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	<title>№1 2022 &#8211; ВОПРОСЫ ЛЕСНОЙ НАУКИ/FOREST SCIENCE ISSUES</title>
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		<title>WILDFIRES AS A FACTOR OF LOSS OF BIODIVERSITY AND FOREST ECOSYSTEM FUNCTIONS</title>
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					<description><![CDATA[Original Russian Text © 2021  A. P. Geraskina, D. N. Tebenkova, D. V. Ershov, E. V. Ruchinskaya, N. V. Sibirtseva, N. V. Lukina  published in Forest Science Issues Vol. 4, No. 2,   Article 82 &#160; A. P. Geraskina*, D. N. Tebenkova, D. V. Ershov, E. V. Ruchinskaya, N.&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/05/Wildfires.pdf"><img loading="lazy" class="size-full wp-image-1122 alignright" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></span></p>
<div style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000; font-size: 10pt;"><span lang="EN-US"><span style="color: #000000;">Original Russian Text © 2021  A. P. Geraskina, D. N. Tebenkova, D. V. Ershov, E. V. Ruchinskaya, N. V. Sibirtseva, N. V. Lukina  published in Forest Science Issues Vol. 4, No. 2, </span>  </span><a href="https://jfsi.ru/4-2-2021-geraskina_et_al/" target="_blank" rel="noopener noreferrer">Article 82</a></span></div>
<p>&nbsp;</p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>A. P. Geraskina</strong><strong><sup>*</sup></strong><strong>, </strong><strong>D. N. Tebenkova</strong><strong>, D. V. </strong><strong>Ershov</strong><strong>, </strong><strong>E. V. Ruchinskaya</strong><strong>, </strong><strong>N. V. Sibirtseva, </strong><strong>N. V. Lukina</strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Center for Forest Ecology and Productivity of the Russian Academy of Sciences </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>117997 Moscow, Russian Federation, Profsoyuznaya st. 84/32 bldg. 14</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><sup> </sup></strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><sup>*</sup>E-mail: angersgma@gmail.com</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 07.07.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 12.08.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 18.08.2021</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Due to the ever-increasing anthropogenic impact and global climate change, wildfires are becoming more frequent and intense all over the world. The wildfire factor is turning into an acute problem for forested countries that requires prompt solutions as the areas of forest ecosystems are reducing catastrophically, which results in an irreparable loss of biodiversity that provides all ecosystem functions and services. Many biologists consider wildfires a factor destructive to biota that results in permanent loss of some species and groups of living organisms; even if it is possible for them to recover after a wildfire, they may need a lot of time to do so. However, some studies argue that wildfires do not reduce the biodiversity in forest ecosystems, but even increase it, thus contributing to species conservation and sustainable functioning of forests.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">This article is aimed at analyzing the studies of how wildfires impact the main components, biodiversity, and functions of forest ecosystems. The authors answer the question of why wildfires while being a destructive factor, are sometimes considered a factor increasing biodiversity. The “positive” influence wildfires have on biodiversity mostly comes down to the formation of mosaic patterns, that is, forest canopy gaps that occur after a wildfire. However, analysis of references shows that the established opinion found in a number of studies that a certain frequency of wildfires is necessary to maintain forest communities may be associated with ignoring or misunderstanding the importance of biotic factors in the functioning of forests. In modern forest ecosystems, populations of keystone large mammal species have disappeared or greatly declined; therefore, there are no microsites they usually form, including large forest canopy gaps (glades) that provide opportunities for photophilous flora and pollinating insects to develop and generally maintain adequate conditions for multi-aged polydominant forest ecosystems with high biodiversity. In the forestry practice, there are measures to maintain mosaics. They include special types of felling, supporting populations of keystone animal species, etc., and are both significantly less catastrophic in comparison with the wildfire factor and substantiated biologically. The authors provide recommendations for the conservation and maintenance of biodiversity and ecosystem functions in modern forests.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Keywords:</strong><em> forest, fires, vegetation, animals, keystone species, greenhouse gases, soil, climate, carbon, ecosystem services, emissions </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires are not only a modern global factor determining the state and functioning of forest ecosystems, exerting a powerful influence on the biogeochemical carbon cycle, hydrological regime and climate change, but also a historical factor in forest formation. The interaction of man and nature has been closely connected with fire since the middle of the Pleistocene (500 thousand years ago): drive hunting, slash-and-burn agriculture, fire clearing for meadows and pastures (Gowlett, 2006; Bowman et al., 2009; Bobrovskij, 2010; Tang, Yap, 2020; MacDonald et al., 2021). Therefore, when assessing the biodiversity of modern forests and the effectiveness of their ecosystem functions, it is necessary to take into account the anthropogenic history, in which fires in many territories were the most important factor of forest formation (Whitlock et al., 2010; Aleynikov et al., 2015). Currently, despite fundamentally different technologies in economic activity, wildfires remain an acute problem for forest countries, which requires solutions both in connection with global climate change and with a number of economic issues, such as loss of ecosystem services provided by forests, loss of forests as an important component amidst decarbonization of the economy. Many biologists consider wildfires as a destructive factor for biota, with slow recovery after exposure. If fragmentary “refugia” are preserved during a wildfire, in which individuals of different species survive, this does not necessarily mean that populations survive (Gongalsky, 2014). Therefore, the following consequences are seen:</span></p>
<ul style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">long-established coordinated functional relationships based on biodiversity are destroyed;</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">plant edificators are suppressed and populations of keystone animal species of above-ground and underground biota are reduced</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">the ecosystem is thrown back to historically earlier stages of development and a round of fire-induced demutational succession is triggered; at a high frequency of wildfires, this leads to persistent digression and the formation of post-fire communities with limited species diversity.</span></li>
</ul>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">At the same time, both in biology and forestry, there are ideas that wildfires are necessary, for example, for the germination of seeds of some plant species (Bell et al., 1993; Keeley, Fotheringham, 2000), the maintenance of pine and oak plantations (Cvetkov, 2013), etc. Currently, authors of some studies claim based on their findings that wildfires not only do not reduce, but also increase the biodiversity of forest ecosystems, and extinguishing large wildfires, in general, is economically impractical (Stephens et al., 2018; Kharuk et al., 2021). One of the arguments is that wildfires had also occurred prior to the beginning of global human influences on nature; therefore, they are necessary as a formation factor of forest ecosystems and even the evolution of biota (He et al., 2019). However, it should be kept in mind that, at present, the frequency, intensity and scale of wildfires (ninety percent of which, according to experts, occur due to human activity even in the most remote areas) have increased significantly, and this is exacerbated by the impact of climate change. The type of evolution of forest ecosystems under the influence of wildfires can be defined as “erasing evolution”, according to the definition of L. G. Bogatyrev (2004), proposed for the development of forest litter.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The objective of this article is to analyze the results of studies of the impact of wildfires on the main components of forest ecosystems, their biodiversity and functions and to answer the question why wildfire as an obviously destructive factor is sometimes considered as a factor of increasing biodiversity.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE SCALE OF WILDFIRES AND FIRE-INDUCED EMISSIONS OF CARBON COMPOUNDS IN THE FORESTS OF RUSSIA</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>The scale of wildfires </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to official statistics, 569.912 sites of wildfires were registered in the territory of the state forestry of the Russian Federation in 1992–2012, which averaged 26.805 foci per year (EMISS, 2021a). In 2009–2020, the area of state forestry lands covered by wildfires amounted to 43.945 million hectares (an average of 3.662 million ha per year) (EMISS, 2021b). Damage from wildfires in 2019, according to official statistics, amounted to 13.5 billion RUB (EMISS, 2021c). At the same time, according to various estimates, the proportion of major wildfires (with the area of more than 200 ha) in Russia is about 5% of the total, but their contribution by area is about 95%. In the forests of Russia, surface fires occur and spread most often, accounting for up to 98% of the total number of wildfires and more than 88% of the area covered by fire, whereas crown fires account for 1–2% and 12%, respectively (Isaev et al., 1995).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The data of satellite monitoring of wildfire areas, provided by various Russian and foreign experts, differ significantly from official statistics. Thus, A. Z. Shvidenko and D. G. Shchepashchenko, who have investigated the influence of climate on the wildfire situation in Russia in 1998–2010, cite data from various sources. On average, according to their estimates, the area of fires during this period was 8.5 million ha per year (Shvidenko, Shchepashchenko, 2013). From time to time, years with an abnormal frequency of fire occurrence with an area of up to 16–18 million ha are registered. Other authors (Lupyan et al., 2017) report that using satellite data, 5 to 20 thousand wildfires were registered annually in 2001–2016 in Russia, damaging forests with an area of 5–20 million ha. Similar estimates are given in the works of other Russian researchers (Ponomarev, Shvecov, 2015; Bondur et al., 2016).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Types of fire emissions and their assessment by surface methods</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A significant contribution to the emissions of greenhouse gases (СО<sub>2</sub>, СН<sub>4</sub>, N<sub>2</sub>O) and gases with an indirect greenhouse effect (CO, NO<sub>x</sub>, non-methane volatile organic carbons) and other compounds are made by wildfires that occur annually in the forests of Russia over vast territories and often turn into natural disasters. The impact of wildfires on the carbon balance is determined by two main processes: the physicochemical process of “rapid” release of carbon compounds formed during incomplete combustion of organic matter (“fire” emissions) and the biological process of “slow” release of carbon compounds due to destruction and rotting of plants that died from wildfire, but had not been burnt (“post-fire” emissions). Fire emissions occur directly during the wildfires and can last from several hours to several days or weeks. Post-fire emissions begin with the death of woody plants and continue for several years or decades.