga (Fig. 1). The samedata was obtained using the historical materials andherbarium findings of 18th and 19th centuries (Bakun,2006).

To evaluate the chance to restore Tilia cordatawithin its late Holocene range, we carried out a com�parison of the mean point values of winter severity(Cr), climate warmth (Tm), climate moisture (Om),and soil richness (Tr) of each geobotanical releve withthe range evaluations of Tilia cordata by the same indi�ces (Fig. 2). The comparison of the data obtained withregard to Cr and Tm factors showed that the majorityof dots that characterize the mean values by these indi�ces were shifted toward the bottom borderline of theTilia cordata geographical range (Cr 5–7 and Tm 6–8points, respectively). These points correspond to theboreal and subboreal conditions where the mean tem�perature of the coldest month ranges from –16 to⎯24°C, which is typical of northern and, partly, themiddle taiga. The bottom borderline of Tilia cordatageographical range is inhabited by the following tall�herb forests that correspond to these indices: LaplandBiosphere Reserve; Pinega Nature Reserve; north areaof Pechoro�Ilych Reserve; Paanajárvi National Park;Gorno�Khadytin and Sobtyegan Reserves of Yamalo�Nenets Okrug, Tyumen’ oblast. A comparison of the

Table 1. Characteristics of habitats of tallherb dark coniferous forest communities

Characteristics of habitats Interfluve complex Valley complex

Elements of relief (based on the field survey data)

Leveled interfluve area, interfluvial slope (incl. swamped areas); closed positive and negative relief forms in the interfluve area (incl. karstic ones); lake terrace

Valley, valley side, first terrace above floodplain, high floodplain (ridges, depressions, subhorizontal surfaces, and natural levees); bottom of the brook valley

Quaternary deposits and parent rocks based on the literature data

Quartzites, quartzite sand, clay–limestone deposits; gabbroids and granitoids intrusions; amphibolites and gneisses; morainic and fluvioglacial deposits (clays, clay loams, boulder clays interlaid with inequigranular sands, sands); basic and ultrabasic rock complexes; red marl beds and sandstones

Alluvial (sands, sand clays, clay loams, peat interbeds, and lenses; sandy–clay and peat�like); Valdai and Moscow glacial deposits: fluvioglacial (pebblebeds and sands; lake and glacial (sands and clays); glacial (boulder clay loams)

Parent rocks (based on the field survey data)

Light, middle, and heavy clay loams (incl. morainic ones); sand; rocky sand (with pebble); limestones; quartzites, granites, and diabases

Alluvial clay loams, sand clays; crypto podzolic sand clays on clay loams; karsting rocks

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data obtained with regard to the climate arid�ity/humidity (Om) and soils poorness/richness (Tr)indices showed that the majority of dots that corre�spond to the mean values of these indices are situatedin the middle part of Tilia cordata range (8–9 and 4–7 points, respectively) (Fig. 2). Thus, the obtaineddata confirm the general idea that winter severity andclimate warmth constitute a constraining factor for theTilia cordata distribution in the northern taiga,whereas it could steadily inhabit the great part of themiddle taiga.

For a long time, representatives of the tallherbgroup that determine the outlook of the tallherb darkconiferous forests have attracted the attention ofresearchers of modern forest vegetation genesis ofRussia. From the florogenetic perspective, these spe�cies were formerly attributed to the “Pleistocene floralcomplex,” the birch or alder forest, and the “betulacenoelement” (Krasheninnikov, 1937; Gorchakovskii,1969; Zozulin, 1973; Kleopov, 1990). Regarding thelight demand as a common trait of the studied species,these authors argued that they had formed in open for�ests of Pleistocene, the ecotonic communities of darkconiferous broad�leaved forests, and in cryogenic

prairies. It is worth mentioning that the number ofspecies of all the above�mentioned units (floral com�plexes) is considerably higher than that of the borealand nitrophilous tallherb species that were revealed inthe forests, and steadily exist providing the presence ofwindows in forest stand that occasionally form as aresult of the death of one or several old trees (Zaugol�nova et al., 2009).

Analyzing the genus composition of the latePliocene tree layer on the Russian Plain, one can seethat, by this time, the forest zone had already beenrepresented by the ancestral forms of the modern darkconiferous, broad�leaved, and small�leaved species(Grichuk, 1989). Clearly, the spontaneous develop�ment of these forests (as any natural forests) wasaccompanied by the constant creation of restorationwindows and windfall–soil complexes; therefore,their lighting, humidity, and soil richness conditionswere suitable for the development of large, light�demanding species. They were able to survive underthe trees canopy due to their tolerance to the low lightconditions and ability to pass into a state of secondarydormancy (Vostochnoevropeiskie lesa …, 2004). Onecan assume that the species of boreal and nitrophilous

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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 rangeof Tilia cordata according to the same indices).



tallherb did not form in the open forests of Pleis�tocene, but rather much earlier, in broad�leaved conif�erous forests of late Pliocene.