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Surface studies of the intensity of combustion and the expenditure of various combustion conductors of forest fuels (FFs) show that the mass of above-ground FFs varies depending on the species and age of plantings, their productivity and degree of closure (completeness), the forest plant zone and the phenological state of vegetation. It usually ranges from 4.0 to 12.0 t × ha<sup>–1</sup>, which corresponds to the stock of needles, dry and small branches in the canopy (crowns) of coniferous stands most susceptible to wildfires (Molchanov, 1954; Kurbatskij, 1972; Grishin, 1981). Taking into account incomplete burning (not completely burnt, partially charred FFs), the mass of above-ground FFs burning during crown fires on average is about 7.0 t × ha<sup>–1</sup>.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The mass of above-ground FFs formed from living ground cover (mosses, lichens, shrubs) and litter (needles, leaves, small branches, etc.) varies widely depending on the species composition, age and closeness of stands, forest type, nutrient and water regime of soils. In most cases, FF stocks in this group range from 2.0 to 15.0 t × ha<sup>–1</sup> (Vonskij, 1957; Konev, 1977). Taking into account incomplete burning, the mass of ground FFs burning during surface wildfires is 5 t × ha<sup>–1</sup>.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The mass of litter and organic soil horizons, consisting of dead parts of plants with varying degrees of decomposition and humus, in forest ecosystems usually varies in the range from 5.0 to 25.0 t × ha<sup>–1</sup> (Molchanov, 1954; Vonskij, 1957). In most cases of crown and surface wildfires, the depth of burning does not exceed half the thickness of the forest litter layer, which corresponds to stocks of 3.0–12.0 t × ha<sup>–1</sup>. In case of ground fires that occur in swamps and swampy forests with a developed peat horizon, the mass of organic materials involved in burning can be up to 150 t × ha<sup>–1</sup> or more (Arcybashev, 1974; Sheshukov, 1979).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The stock of FFs from deadwood residues (deadwood, dead standing trees, stumps, dry branches) can reach several tens of tons per hectare. Most often, no more than half of the available stock of deadwood residues are burnt, which is commensurate in weight with the stock of living ground cover in forest areas.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Taking into account the above assumptions and stocks of the main FF groups, the mass of burning organic materials per ha of the area covered by wildfire is 30 t × ha<sup>–1</sup>, 12 t × ha<sup>–1 </sup>and 120 t × ha<sup>–1</sup>, respectively, for crown, surface, and ground wildfires.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Remote estimates of carbon emissions from wildfires</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Quantitative estimates of direct fire emissions of carbon compounds and other greenhouse gases using satellite data differ by different researchers and are related to the methods of wildfire recognition and their consequences, models for measuring and estimating greenhouse gas emissions, as well as auxiliary data on Russian forests (maps of vegetation, woody fuels, etc.).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Direct measurements of fluxes and concentrations of gases (the “top-down” approach) in the Earth’s troposphere are performed using satellite instruments (Amiro et al., 2001a; Liu et al., 2005).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The conventional common “bottom-up” approach is also used, which is based on post-processing of satellite data on fires (area and degree of fire damage of vegetation) and data on stocks of plant combustion conductors of various types of wooody fuels (Isaev et al., 2002; Kasischke, Bruhwiler, 2003; Soja et al., 2004; Wiedinmyer et al., 2006; Sochilova, Ershov, 2007).</span></p>
<ol style="text-align: justify;" start="3">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">I. Ponomarev et al. use brightness temperature in the 3rd MODIS thermal channel (3.93–3.99 μm) to assess the intensity and type of wildfire, as well as its relationship with the FF consumption for various wood residues estimated according to literature (Ponomarev et al., 2017). The estimates of direct carbon emissions presented by the author for the time period 2002–2016 averaged 83 ± 21 Mt C per year<sup>–1</sup>. The range of variation of direct carbon emissions in different years was 20–227 Mt C per year<sup>–1</sup>. A. Z. Shvidenko and D. G. Shchepashchenko estimate the number of carbon emissions during 1998–2010 due to wildfires in Russia at 121 ± 28 Mt C per year<sup>–1</sup> with annual variability of 50 (2000) to 231 (2003) Mt C per year<sup>–1</sup> (Shvidenko, Shchepashchenko, 2013). Looking at some rough estimates of post-fire carbon emissions from wildfires of approximately 90–100 Mt C per year<sup>–1</sup> (Shvidenko et al., 2010), the authors estimate total carbon emissions due to wildfires in recent decades at 180–200 Mt C per year<sup>–1</sup>.</span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to our estimates, direct fire carbon emissions in 2002–2018 amounted to 34 ± 19 Mt C per year<sup>–1</sup>, ranging from 12 (2009) to 127 (2003) Mt C per year<sup>–1</sup> (Ershov, Sochilova, 2020). At the same time, the areas of forest damage and the intensity of direct fire-induced carbon emissions increased 1.4 times after 2012. Until 2012, the average damage area and emissions were 3.95 million ha and 29.18 Mt C, whereas over the past 9 years those figures were 5.73 million ha and 41.07 Mt C, respectively. Differences in estimates as compared to other authors are due to the fact that only data from forest ecosystems (forested areas) are used, and there are no direct emissions data for large wood residues due to the lack of spatial data throughout Russia.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, the extent of the forest area covered by wildfire and the amount of direct fire-induced emissions are evidences of a significant impact of wildfires on the state and biological diversity of forest ecosystems in Russia. Surface fires occur and spread most often in the forests of Russia, both in terms of the total number of wildfires and the area covered by fire, whereas major wildfires (with an area of more than 200 ha) make a significant contribution to the emissions of carbon compounds and other greenhouse gases. In addition to fire emissions corresponding to the duration of forest burning, post-fire emissions occur, which last for several years or decades.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>PREREQUISITES OF IDEAS ABOUT WILDFIRES AS A FACTOR INCREASING BIOLOGICAL DIVERSITY</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Modern forest ecosystems differ significantly from pre-anthropogenic forest-meadow systems that existed before the beginning of the Holocene when mass destruction of keystone animal species by humans occurred during the development of appropriating economy (Smirnova et al., 2021). In modern forests, biological diversity, including functional and structural, is reduced as compared to prehistoric forests (Vera, 2000; Orlova, 2013; Korotkov, 2017; Lukina et al., 2020). The mosaic of microsites of pre-anthropogenic forests was a result of treefalls or breaks due to either the natural death of trees or the activity of large vertebrates, which formed much larger gaps (breaks in the canopy of the forest) and clearings than the falls of single trees. Large phytophages had a great influence on the undergrowth of trees and shrubs through uneven grazing and trampling. As a result, a stand of different composition and different ages was formed (Vera, 2000). The renewal of light-demanding flora was not limited by the lack of light. Mammals and birds contributed to the spread of seeds, created additional micro-habitats for companion species, such as small mammals, insects and other invertebrates. Mosaic nanorelief was formed with different soil moisture and composition of soil fauna (Puchkov, 1992).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Currently, especially in boreal forests, the renewal of light-demanding flora is limited by a lack of light due to the continuous canopy of dark coniferous tree species, which is probably why a number of works claim that the preservation of modern pine, oak, and larch plantations is ensured by wildfires (Sannikov, 1997; Cvetkov, 2013; Robertson et al., 2019; Matveeva, 2020). However, there are studies showing that wildfires of any intensity also inhibit the renewal of pine trees (Allen et al., 2002; Makarov et al., 2016). According to available data, intra-forest clearings make a significant contribution to the floristic diversity of forest ecosystems (Smirnova et al., 1997; Evstigneev et al., 1999; Gornov et al., 2020). Succession changes of woody vegetation occur in the direction from light-demanding species to shade-tolerant, and a new demutation process is started after disturbances, such as blow-down, fire, logging, insect epidemics. However, after such large-scale disturbances, an even-aged stand with a small set of tree species that is vulnerable to external factors will be formed again.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Great importance in modern forests is assigned to deadwood as a common microsite of old-growth forests. Deadwood supports floral diversity (Evstigneev et al., 2012; Evstigneev, Gornova, 2017; Khanina, Bobrovsky, 2021), is a favorable habitat for dozens of species of vertebrates and hundreds of species of invertebrates, as well as fungi and bacteria (Goncharov, 2014; Geraskina, 2016; Ashwood et al., 2019; Evstigneev, Solonina, 2020; Jacobsen et al., 2020), which is especially relevant in the face of accelerating rates of loss of biological diversity (Lukina et al., 2021). Despite the fact that deadwood, especially in the late stages of decomposition, usually has higher humidity than the surrounding soil, it is also currently considered as a factor of increased fire danger (Paletto et al., 2012). This indicates a high degree of disturbance and vulnerability of modern forests since they practically lack such keystone species as moose, bison, beavers, etc., therefore, no natural barriers to the spread of wildfire are created due to the formation of gaps, trails, understocking, or intra-forest reservoirs. Felling of individual trees and creating gaps in order to prevent the spread of wildfire is recommended as one of ecological principles of wildfire protection (Allen et al., 2002).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Since fire is a historically long-standing factor, adaptations to wildfires have formed in a number of plants, i. e. significant thickening of external protective tissues of woody plants, activation of the seed bank of flowering plants under the influence of high temperatures (Keeley, Fotheringham, 2000; Lamont et al., 2018; Soos et al., 2019), opening of cones of gymnosperms (Sannikov, 1997; Agapov, 2019). For example, the giant sequoia (<em>Sequoiadendron giganteum</em>) is, in the big scheme of things, a fire-dependent plant, since it is generally believed that the cones of this species open only after exposure to wildfire (Harvey, Shellhammer, 1991). However, there are also natural biotic factors that ensure the spread and germination of seeds. Pine and cedar cones are eaten by birds (nutcrackers, jays), mouse-like rodents and squirrels, who release seeds from under the dense scales and make a stash in the litter and burrows, a large part of which most often is not found, so the seeds germinate (Rejmers, 2015). Giant sequoia cones serve as food for Douglas squirrel (<em>Tamiasciurus douglasi</em>), whose main food is the green scales of young sequoia cones, because the seeds are very small and have less nutritional value than large scales. The longhorn beetle (<em>Phymatodes nitidus</em>) is trophically very closely related to the cones of the giant sequoia: female beetles lay eggs on the surface of the cones, and hatching larvae eat the scales of the cones and release seeds (Weatherspoon, 1990). Besides, scales of cones dry and crack and the seeds fall down after exposure not only to wildfire, but also to direct sunlight, however, under the closed canopy of the stand due to the lack of open spaces as a result of extermination of large forest animals, this mechanism is often not implemented (Harvey et al., 1980).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The positive impact of wildfire on forest biodiversity is also believed to include:</span></p>
<ul style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">reduced root competition among different tree species (Matveeva, 2020),</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">improved seed germination due to burning of the forest litter to the mineral layer (Karnel’, Zabelin, 1978) and a decrease in number of small mammals that may damage seeds and plant sprouts (Farber, 2012);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">accelerated mineralization of organic matter (Wells et al., 1979);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">antiseptic effect of high temperatures on soils (Sokolov, 1973);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">reduced competition for light and precipitation on the burnt landscape (Agapov, 2019).</span></li>
</ul>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">All these arguments are quite well supported by functional losses in the biodiversity of modern forests, since these effects implement biotic relationships between the components of forest ecosystems: the destruction of litter is provided by invertebrate saprophages and saprotrophic microorganisms, which also complete its mineralization and have a “sanitation” effect on soils, regulating the balance of different groups of bacteria (Byzov, 2005), the formation of structural diversity and reduction of competition between plants, including underground (root systems) provide zoogenic mechanisms in forest regulation (Puchkov, 1992; Vera, 2000; Smirnova et al., 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, in modern forests, where keystone species of large mammals have been lost together with the microsites formed by them and providing opportunities for the formation of multi-age polydominant forest ecosystems, wildfires are often considered as an important and necessary factor in maintaining biodiversity. Wildfires trigger positive feedback mechanisms; therefore, some forest communities (for example, pine forests) are now classified by researchers as fire-dependent. A number of plants have developed adaptation mechanisms to fire exposure. However, biotic factors play a high role in the functioning of forest ecosystems and the maintenance of biodiversity, and it must be taken into account when considering approaches to sustainable forest management and, if possible, lost ecosystem components should be restored.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE IMPACT OF WILDFIRES ON PLANT COMMUNITIES</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfire affects plants directly by destroying them completely or partially, as well as indirectly through changes in living environment. Therefore, short-term and long-term effects of wildfires are distinguished. The short-term ones include the combustion of forest fuels, including phytomass, heating of the soil, burns (fire wounds) or death of plants, terrestrial vertebrates and soil animals, microorganisms (Melekhov, 1948; Wildland&#8230;, 2000; Il’ina, 2011; Suhomlinov, Suhomlinova, 2011, etc.). The long-term consequences of wildfires include fire-induced soil transformation, reduction of soil biota diversity, drying out and death of trees, accumulation of phytomass, post-fire succession of vegetation (Kuleshova et al., 1996; Monitoring&#8230;, 2002; Tyler, Spoolman, 2011; Gorbunova et al., 2014; Ivanova et al., 2018, etc.).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Crown wildfires, when the fire spreads from the soil to the tops of trees, are the most destructive ones for forest vegetation. Crown fires can be running and independent (Zalesov, 2011; Il’ina, 2011). An independent wildfire is a disaster for the entire plant community, as it affects all its components. After the death of forest due to impact of a wildfire, there are sharp changes in the microclimate, hydrological and soil conditions, which, in turn, affect the formation of a new community depends, i. e. cause a change of phytocenoses. In some cases, the stand dies completely and falls out in a short time, forming blockages (Nesgovorova et al., 2015). Sometimes vegetation recovery is delayed due to severe burning of soils and lack of seed sources.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In case of surface wildfires, plants of the lower layers (moss-lichen and grass-shrub tiers, understory and undergrowth), as well as litter and humus horizon partially or completely burn out. Root systems are damaged, fire wounds form on tree trunks (Devyatova et al., 2014; Richter et al., 2019), deadwood, stumps and felling residues partially burn out. Surface wildfires, under some circumstances, can turn into crown wildfires. Fire-damaged and weakened trees are more severely damaged by insects and fungi (Melekhov, 1948; Popov, 1961; Parker et al., 2006). However, some studies argue that low-intensity wildfires can have a positive effect on the ability of some trees to protect themselves from insects, for example, the Eastern larch beetle (<em>Dendroctonus simplex</em>) (Hood et al., 2015). After surface fires, the understory mostly dies. The study by K. V. Levchenko (2017) emphasizes that the resistance of coniferous forests to surface wildfires is very low. In communities with understory and undergrowth, in the presence of slopes, a surface wildfire can turn into a crown one, and all components of the phytocenosis, including the ground cover, are completely destroyed.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Surface wildfires of different intensity affect vegetation differently (Pourreza et al., 2014; Ivanova et al., 2018). There are wildfires of low, medium and high intensity, which differ in the degree of burning out of litter and soil. After a weak impact, the stand is preserved, while the fire hazard of the territory is reduced for some time due to a decreased supply of fuels. After low-intensity wildfires, the abundance and diversity of grasses and mixed herbs may increase (Hutchinson et al., 2005). This is believed to be associated with the emergence of new ecological niches (Rosenzweig, 1995; Gorbunova et al., 2014). Medium-intensity wildfires, as well as low-intensity wildfires, weaken the stand and lead to the loss of trees (Ivanova et al., 2018). After high-intensity wildfires, the recovery time of the post-fire community is many times more (Ivanova et al., 2017). They significantly disrupt landscapes (Collins, Stephens, 2010) and lead to pronounced homogenization of the habitat, which significantly reduces biodiversity (Hessburg et al., 2016; Shive et al., 2018, Steel et al., 2018). Besides, after intense wildfires, the reserves of ground-based fuels increase and may exceed the pre-fire figures several times, providing conditions for the recurrence of a high-intensity wildfire (Ivanova et al., 2017). Sometimes, after such fires in high-light conditions, massive sprouting of woody plants is observed (Ivanova et al., 2018). However, due to increased soil temperature, insufficient moisture and infection with phytopathogens, these seedlings die. The understory is restored after 12–14 years.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Often after wildfires in forest communities, the proportion of light-loving plants increases, i. e. of pine forest and meadow species (Ivanova, Perevoznikova, 1996; Bizyukin, 1998), whereas, in some cases, the proportion of meadow-steppe species is increased (Shpilevskaya, Katkova, 2011)<em>. </em>Moreover, the so-called pyrophytes often actively invade the burnt-out areas (Vostochnoevropejskie&#8230;, 2004; Afanas’eva, Berezina, 2011), the emerging “diversity” being qualified as pyrodiversity (Не et al., 2019). It is believed that some plants have adapted to survive wildfires (Kelly, Brotons, 2017). These include, for example, the structure of seeds, which keeps the embryo alive after being exposed to wildfire, as well as the thick bark of trees that protects the cambium (Il’ina, 2011). Often, pyrophytes include fireweed (<em>Chamaenerion angustifolium</em>), which inhabits post-fire areas and forms closed plant aggregations (Bizyukin, 1998; Afanas’eva, Berezina, 2011; Shpilevskaya, Katkova, 2011). Pyrogenic communities may also be invaded by adventitious and ruderal species (Goryainova, Leonova, 2008; Shpilevskaya, Katkova, 2011).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfire causes change in the vegetation composition of affected areas; that is, it leads to the formation of post-fire (pyrogenic) successions. They depend on the composition and condition of the initial community, fire intensity and duration (Kuleshova et al., 1996; Ivanova et al., 2017; Miller et al., 2019). At the first stages, the community is populated by pioneer (reactive) species, “pyrophytes” can often spread. Diasporas can stem from a soil seed bank and plants from undamaged sites. With no adult woody plants, the settlement of burnt areas will depend on seed transfer by animals (birds and small mammals) (Diaci, 1994). The importance of vegetative reproduction of plants increases as well (Ivanova, Perevoznikova, 1996; Kovaleva et al., 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Although areas with higher illumination are invaded by pyrophyte species, wildfires always result in a decrease in plant species diversity (Chibilev, 1998; Il’ina, 2011; Richter et al., 2019). After wildfires, the stocks of seeds in the soil are significantly reduced (Il’ina, 2011; Miller et al., 2013). Rare flora may disappear entirely after wildfires (Kryukova, 2009; Makarov et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Post-fire recovery can take from several years to decades (Telicyn, Ostroshenko, 2008). Modern ecosystems are modified to varying degrees and are subject to anthropogenic impact (Richter et al., 2019). Therefore, the impact of wildfire on forests can manifest in different ways, depending on the composition of the original community and the history of wildfires in the specified area (Miller, Safford, <a style="color: #000000;" href="https://esajournals.onlinelibrary.wiley.com/doi/full/10.1002/ecs2.2882#ecs22882-bib-0056">20</a>20). In the review of D. A. Driscoll et al. (2021), wildfires and fragmentation of communities were shown to interinfluence depending on the conditions of interaction and its scale. For instance, after a wildfire, landscapes often become heterogeneous, while communities that have already survived such impact can restrain the spread of fire due to the areas covered by fire. Short-term increase in biodiversity that is observed in some cases is mainly due to the marginal effect.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires as a very powerful factor in development of forest ecosystems have had a huge impact on the modern appearance of boreal forests in both North America (Payette, 1992) and Eurasia (Gorshkov, 2001; Neshataev, 2017). Many researchers of boreal forests register the fact that in the modern vegetation cover of the taiga zone, most of the light and dark coniferous forests are not indigenous stands but various stages of forest recovery in the areas covered by fire (cited by Neshataev, 2017).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Modern dendrochronological studies show the influence of long-standing large wildfires (those that occurred more than a hundred years ago) on forest ecosystems. For example, the influence of a large wildfire in 1896 can still be seen in the growth pattern of trees and the depth of seasonal permafrost melting in Central Siberia. After the death of the stand and ground cover, there was a decrease in thickness of the organic soil horizon and an increase in thickness of permafrost, resulting in slow forest recovery after wildfires in most circumpolar boreal zones (Kirdyanov et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Amid climate change, the number of wildfires and their frequency will increase (Flannigan et al., 2000, 2006; Camia et al., 2017; Molina et al., 2019). Some post-fire systems may not restore the original composition of vegetation due to changes in soil conditions and the formation of deflation zones, despite reforestation already carried out (Gyninova et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, wildfires of any intensity have a direct and indirect impact on the stand, understory and ground cover. Wildfires change the functioning conditions of all components of plant communities and make them more vulnerable to other environmental factors. The state of coenopopulations of plants that prevailed in pre-fire ecosystems deteriorates. The advent of light-demanding “pyrogenic” species does not make up for the overall level of decline in biodiversity after wildfires. Post-fire vegetation restoration requires considerable time, available diaspora sources and carriers.