A comparison of the ranges of 45 boreal and nitro�philous tallherb species identified so far using 850 geo�botanical releves showed that the majority of thembelonged to Eurasian and Holarctic groups (35); aconsiderably minor part belonged to the Europeangroup (6), and just a small part fell into Ural (2), Euro�Siberian (2), and Meridional groups (Table 2). As isclear from the maps of the high�constant (5–4 pointsin releves) and constant (4–3 points in releves) spe�cies, the use of the previously created arealogical base(by T.S. Prokazina) allowed us to considerably aug�ment the former data on the boundaries of ranges andto identify the areas of highest species concentration(Figs. 3, 4). The constructed maps of the tallherb spe�cies combined with the high�resolution pictures forma necessary basis for further study and description oftallherb dark coniferous forests in Russia. Surely fur�ther geobotanical and floristic studies will allow us torefine both the ranges of the tallherb species and tall�herb dark coniferous forests; however, it is possibleeven now to consider the tallherb species to be an inte�gral component of dark coniferous forests that possesswell�manifested mosaics of restoration windows andwindfall–soil complexes.

The compilation of the earlier data, as well as theresults of studying tallherb dark coniferous forests dis�tribution, ecotopic diversity, and dependence on themain ecological factors, allows us to consider theproblem of dark coniferous taiga origin and the devel�opment from a somewhat different angle.

Two main hypotheses of taiga origin, namely, theautochthonous and allochthonous hypotheses, havelong been hotly debated in the literature (Yurtsev,1972). The hypothesis of the autochthonous origin ofmodern taiga zone from the Arcto�Tertiary forests thatfollow gradual climate change from late Pliocene tolate Pleistocene was based on the studies of the broad�

leaved coniferous forests of Far East and Caucasus thatwere considered to be refugia of natural forests(Leskov, 1943; Sochava, 1946; Vasil’yev, 1958). Theidea of the autochthonous origin of taiga forests wasalso voiced by paleobotanists (Baranov, 1954; Doro�feev, 1963, 1964; Grichuk, 1989). According to them,the Arcto�Tertiary flora was changing rather slowlyduring the Pleistocene and the first half of Holocene:throughout the entire cycle, the tree flora was of amixed, nemoral–boreal type.

Taking into account the small distances that can becovered by the seeds of late�successional trees andmany herbs (Udra, 1988; Evstigneev, 2004), one mustadmit that the permanent existence of a constantlydepleted Arcto�Tertiary Geoflora was only possibledue to the significant number of refugia where it couldsurvive during the unfavorable stages of Pleistoceneand early Holocene. The existence of refugia and theirpossible age have frequently been discussed in the lit�erature (Lavrenko, 1930, 1938; Udra, 1988). Recently,the data on the location of refugia in Eastern Europeduring the Late Valdai glacial maximum has beenobtained using a large amount of palynological mate�rial (Morozova and Kozharinov, 2001).

The data on the mixed (nemoral–boreal) nature ofthe grass–shrub layer of the tallherb dark coniferousforests inhabited by various boreal species (from dwarfshrubs such as Vaccinium myrtillus to tallherbs, namely,Paeonia, Aconitum, and Delphinium) and nemoral spe�cies (summer vegetative and ephemeroids), may alsobe regarded as an argument in favor of the autochtho�nous origin of modern taiga. The following conditionsare required for the development of nemoral–borealand grasses–shrubs layer: the fall of leafy trees, sincethe nemoral species often live under their canopy; thepresence of sags usually inhabited by the nitrophilousherbs and hemiboreal mosses; the presence of debris atthe second or third stage of filling by vegetation popu�lated by small boreal herbs, dwarf shrubs, and greenboreal mosses; intercrown areas dominated by the tall�

Table 2. Peculiarities of geographical range of boreal tallherb species

Meridional range groups Species Number

of species

European Aconitum lasiostomum, Angelica archangelica, Crepis paludosa, Trollius europaeus, Valeriana officinalis, Cicerbita alpina

6

Ural Cicerbita uralensis; Knautia tatarica 2Euro�Siberian Actaea spicata, Cirsium heterophyllum 2Eurasian Aconitium septentrionale, Actaeae erythrocarpa, Angelica sylvestris, Atragene sibirica,

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

31

Holarctic Athyrium filix�femina, Athyrium distentifolium, Cinna latifolia, Chamaenerion angustifolium

4

500


SMIRNOVA et al.