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE EFFECT OF WILDFIRES ON VERTEBRATES</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Despite high relevance, there are not so many studies of the impact wildfires have on vertebrates, which is stated in a number of works (Strategiya&#8230;, 2011; Pushkin, 2014; Barlow, Peres, 2006; Pastro et al., 2014; Gertini et al., 2021). Assessment of the impact of wildfires on animal populations is mainly based on change in their density over time: if population density increases in a certain area, a conclusion about the positive impact of wildfire is usually drawn, and if population density decreases, wildfire is believed to have a negative impact; alpha and beta diversity as well as spatial distribution of animals are analyzed as well (Revuckaya et al., 2018; Belyh et al., 2021; Cleary et al., <a style="color: #000000;" href="https://onlinelibrary.wiley.com/doi/full/10.1111/geb.12195#geb12195-bib-0013">2004</a>; Pastro et al., 2011; 2014).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires destroy habitat and food resources for vertebrates and increase the efficiency of predator hunting in post-fire landscapes (Letnic et al., <a style="color: #000000;" href="https://onlinelibrary.wiley.com/doi/full/10.1111/geb.12195#geb12195-bib-0039">2005</a>; Green, Sanecki, <a style="color: #000000;" href="https://onlinelibrary.wiley.com/doi/full/10.1111/geb.12195#geb12195-bib-0029">2006</a>; Kodandapani et al., <a style="color: #000000;" href="https://onlinelibrary.wiley.com/doi/full/10.1111/geb.12195#geb12195-bib-0036">2008</a>). Wildfires can be detrimental to the physiology of small mammals, for example, making it difficult for them to reproduce, as it has been shown in Australia regarding some quolls and antechinus species. In fact, major environmental changes destabilize animals at such stages of reproductive behavior as courtship, pregnancy, and offspring care (Banks et al., 2007). The impact of fire on individual animal species depends on the intensity and scale of wildfires (Cleary et al., <a style="color: #000000;" href="https://onlinelibrary.wiley.com/doi/full/10.1111/geb.12195#geb12195-bib-0013">2004</a>; Pastro et al., 2011).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the oak forests of Pennsylvania, 4–12 months after the fire, the number of small mammals in the burned forests was significantly less than in the unburned forests, and two rodent species, i. e. <em>Microtus pennsylvanicus</em> and <em>Clethrionomys gapperi, </em>were not found at the fire sites (Kirkland et al., 1996). In a burned-out area of 15.000 ha in Arizona, the number of rodents of the Cricetidae family declined due to fire-induced disturbance of grass cover and returned to the pre-fire level only 6 years later (Bock et al., 2011). The abundance and diversity of small mammals in some parts of the eucalyptus forest in Australia recovered at least 9 years after a wildfire (Fox, McKay, 1981).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In some cases, the “benefits” of wildfires are listed for animals such as <em>Cervus elaphus</em> and <em>Alces alces</em>, which feed on herbaceous plants and understory of trees that appear on overgrowing fire sites (Kharuk et al., 2021). At first glance, this is corroborated by the established positive correlation between the increase in number of herbivores and the area covered by fire (Belyh, Sadovskaya, 2021). However, according to the authors of the study themselves, such a correlation may be due to forced migration of animals to the burnt-out areas from areas where the forest is still burning, in an attempt to escape the fire. The same explanation might be true for animals of the Canidae, Felidae, Ursidae, and Phasianidae families (Belyh, Sadovskaya, 2021). In the boreal forests of North America, foxes are more common at fire sites than wolves, which, however, reclaim the territories quite quickly. The dynamics of lynx population is largely determined by the population density of hares that are lynx’s main prey (Fisher, Wilkinson, 2005).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In some studies, authors suggest that the effect of wildfires on animals is neutral (Pastro et al., 2014). For example, E. P. Lipatnikov, O. P. Vin’kovskaya (2012) did not find any dependence of the population of wild boar (<em>Sus scrofa sibiricus</em>) on the size of the areas covered by fire. At the same time, the very activity of wild boars affects wildfires: rooting damage caused by wild boar limits the spread of surface fires and protects woody understory (Lipatnikov, Vin’kovskaya, 2012), acting as a mineralized shelterbelt. At the same time, O. L. Revuckaya et al. (2018) found that the highest population density of wild boar, as well as Manchurian wapiti (<em>Cervus elaphus xanthopigus</em>), is recorded in areas with the least frequency of fire occurrence. Studies of the effect of controlled burning in <em>Pinus palustris</em> communities in the south-eastern United States on small mammals and amphibians have not revealed significant differences in the number of animal species depending on the frequency of ignition: intervals of 1–3, 3–5 and more than 5 years (Darracq et al., 2016).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires have an extremely negative impact on Siberian musk deer (<em>Moschus moschiferus</em>) as their population on the burnt areas declines sharply, sometimes to the extent of disappearance, and does not recover for a long time (Domanov, 2017); sun bear (<em>Helarctos malayanus</em>) in South-East Asia (Fredriksson et al., 2007), tiger (Joshi et al., 2015), Indian elephant (Joshi et al., 2015), Amur leopard (Pikunov et al., 2009) and other rare mammals.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Most researchers are unanimous in their negative assessment of the impact of landscape wildfires on representatives of the Mustelidae family, in particular, sables (<em>Martes zibellina</em>) (Naumov, 2014; Pushkin, Mashkin, 2014; Revuckaya et al., 2018; Fedorova et al., 2020; Belyh, Sadovskaya, 2021). The work “Wildfires in the Siberian taiga” (Kharuk, 2021), on the contrary, argues that sables are attracted by overgrowing fire sites due to growing populations of hares and small mouse-like mammals they feed on. However, in the years of the maximum number of wildfires, there is a decrease in the number of sable populations (Fedorova et al., 2020; Belyh, Sadovskaya, 2021). Apparently, this is due to sable behavior in a wildfire. According to P. P. Naumov (2014), during a wildfire, sables do not try to escape from the impending fire, but hide. Therefore, they die from exposure to fire or smoke. During a crown wildfire, up to 100% of sables die (Naumov, 2014). Huge empty spaces remaining in the areas covered by crown fires cause damage to sable populations, hindering their reproduction and creating prerequisites for reduction of their range and population (Naumov, 2014). Damage caused by the destruction of the habitat of sables as a result of wildfires of 2019 in Krasnoyarsk Krai is estimated at more than 22 billion RUB (<a style="color: #000000;" href="https://greenpeace.ru/author/mihail-kreindlin/">Krejndlin</a>, 2019). These calculations show the lameness of conclusions about economic inexpediency of extinguishing wildfires. A negative effect of wildfires was also found for <em>Sciurus vulgaris</em> (Revuckaya et al., 2018) and <em>Lynx lynx</em> (Bekshaev, 2016).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires have a negative impact on populations of forest birds, especially highly specialized species (Bendel et al., 1974; Gil-Tena et al., 2009). Given the practice of burning felling residues, studies are being conducted on effects of such burning on birds nesting on clear cutting sites. Destruction of nests and death of broods are often noted, as well as forced abandonment of their nests by birds, including those that nested near the territory exposed to fire. However, despite the data obtained, some authors recommend the “method of controlled burning of felling residues on clear cutting sites in mountain forests as not causing significant change in animal communities” (Timoshkina, 2004). Considering that burning of felling residues during a fire season often leads to large wildfires (Yaroshenko, 2021), the negative effect of such burning can significantly increase.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Representatives of the herpetofauna (amphibians and reptiles) die from fire, smoke and oxygen starvation, despite the fact that these animals can potentially escape its influence. However, even fast-moving snakes and lizards get irreversible injuries, their shelters are destroyed and their food supply is depleted (Pausas, 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, open spaces with green food, including those that result from wildfires, can indeed attract large phytophagous animals and predators that feed on them. But in ecologically balanced ecosystems, such spaces arise and are maintained at the expense of keystone species (Vostochnoevropejskie lesa&#8230;, 2004). The heterogeneity of environmental conditions necessary to maintain biodiversity is created as a result of the population life of animals and plants, whose activities do not lead to catastrophic disturbances and losses that are inevitable after exposure to wildfire. In addition, often as a result of a large wildfire, huge homogeneous open spaces are formed, leading to the destruction of the natural heterogeneity of the living cover and, as a result, to a steady decline in biodiversity, including vertebrates.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE EFFECT OF WILDFIRES ON SOIL PROPERTIES</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Pyrogenesis is one of the leading processes in forests that affect soil properties. Wildfires cause changes in morphological and physicochemical properties, the composition of organic matter and mechanical composition of soils (Sapozhnikov, 1976; Trofimov, Bahareva, 2007; Kawahigashi et al., 2011; Dymov et al., 2014). Changes in morphological properties of soils are caused by burning out of organogenic horizons, loss of growing forest, deadwood and other plant residues and include formation of a pyrogenic horizon or appearance of signs of pyrogenesis in soil horizons. It has been found that morphological signs of fire influence can be found at a depth of up to 0.3 m (Dymov et al., 2018). Signs of pyrogenesis are manifested in the form of carbon-bearing inclusions in the lower part of the litter and mineral horizons, pyrogenic morphones. Signs of pyrogenesis include darkening of mineral horizons due to pyrogenic organic matter capable of active migration. The podzolic horizon becomes impregnated with mobile organic matter, hydrophobization is observed, and the upper mineral horizons are over-compacted.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires lead to decreased acidity of the litter and, on the contrary, increased acidity of the mineral horizons of soils, an increase in the content of exchangeable calcium in the mineral horizons of soils and their enrichment with carbon and nitrogen, a short-term increase in the availability of nutrients, a decrease in the biological activity of soils and the proportion of carbon of water-soluble compounds, a narrowing of the C : N ratio in the litter and other horizons that have experienced pyrogenic effects (Sapozhnikov, 1976; Sorokin et al., 2000; Certini, 2005; Bezkorovajnaya et al., 2007; Cibart, Gennadiev, 2009; Lukina et al., 2008; Dymov et al., 2014; Ludwig et al., 2018). The decrease in litter acidity on fire sites is associated with the influence of low-molecular organic compounds present in the soil solutions of the fire sites (Sapozhnikov et al., 2001). An increase in the carbon content is associated with its intake from burnt wood, an increase in the nitrogen content and exchangeable calcium is believed to be due to the massive intake of a large number of plant residues resulting from the impact of wildfires on woody and other plants.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Recent assessments of the effect of prolonged use of prescribed burning on the soils of south-western coastal plain pine forests in the United States demonstrate similar changes in their physicochemical properties. With an increase in the frequency of wildfires, the content of mobile calcium and manganese increases, the actual acidity, the content of potassium and sulfates in the ten-centimeter soil layer decreases (Coates et al., 2018). The authors believe these changes to be temporary. However, other authors demonstrate by the example of pyrogenic succession series lasting several hundred years that the effects of wildfires in the soils of forests in South Australia are observed after eighty years or more, and include depletion of soils with nutrients, in particular available phosphorus compounds and nitrates (Bowd et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">During wildfires in taiga biogeocenoses, there is a change and redistribution of organic matter pools between ecosystem components: a decrease in carbon and nitrogen reserves in the litter with their increase in the upper mineral horizons (Dymov et al., 2018). However, it should be emphasized that this increase in the carbon stock in the upper mineral horizons is accompanied by its huge fire-induced emissions into the atmosphere (section: The scale of wildfires and fire-induced carbon emissions in the forests of Russia).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires lead to change in the composition of soil organic matter. Due to fire, the content of hydrophilic organic compounds decreases and the content of hydrophobic compounds increases (Certini, 2005; Dymov et al., 2015a). Increased soil hydrophobic properties lead to an increase in surface runoff and intensification of soil erosion processes. Wildfires contribute to an increase in the pyrogenic horizons of the content and proportion of polycyclic aromatic hydrocarbons (PAHs), which have carcinogenic and mutagenic properties. Naphthalene, whose content increased especially significantly, was also found in pyrogenic morphones at a depth of more than half a meter (Dymov et al., 2015b).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The depth and scale of fire-induced changes in soil properties are, on the one hand, due to the nature of fire, its intensity, and on the other hand, due to the conditions (the level of soil moisture, precipitation, etc.) in which forests are formed, as well as types of forests.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In a changing climate, the frequency and intensity of wildfires are increasing. They lead to the release of carbon compounds from the buried organic matter of soils (legacy carbon) of boreal forests, which causes an increase in greenhouse gas concentrations and warming (Merzdorf, 2019). It has been shown that the restoration of litter in boreal forests after wildfires takes a lot of time (from 120 to 190 years) (Gorshkov et al., 2005).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Therefore, wildfires, the frequency and intensity of which are increasing in the modern circumstances of climate change, have a significant and negative impact on the properties of forest soils. As studies of long-term effects show, wildfires lead to reduced soil fertility, namely, to depletion of soils with available phosphorus and potassium compounds, to the release of carbon buried in the mineral horizons of soils, which causes a further increase in greenhouse gas concentrations. Wildfires contribute to an increase in soil hydrophobic properties and lead to an increase in surface runoff and intensification of soil erosion processes, as well as to an increase in the content of polycyclic aromatic hydrocarbons in soils, that have carcinogenic and mutagenic properties with inevitably detrimental effect on soil biota.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE IMPACT OF WILDFIRES ON MICROBIOTA AND SOIL INVERTEBRATES</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires have a destructive effect on the soil biota (Bowman, 1998; Doamba et al., 2014; Certini et al., 2021). Both crown and surface wildfires are dangerous, since both lead to the xerophytization of forest communities, which significantly changes the habitat conditions of both soil fauna and microorganisms. Charred wood (deadwood and tree trunks damaged by fire) is an unfavorable substrate for settlement of soil biota. Even among fungi, few species are known that can ensure the successful development of the pioneer stages of pyrogenic successions on wood (Safonov, 2006). In addition, direct burning of litter and deadwood leads to habitat loss for most species of soil biota. In general, wildfires reduce the biological activity of soils (Sorokin et al., 2000; Bezkorovajnaya et al., 2007; Sorokin, 2009; Sorokin, Afanas’eva, 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Various studies focused on the effect of wildfires on microorganisms, most of which were short-term and conducted in the first years after the fires (Ahlgren, Ahlgren, 1965; Min, Haiqing, 2002; Mataix-Solera et al., 2009; Silva et al., 2020). Wildfire can affect the soil microbiome directly, through heating, and indirectly, changing the properties of the soil. The most important factors include the intensity and duration of wildfire, as well as soil properties. In the event of an intense, prolonged fire, the top layer of soil can undergo complete sterilization. The activity of soil microorganisms also decreases due to changes in the quality of organic matter. After depletion of easily mineralized organic compounds, the initial increase in microbial basal respiration quickly goes into a decrease, since the preserved forms of carbon and nitrogen are more resistant to the effects of microbiota. The increase in pH (due to deposition of ash) is the reason for the increased bacteria/fungi ratio (Mataix-Solera et al., 2009; Pressler et al., 2019). After medium- and high-intensity wildfires, rapid recolonization of the soil by photoautotrophic microorganisms (algae) can occur (Mataix-Solera et al., 2009).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the middle taiga and southern taiga pine forests of Central Siberia, wildfires of medium and, especially, high intensity in the first year had a negative impact on the structure and functional activity of microbial complexes of sandy podzols. The number and biomass of nitrogen-carbon cycle microorganisms decreased, the qualitative composition became poorer, the enzymatic activity and intensity of microbial respiration decreased, the oligotrophicity of soils with respect to nitrogen increased (Bogorodskaya, 2006). A surface wildfire of moderate intensity led to the decreased metabolic activity of the microbial community in the litter of the pine forest of the Novosibirsk region in the first two years after exposure (Naumova, 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Analysis of the microbial community of Cambic Leptosols soils of Tolyatti pine forests after fires also showed that wildfires have a negative impact on the structure and metabolic activity of the microbial community of post-fire soil. It was found that the carbon content of microbial biomass and the rate of microbial respiration of the soil (in the upper organogenic horizons) of the sites after the wildfire significantly decreased as compared to the background figures (6.5 and 3.4 times, respectively). At the same time, at a depth of 10 cm in the soil, the effect of wildfire on these microbiological indicators has not yet been revealed (Maksimova et al., 2017).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires lead to a reduction in mycocenosis species diversity due to reduced quantity and quality of substrates (litter, wood residues) serving as a bank of spores and mycelium of fungi. The direct impact of wildfire on mycocenoses leads to a decrease in the species diversity of fungi. Burnt wood is slowly populated by xylotrophic fungi. As the deadwood accumulates after the fire, further development of mycocenosis occurs, but it goes in a direction different from the initial one (Safonov, 2006). Fungi are more sensitive to wildfires than bacteria (Pressler et al., 2019). Most studies of fungi forming arbuscular mycorrhiza have shown a negative effect (Mataix-Solera et al., 2009).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A meta-analysis of 1.634 field and 131 empirical studies of the impact of wildfires on microorganisms and mesofauna showed that wildfires have a strong negative impact on biomass, diversity, and distribution of soil biota. Wildfire reduces species richness and diversity of soil microorganisms and mesofauna by 88%–99%. The number of nematodes after wildfires is reduced by 88% (Pressler et al., 2019), Enchytraeidae — by 30–65% (Malmström et al., 2009), population and diversity of microarthropods are also reduced (Krasnoshchekova et al., 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The monograph of K. B. Gongalsky (2014) focuses on the influence of wildfires on soil fauna and provides an overview of the world literature on the influence of wildfires of different scales on soil fauna. The results of field experiments on artificial burning of forest areas are presented, which showed 100% death of invertebrates of the litter and upper mineral soil horizons (Wikars, Schimmel, 2001); laboratory experiments with direct fire exposure to soil samples for 1 minute without subsequent extinguishing showed a 46% decrease in the total number of macrofauna; the survival rate of spiders was 49%, rove beetles — 27%, larvae of soldier beetles, click beetles and chironomids — 58–62%, whereas all cicadas, caterpillars (Noctuidae and Pyralidae) and molluscs were killed by wildfire (Gongalsky et al., 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">During surface fires, the inhabitants of the litter and mineral horizons of soils at a depth of 2–3 cm below the burning area are killed; death occurs both directly from high temperatures during a wildfire, and in the first few days after the fire due to intoxication by combustion products (Wikars, Schimmel, 2001). In the wildfire zone, mass mortality of ticks, collembolans, testate amoebas, insects and earthworms, i. e. groups closely related to the organogenic horizons of the soil, is recorded. “Mobile” insect groups are more resistant to wildfires, i. e. flying zoophages and phytophages (Moretti et al., 2006). At the same time, at the egg stage, almost 95% of insects die, at the larva and imago stages — 60% (Gongalsky, 2014).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Surface wildfires of any intensity have a negative impact on earthworms. During field studies in European forests after wildfires, it was expected that epigeic earthworms would suffer the most, since they are closely related to the litter, but it turned out that endogeic worm populations declined most and were extremely slow to recover due to the fact that cocoons and juvenile individuals of this group are located in the uppermost horizons of the soil. Wildfires also had a negative impact on the anecic earthworms group (Certini et al., 2021). At the same time, epigeic worms, as more mobile, probably found refuge in the trees and other fragments of woody remains in the forests. In the forests of the Russian Far East, significant differences in the population of earthworms in terms of decreased number, biomass, species diversity and composition of morpho-ecological groups have been revealed in forests often prone to wildfires, as compared to less disturbed forests (Geraskina, Kuprin, 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Influence of wildfires on different taxonomic groups of meso- and macrofauna is a subject of numerous studies (Neumann, Tolhurst, 1991; Collett et al., 1993; Saint-Germain, 2005; Sackmann, Farji-Brener, 2006; Trucchi et al., 2009; Pressler et al., 2019; Gertini et al., 2021). Authors mostly report negative direct effects of wildfire on the density and species diversity of soil fauna, emphasizing their vulnerability and close relationship with the habitat. However, taking into account the indirect effects of wildfires, such as the emergence of open spaces, short-term development of microorganisms on mineralized due to fire organic residues, lack of competition in the first few years after a wildfire, etc., some authors report more favorable trophic and topic resources for individual taxonomic groups in the first years after a wildfire. For example, a number of Russian works show an increased diversity of ground beetles on fire sites in spruce forests: forest-meadow, meadow and field species appear, whereas the population of forest species of ground beetles decreases (Potapova, 1984; Uhova et al., 1999). At the same time, in the pine forests of Minnesota (Ahlgren, 1974) and the Spessart mountain range in Germany (Bauchhenss, 1980), a decrease in the diversity and population of ground beetles in the first two years after the fire was shown. A decrease in density and diversity of ground beetles in pine forests and an increase of these factors in spruce forests was found in Sweden, and the authors attribute this to better preservation of litter in spruce forests and its high humidity in comparison with pine forests. Insect larvae, earthworms, collembolans that ground beetles feed on have been preserved in the wetter litter. At the same time, the preservation of the diversity of ground beetles was directly correlated with the intensity of wildfire in both types of forest (Gongalsky, 2014).