(a)

(b)

(c)

0 1500 3000 6000 km

1234

0 1500 3000 6000 km

123

0 1500 3000 6000 km

123

Fig. 3. Reconstructed dot area of high constant tallherb species in the studied dark coniferous forests: (a) Aconitum septentrionaleKoelle: 1, dot location of a species according to Hulten and Fries (1986), 2, dot location of a species according to the herbariumlabels data, 3, boundary of sporadic distribution of a species according to Hulten and Fries (1986), 4, continuous area of a speciesaccording 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).



herbs; etc. It is worth mentioning that the presence ofnemoral herbs species in north and middle taiga onlyinhabited by the small�leaved trees may implicitly sug�gest the former existence of the broad�leaved tree spe�cies there.

The hypothesis of the modern taiga allochthonousorigin was also voiced by different researchers. Thestarting point was believed to be located in the ancientland close to the Bering Sea (Beringiya, B.K. Shteg�man, 1931), southern Palearctic mountains (Tolma�chev, 1954), and the areas around the North Pole (sub�polar hypothesis), M.G. Popov, 1957). According tothe hypotheses of subpolar and Bering origin (Veresh�

chagin and Baryshnikov, 1980), taiga was consideredthe young formation, while the process of taiga land�scape development was seen as the movement ofalready formed complexes from north to south. Inaccordance with the hypothesis of mountainous ori�gin, taiga was considered the ancient formation thatexisted, at least, at the same time with the Arcto�ter�tiary flora or even had developed earlier and thenreplaced it in late Pliocene or in Pleistocene as a resultof a climatic deterioration.

The stages of origin and development of dark conif�erous taiga, its natural outlook, floristic variety, andother characteristics are described in the monograph

(b)

(a)

0 1500 3000 6000 km

0 1500 3000 6000 km

123

1234

Fig. 4. Reconstructed dot area of constant tallherb species in the studied dark coniferous forests: (a) Delphinium elatum L.: 1, dotlocation 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, dotlocation 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 accordingto Hulten and Fries (1986).

502


SMIRNOVA et al.

by A.I. Tolmachev (1954). His ideas about the naturaloutlook of dark coniferous taiga were based on thegeobotanical studies of the abundant mono� or oligo�dominant floristically quite poor forests of the RussianPlain, Urals, and Siberia dominated by the evergreenvascular plants and boreal mosses (Rastitel’nyi …,1956; Rastitel’nost’ …, 1980). These studies led him tothe conclusion that “the typical dark coniferous taigais characterized by not only a limited number of for�est�forming species, but also by a generally poor floracomposition. The limited composition of species alsoagrees with the scarcity of vegetation cover…. Theplants that inhabit the canopy of the dark coniferousforest represent only a negligible portion….” (Tolma�chev, 1954, p. 11). However, the studies of refugia oftaiga forests carried out during the last decades andbased on the modern concepts of synecology and his�torical ecology (Vera, 2000; Bobrovskii, 2010) lead usto the conclusion that the natural outlook of darkconiferous taiga was mostly determined by the tallherbforests.

Thus, the results of previous studies of successionalprocesses in taiga forests, along with the data on theranges of tallherb species and the tallherb forests dis�tribution, support the autochthonous hypothesis ofdark coniferous taiga origin.

CONCLUSIONS

The distribution of tallherb forests on the large areafrom the northern boundaries of forest belt to thesouthern taiga and subtaiga, their considerableecotypic variety, and the ability to survive under severeclimate conditions while preserving the nemoral andboreal flora species allows us to assume that these par�ticular forests represented in large part the prehistori�cal outlook of taiga forests in the studied area.

This suggestion may be regarded as an argument infavor of the autochthonous origin of forests. On theother hand, the earlier data on the highest degree ofbiological variety, productivity, and intensity of biolog�ical turnover among the so�far analyzed dark conifer�ous taiga forests of the studied area allows us to con�sider tallherb forests to be the most important objectsin estimating the ability of the forest cover to fulfill itsecosystem functions.

ACKNOWLEDGMENTS

This work was supported by the Russian Founda�tion for Basic Research, project no. 10�04�00355a,and the Fundamental Research Program, RussianAcademy of Sciences Presidium “Wildlife: CurrentStatus and Problems of Development.”

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Translated by S. Korobkova

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