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the first years after wildfires, irruptions of ants can be observed in the fire sites, which is believed to be due to the presence of a large amount of wood residues and high adaptation of ants to xerophilic conditions (Bess et al., 2002; Krugova, 2010). At the same time, it is known that even crown wildfires have a negative impact on some species of ants (Arnan et al., 2006).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The restoration of soil biota diversity after a wildfire is very slow, especially in groups of animals with low migration abilities, such as earthworms, millipedes, or molluscs (Gongalsky, 2014). Restoration of the soil population is possible due to the heterogeneity of the soil cover and the preservation of perfugiums — areas poorly affected by wildfire, where some invertebrates survive during a wildfire. Along with the inhabitants of the deep layers of soil, they are the first to populate the fire sites (Gongalsky, 2006; 2014). Mobility of invertebrates is of great importance for the subsequent recovery of population; for example, recovery of collembolan groups living in mineral horizons is much slower compared to the population of ground beetles living in the litter (Mordkovich, Berezina, 2009). It has been shown that spring burnings are more dangerous than autumn ones for collembolans, larvae of dipterans, butterflies, parasitic wasps and earthworms. After spring burnings, most of the taxa recover within one year, the earthworm population — within 3 years after the fire (Neumann, Tolhurst, 1991).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The long-term effects of wildfires on soil fauna have been studied in less detail than the short-term effects (Gongalsky, 2014). It takes at least 10 years to restore micro- and mesofauna (Pressler et al., 2019). It has been shown that, for example, in the fire sites in the Oka Nature Reserve (Ryazan region, Russia), no complete restoration of the soil fauna occurred 20 years after the wildfire due to the fact that the litter horizon did not return to its pre-fire state (Potapova, 2002).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Therefore, xerophytization of forest communities after a wildfire, loss of microhabitats, direct impact of fire and smoke on soil biota and indirect influence through changes in soil properties and destruction of trophic relationships has a negative impact on the biotically consistent structure of soil fauna. Irruptions of individual species or an increase in the diversity of individual groups (ground beetles, ants and other insects) are of a short-term nature, limited by trophic resources that are rapidly depleted on fire sites, and occur due to the formation of open spaces available for settlement by species with high migration abilities from neighboring biotopes.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>THE IMPACT OF WILDFIRES ON ECOSYSTEM FUNCTIONS AND SERVICES OF FORESTS</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Consideration of fire issues in the context of related socio-ecological systems that recognize the links between people and their natural environment is very relevant in the light of the increase in the world’s population and, as a consequence, the increased demand for goods and services of forests. The terms “ecosystem functions” and “ecosystem services” are key in the concept of functional biodiversity. Ecosystem functions are a set of physical, biological, chemical and other ecosystem processes that support the integrity and conservation of ecosystems (Ansink et al., <a style="color: #000000;" href="https://link.springer.com/article/10.1007/s10531-017-1453-2#CR2">2008</a>). Ecosystem services are the benefits that people obtain from ecosystems, including provisioning services (fiber, wood, food, etc.), regulating services (erosion control, climate regulation, pollination, etc.), supporting services (soil formation, photosynthesis, etc.), cultural services (spiritual and religious, recreational, educational, etc.) (MEA, 2005). Forests simultaneously render forest ecosystem services (FESs) of all four categories, i. e. they are multifunctional (Byrnes et al., 2014; Manning et al., 2018; Van der Plas et al., 2018; Teben’kova et al., 2019). The transition to multifunctional forest management is considered as one of the key directions for achieving sustainable development of the forest-based sector (Bol’shakov et al., 2013). The multifunctional performance of forests can be considered at two levels: (1) the multifunctional performance of ecosystem functions, which are evaluated by fundamental studies of biological, geochemical and physical processes occurring in ecosystems; (2) the multifunctional performance of ecosystem services, which is defined as the joint provision of a number of ecosystem benefits in response to a request from society (Manning et al., 2018; Lukina et al., 2021). Taxonomic, functional, and structural biodiversity is the basis of multifunctional performance (Lukina et al., 2021). It has been shown that a greater number of species are needed to ensure multifunctionality than for single functions and services (Hector, Bagchi, 2007).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Later on, the impact of wildfires on each category of FESs is briefly reviewed.</span></p>
<ol style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Provisioning FESs</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Provision with wood.</em> Due to wildfires, there is a loss of wood biomass as a result of its complete or partial burning out, loss of value of wood resources due to trunk damage by fire and due to subsequent damage by wind, fungal diseases, and insects. In the case of a weak surface wildfire, when cambium is not damaged along the entire circumference of the trunk, its vital activity is partially preserved, and wood with a highly developed resin-forming apparatus begins to form, which is a response to fire damage. An increase in the number of annual rings was noted in the newly formed annual ring wood after damage. During a strong surface wildfire with a scorch height of 6–8 m, the tree loses its viability. Anatomical elements of the wood, most notably the resin canals, are completely or partially destroyed. The resin strongly impregnates the butt end of the trunk, which increases its density. Due to the destruction of anatomical elements, the sapwood of the upper part of the trunk shows a slight increase in the water absorption by wood and its decrease in the lower part due to resinosis. This affects the technology of storing lumber from fire-damaged forests (Isaenko et al., 2016). Moreover, favorable conditions are being created for the development of fungal diseases. After a severe wildfire, small and medium-sized roundwood has poor quality already in the first months after the wildfire and cannot be used as industrial wood (Kur’yanova et al., 2011). After wildfires, the growth of trees in the main canopy slows down, the understory and undergrowth are damaged (Gardiner et al., 2010). Moreover, this damage affects the economic aspects of the sale of biomass. For example, due to increased costs for timber harvesting and reforestation after a damage, the market is demoralized as a result of supply impulses (Prestemon, Holmes, 2004). After a wildfire, species composition of the forest changes, locations of raw-material bases are redistributed, which directly leads to changes in raw materials supplies to markets (Kogler, Rauch, 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Provision of non-wood FESs.</em> Since wildfire creates open spaces, despite its catastrophic effects on the ecosystem, fire is used to stimulate and increase the production of non-wood forest products, such as mushrooms, asparagus, medicinal and aromatic herbs, wild berries, nuts, etc. (Skulska et al., 2014). It is assumed that a low-intensity wildfire has a positive effect on regrowth of shoots of common hazel (<em>Corylus avellana</em>), raspberry (<em>Rubus idaeus</em>), mountain ash (<em>Sorbu</em>s <em>aucuparia</em>), prickly wild rose (<em>Rosa acicularis</em>), etc. (Johnston, Woodard, 1985; Panin, Zalesov, 2018). Under the influence of fire, the yield of the California hazelnut (<em>Corylus cornuta</em> var. <em>californica</em>) twigs increases, which are used for weaving (Marks-Block et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">After low-intensity running and surface wildfires, the amount of lingonberries reaches the pre-fire level in 2–3 years and bog bilberries — in 3–5 years, after wildfires of average intensity — in 4–6 and 6–8 years, respectively, and after strong-intensity wildfires — in 10 and 15 years. The yield of berries increases in comparison with the pre-fire level by 30–60% due to improved lighting, temperature conditions and soil moisture. At the same time, subsurface and crown wildfires of high intensity lead to almost complete loss of berry plants from the ground cover of forest phytocenoses (Ostroshenko, 2012; Duchesne, Wetzel, 2004). In the areas covered by fire, European blueberry is actually eliminated from economic use for a long time (Panin, Zalesov, 2018; Duchesne, Wetzel, 2004).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The composition of fungal communities changes greatly under the influence of wildfire, which reflects changes in physical, chemical and biochemical properties of soils (Dahlberg et al., 2001). Wildfire intensity, stand age, soil pH, humidity, and C : N ratio are considered to be the main drivers of these changes (Waldrop, Harden, 2008; Reazin et al., 2016; Day et al., 2019). Moreover, the loss of vegetation cover and changes in plant composition are closely related to fungal communities that have symbiotic/saprophytic relationships with them (Cairney, Bastias, 2007). It is reported that in some cases, after wildfires, the number of carbotrophs increases, which is a special group of fungi using ash and charred wood as a substrate, as well as saprotrophs — fungi that feed on dead organic matter, and xylotrophs — fungi that feed on the wood of living and dead trees. Some morel species (saprotrophs) bear fruit abundantly in the first year after a fire (Larson et al., 2016). Most of the marketable yield in western North America consists of morels harvested in the first year after wildfires (Pilz et al., 2007). However, these effects are short-term and not always marked. Most often, after wildfires, there is a significant reduction in the number and biomass of edible and edible mycorrhizal fungal species (Gassibe et al., 2014). Fungal communities of boreal forests are the most vulnerable. One year after the wildfire, mycorrhizal fruit bodies were not found in these forests (Franco-Manchón et al., 2019). The number of species associated with mature trees is also decreasing. Restoration of symbiotic fungi is directly related to tree restoration. Boletus and saffron milk caps appear a few years after the wildfire at sites of self-sown pines (Smith et al., 2021). Fruit bodies of xylotrophic fungi, collected also in places covered by wildfires, are used in medicine. For example, a number of polypores are used in medicine, such as sulphur polypore, <em>Ganoderma applanatum</em>, <em>Ganoderma lucidum</em>, medicinal polypore, and chaga (Kochunova, 2014).</span></p>
<ol style="text-align: justify;" start="2">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Influence on regulatory FESs</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Regulation of carbon cycles.</em> Wildfires lead to emissions into the atmosphere of large amounts of greenhouse gases and gases with an indirect greenhouse effect either directly as a result of burning out of living and dead wood, litter, as well as during the subsequent decomposition of dead wood, mineralization of litter and soil organic matter. Therefore, wildfires play an important role in the carbon cycle. It is wildfires, according to D. G. Zamolodchikov et al. (2013), that are the main cause of year-to-year variations in the carbon balance of forests in Russia. The negative impact of wildfires on carbon deposition is more often reported in the literature, mainly due to the reduction of aboveground biomass in the ecosystem (Bond-Lamberty et al., 2007; Bartalev et al., 2015; Zamolodchikov et al., 2017; Ershov, Sochilova, 2020), less often due to the burning of soil organic matter (Walker et al., 2018, 2019). It has been found that the time since the damage and wildfire intensity have an impact on the stocks of all carbon pools. So, on average, the differences in carbon stocks as compared to forests undisturbed by fire are –91.3 and +155.5% in the first year after the fire for live and dead wood, respectively, and increase by 0.6% for live and decrease by 1.4% for dead wood every year after the damage (Thom, Seidl, 2016). The study of the relationship between phytomass consumed by fire and mortality rate of trees in stands of mixed conifers and western yellow-pine (<em>Pinus ponderosa</em>) showed that burning of up to 13% of the available ground biomass led to mortality rate of 22%, while burning of 13%–35% was associated with mortality rate of 54% and of over 35% — with mortality rate of 98% (Meigs et al., 2009). Over time, forests restore biomass and, accordingly, the carbon stock that has been lost during the fire. This process depends on fire intensity and the resulting environmental conditions (soil-related, hydrological, light-loving vegetation overgrowth, etc.). For example, after a small surface wildfire, the Sierran mixed coniferous forest restores lost carbon in less than seven years, which is comparable to the historical interval between fires in such forests (Hurteau, North, 2009); Yellowstone National Park pine forests recovered about 90% of carbon within 100 years after the fire, with a historical average fire interval of 150–300 years (Kashian et al., 2013). This occurs not only due to active growth of woody plants, but also due to a decrease in soil respiration (Perez-Quezada et al., 2021), due to changes in the structural and functional organization of soil microbiocenosis against the background of pyrogenesis (Medvedeva et al., 2020). Wildfires reduce the rate of carbon mobilization by soil biota. Shifts in soil trophic webs caused by wildfires have a significant short-term impact on the carbon cycle in forest soil; these effects vary depending on the type of forest and its geographical location (Gongalsky et al., 2021). Thus, if the frequency of fire occurrence will not increase significantly and become less than time needed for restoration of a ripe forest, wildfires should not cause net carbon emissions into the atmosphere (Campbell et al., 2012). But it also follows that if forests do not recover after a wildfire, the frequency of fire occurrence is high and there is not enough time to restore carbon stocks or there is a constant change in forest structure, leading to low carbon stocks, there will be a net loss of carbon over time. Therefore, it is so important to take measures to develop systems of forecasting, rapid fire detection and extinguishing.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">However, it is believed that the protection of forests from wildfires increases the risk of fire. It has been shown that an effective fire detection and extinguishing system contributes to a significant accumulation of fuel in forests, which usually burns down during wildfires of low and moderate intensity. In combination with climate change, this can lead to a sharp increase in the frequency of fire occurrence. With such a system, in the case of large mega-fires, emission may exceed carbon deposition. Thus, in a number of countries, prescribed wildfires are used as a method of reducing the amount of fuels in such forests to reduce the risk of large catastrophic fires (Adams, 2013). At the same time, it is obvious that the trade-off with risks for environmental assets, such as biodiversity and ecosystem services, when using such a system is not entirely clear (Moritz et al., 2014; Harper et al., 2018). Prescribed burning leads to even greater frequency of fire occurrence (Yaroshenko, 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Regulation of water regime.</em> In many parts of the world, forests provide people with fresh water for domestic, agricultural, industrial and environmental needs. Forest stands affect the quantity and quality of water runoff by absorbing cations and anions from the solution, improving the bacteriological properties of water, purifying it from suspended solids and having an impact on the temperature regime of water bodies. Forest reduces peak loads of surface runoff, transforming it into underground one, and thereby reducing the risk of flooding (Rybalova, 2007). Wildfires can have devastating consequences for aquatic ecosystems and the potable water supply of the population. They can influence hydrological processes (interception, infiltration and evapotranspiration), which in turn affect the time and magnitude of river flow (base flow, peak flow and annual water production) (Shakesby, Doerr, 2006). The destruction of forest vegetation by wildfire reduces evaporation by intercepting precipitation and evapotranspiration, thereby increasing the amount of rain and snow reaching the ground and increasing soil moisture, runoff and volumes of water flowing into water bodies (Neary et al., 2003). Due to the greater amount of solar energy reaching the snow cover in the burned areas, there is a twofold increase in the rate of snow melting (Burles, Boon, 2011). Moreover, the thickness of snow cover in the areas covered by fire is less than in the undamaged areas (Maxwell et al., 2019). When the ground cover is damaged by wildfire, the natural water-repellent soil layer can be exposed (Doerr et al., 2009), which can reduce the infiltration of precipitation into the soil during heavy rains or snowmelt, contributing to an increase in surface runoff (Huffman et al., 2001). A two- to five-fold increase in peak runoff over 6–7 years is reported as a result of fire influence (Moody, Martin, 2001a). There is evidence that a combination of medium- and high-intensity wildfires in the context of intense short-term precipitation can increase peak runoff values up to 870 times (Neary et al., 2003; Moody, Martin, 2001b).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">After wildfires, the role of forest canopy in the processes of precipitation interception decreases sharply, and the qualitative composition of the runoff changes. The consequence of this is an increase in the intensity of water, wind and soil erosion. As a result, the amount of dissolved substances, phosphorus, nitrogen, dissolved organic carbon, sulfates, chlorides, calcium, magnesium, sodium and potassium that are removed from the forest catchment increases sharply, which leads to an increase in their content in surface waters (Mikkelson et al., 2013, Smith et al., 2011; Emelko et al., 2011). As a result, the concentration of pollutants, including heavy metals and pathogenic microorganisms, may increase (Stone, Droppo, 1994), as well as the amount of sediment and debris in reservoirs, which leads to silting (Smith et al., 2011). For example, after the Hayman Fire in Colorado in 2002, twice as many nitrates were recorded in river water, and turbidity increased fourfold as compared to basins whose areas burned to a lesser extent; these indicators remained elevated for 5 years after the fire (Rhoades et al., 2011). This, in turn, affects the biological population of reservoirs, including valuable commercial fishery species. In Australia, populations of fish decreased by 95–100% due to an increase in bottom sediments after the fire and a subsequent decrease in dissolved oxygen levels in river water (Lyon, Connor, 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">From the perspective of water supply, wildfires increase the likelihood of impairment of water quality (taste, smell, color, chemical composition), deterioration of potable water purification processes and shortening of the working lifespan of the water intake and treatment system (Emelko et al., 2011). This is very important because, for example, it is known that almost two-thirds of municipalities in the United States and about one-third of the largest cities in the world, including Tokyo, Melbourne, Los Angeles and Rio de Janeiro, receive most of their potable water from forest catch basins (National Research Council, 2008). As a result of a heavy post-fire downpour in south-eastern Australia, for example, the concentration of arsenic, iron, lead and chromium in drinking water increased to levels exceeding the recommendations of the World Health Organization (Leak et al., 2003). Similarly, during the first two years after the Lost Creek Fire, total mercury concentrations in potable water during storms were many times higher than permissible sanitation standards (Emelko et al., 2011). Elevated concentrations of mercury were also found in fish (Garcia, Carignan, 2005).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Rapid reforestation can offset the negative effects of wildfires on aquatic ecosystems. In the first decade after large wildfires, as compared to mature intact forests, water consumption by forest stands more than doubles during their restoration, followed by a decrease for many decades (Lane, Feikema, 2010; Buckley et al., 2011; Benyon et al., 2007). This can be down not only to an increase in the area of foliage in total (“Kuczera effect”, Kuczera, 1987), but also to the fact that, firstly, the stomatal conductance of newly developing and young leaves is much higher than that of the leaves of adult trees; secondly, both the sapwood area and the leaf area are significantly larger in young stands; and thirdly, night transpiration in young trees is also higher than in mature stands (Buckley et al., 2011).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Protection from avalanches, mudslides.</em> An important regulatory function of forests, also related to water, is the protection of society and infrastructure from natural hazards, such as floods and avalanches. Disturbances weaken the buffer effect of forests on water runoff and increase the risk of avalanches and their collapse (Zurbriggen et al., 2014). Accelerated erosion combined with the emergence of hydrophobic soils, decreased rate of water infiltration, surface runoff or massive soil disturbance on hillsides can also sometimes lead to catastrophic mud streams (Doerr et al., 2009). It is estimated that the volume of sediments from mudslides after wildfires is 2–3 orders of magnitude higher than the annual rates of background erosion from areas of undisturbed forests. The volume of mudslides from slopes with a steepness of 18–62 percent varies from 539 to 33.040 cubic meters (Nyman et al., 2015). There are models for predicting mudslides that help make management decisions, such as RUSLE (Ying et al., 2021) or the US Geological Survey (USGS) Post-Fire Hazard Model (Ellett et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Air quality regulation.</em> Since the late 1970s, wildfires have been recognized as an important source of air pollution (<a style="color: #000000;" href="https://www.sciencedirect.com/science/article/pii/S1352231011004821#bib15">Crutzen et al., 1979</a>; Rogers et al., 2020), and in the context of a changing climate, this contribution could soar due to increasing areas of wildfires (<a style="color: #000000;" href="https://www.sciencedirect.com/science/article/pii/S1352231011004821#bib1">Amiro et al., 2001</a>b; Carvalho et al., 2011). It is known that when burning biomass, many different particles and gases are formed that affect atmospheric processes. These include carbon dioxide, carbon monoxide, methane, volatile and semi-volatile organic compounds (toluene, benzene, acetone, methanol, acetonitrile, isoprene, methyl vinyl ketone, etc.), nitrogen and sulfur compounds, halogenated hydrocarbon, solid volatile particles (soot, black carbon, etc.) (Yadav, Devi, 2018; Butt et al., 2020). The impact of these emissions can be seen at different levels: from temporary local atmospheric pollution (Miranda, 2004; Hodzic et al., 2007) to the global contribution to the greenhouse effect (<a style="color: #000000;" href="https://www.sciencedirect.com/science/article/pii/S1352231011004821#bib47">Simmonds et al., 2005</a>). Emissions of CO, CH<sub>4</sub> and volatile organic compounds into the air affect the oxidizing ability of the troposphere by reacting with OH· and NO· radicals, which leads to the formation of ozone and other photo-oxidants. CH<sub>3</sub>Br emission causes ozone photodegradation in the stratosphere. Solid particles in the air can cause acidification of clouds, a change in the radiation balance of the Earth due to absorption and scattering of incoming solar radiation or formation of cloud condensation nuclei. This leads to a decrease in the size of cloud droplets, thereby increasing the albedo of clouds, which ultimately affects the nature of precipitation and the hydrological cycle (Yadav, Devi, 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Smoke with dangerous fine solid particles and gaseous compounds resulting from biomass burning is one of the main atmospheric components affecting air quality in vast territories due to its massive plumes that can travel thousands of kilometers with the wind (Chen et al., 2017; Beig et al., 2020).</span></p>
<ol style="text-align: justify;" start="3">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Cultural services</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Recreation and meeting of spiritual needs.</em> Recreational value of forest landscapes can be greatly reduced due to wildfires (Sheppard, Picard, 2006), because dead trees are often perceived as less picturesque than living stands and pose a danger to tourists. Therefore, recreational areas such as camping sites and trails are often closed after serious damage due to the risk of trees falling. On the other hand, wildfires provide researchers with opportunities to study a variety of issues, thereby contributing to the production of scientific knowledge. Moreover, many indigenous and traditional societies have a long experience of living with fire (i. e. cultural knowledge) and therefore can share it (Fowler, Welch, 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>The impact on people’s health.</em> The annual global mortality rate from the smoke of plant fires is estimated at about 339 thousand deaths per year (Cascio, 2018). Systematic reviews show that there is a positive association between exposure to wildfire smoke and mortality from respiratory diseases (Arriagada et al., 2019; Reid, Maestas, 2019; Xu et al., 2020). In a number of cases, an association has been recorded with the frequency of cardiovascular diseases, premature birth (Reid et al., 2016; Black et al., 2017), increased incidence of influenza (Landguth et al., 2020), the frequency of visits of patients with diabetes mellitus (Yao et al., 2020). In the areas surrounding a wildfire, cases of carbon monoxide poisoning are recorded very often (Tao et al., 2020; dos Santos et al., 2018). Heavy smoke can cause eye irritation and corneal damage (Finlay et al., 2012). Residents of affected areas are at greater risk of mental illness, including post-traumatic stress disorder, depression and insomnia (Belleville et al., 2019). The psychological effects of wildfires can persist for years (Bryant et al., 2018), and children and adolescents are particularly vulnerable (Brown et al., 2019). Experienced wildfires in childhood are associated with an increased likelihood of mental illness in adulthood (McFarlane, Van Hooff, 2009). Moreover, wildfires are associated with a subsequent decrease in the academic performance of children (Gibbs et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is estimated that in the United States in 2008–2012, health care costs resulting from short-term exposure to particulate smoke from wildfires ranged from 11 to 20 billion US dollars per year, while the costs associated with long-term exposure to this factor range from 76 to 130 billion US dollars per year (US dollars in 2010) (Fann et al., 2017). In Tanzania, in 2010–2019, the total cost of health care related to the effects of wildfires amounted to 76 Australian dollars per day, which corresponds to 5.2% of annual health costs associated with smoking (Borchers-Arriagada et al., 2020).</span></p>
<ol style="text-align: justify;" start="4">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Supporting services</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Net primary production </em>(<em>NPP</em>)<em>.</em> After disturbances, NPP remains low for several years, partly due to the low leaf area index and their number; it reaches a maximum when the canopy closes and decreases slightly as the stand matures (Odum, 2014; Gower et al., 1996; Ryan et al., 1997; Howard et al., 2004; Goulden et al., 2011). In addition, repeated disturbances associated with stand replacement can prevent forests from reaching maximum NPP values (Gough et al., 2007), causing nitrogen losses due to leaching or a decrease in the amount of organic matter and soil fertility in general (Latty et al., 2004). The impact of fire frequency on NPP is particularly pronounced for coniferous forests which have a longer leaf lifespan and a longer recovery period (Peters et al., 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Soil formation </em>(See also the section “Influence of fires on the morphological and physico-chemical properties of soils”). During wildfires, there is a change in soil-forming processes (pyrogenesis of soils). Short-term and long-term post-fire changes are identified. During a wildfire, under the influence of high temperatures, the surface layers of soils lose organic matter, and roots, invertebrates, microorganisms, etc. die. Soil fertility depletion is observed. The contribution of organnic horizons to the total stock of soil carbon is reduced. In the soils of wildfire sites, aeration improves and oxidative processes, ammonification and nitrification are intensified, the degree of decomposition of litter fall within the soil and loss of total carbon increases. In the surface mineral horizons, the pH and base saturation increases as well as the content of mobile organic and mineral compounds increases. Wildfire changes the composition of carbon forms, increasing the proportion of hydrophobic compounds, which affects the structure of the soil system, and the biochemical composition and population of microorganisms in particular (Nadporozhskaya et al., 2020). The strongest impact on the soil has not the fire itself, but post-fire secondary changes in the biogeocenosis associated with the post-fire transformation of vegetation cover (Sapozhnikov et al., 2001). However, it is difficult to make prognoses of composition of vegetation after a fire, because it is influenced by many factors, such as the degree and area of the fire, the distribution of surviving trees, the volume of the seed bank, landscape fragmentation, climate change, invasion of species, the number of herbivores, changing accessibility of the territory, subsequent disturbances (McLauchlan et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Pollination</em>. Since wildfires form open spaces, where populations of flowering plants are usually more represented than under the forest canopy, the density of pollinating insects is higher (Campbell et al., 2007; Hanula et al., 2015). Therefore, it has become more and more accepted that landscape mosaic with a variety of fire regimes and stand ages after wildfires contributes to the diversity of flowering plants and pollinators (Ponisio et al., 2016; Brown et al., 2017; Lazarina et al., 2019), which can also increase crop yields (Winfree et al., 2018; Mola, Williams, 2018). However, open spaces can be created by humans in ways that are less destructive to the ecosystem, for example, by logging, which also contributes to improving pollination efficiency (Goulson et al., 2015).</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Economic damage from loss of ecosystem services as a result of wildfires</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Despite the great economic importance of forest ecosystem services, there are few quantitative estimates in monetary terms of the impact of wildfires on forest ecosystem services (Lee et al., 2015). According to San Diego State University, the total economic impact of the 2003 wildfires in San Diego County is estimated at $2.45 billion, of which the cost of extinguishing is less than two percent of the total losses. This does not take into account the long-term impacts of wildfires on the affected catch basins (<a style="color: #000000;" href="https://www.sciencedirect.com/science/article/pii/S0921800915002050#bb0285">Rahn, 2009</a>). The Western Forestry Leadership Coalition estimates the true cost of wildfires in the western United States from two to thirty times higher than the cost of extinguishing (The true&#8230;, 2014). In our country, using the example of territories of two protected areas in the Irkutsk oblast, quantitative calculations of losses of ecosystem services of forests as a result of wildfires are given (Volchatova, 2019): for the Baikal National Park, annual total damage averages 136.26 million RUB, while for the Baikal-Lena Reserve — 1081.71 million RUB. It is emphasized that the territory of Siberia is extreme in terms of fire. For example, in Irkutsk Oblast, 77% of the forest fund is classified as the first three classes of natural fire danger. The situation is aggravated by the climatic and light conditions of the region — a sharply continental climate with a hot and arid summer period, sunshine over 2 thousand hours per year. An additional factor contributing to vulnerability of the forests of these protected areas is the predominance of pine forests in dry habitats with easily ignitable ground cover and high frequency of fire occurrence in pine stands. Damage caused by wildfire includes not only loss of standing wood, but also decreased ecological functions of the forest, pollution by combustion products, death of biota, which increases the amount of regulatory and support services of forests that were not received.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, at present, wildfires are one of the leading factors regulating the functioning of forest ecosystems. Wildfires of any intensity have an impact on forest ecosystem functions and services of all categories. The short-term increase observed in some cases in the provision of non-wood products (berries, mushrooms, medicinal herbs) and such a supporting function as pollination resulting from the mosaic pattern of forest cover created by wildfires does not make up for the loss of other provisioning (wood, fibers), supporting (net primary production, soil formation, habitat maintenance), regulatory and cultural services. The extent of economic damage caused by wildfires, especially those of high intensity, is difficult to assess, since there is no clear understanding of the long-term effects of wildfires on biodiversity and ecosystem functions and services of forests as of yet. However, it is extremely important to take into account the impact of fire consequences on the functioning of ecosystems and economic development in the context of climate change when making management decisions.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The results of the studies regarding the fire impact on forest ecosystems show devastatingly powerful and long-term destructive impact of wildfires on the biodiversity and functions of forests. According to official statistics, in the last decade, hundreds of thousands of wildfires have been detected in Russia alone, with the total area covered by fire estimated at millions of hectares. Currently, the proportion of large wildfires (those with an area of more than 200 hectares) has increased. Due to global climate change, an increase in the frequency and intensity of wildfires is expected. The most common fire type in the forests of Russia are surface wildfires that have a destructive impact on soil and soil inhabitants, which leads to impaired soil formation and, consequently, decreased efficiency of all ecosystem processes. The restoration of litter in boreal forests after fires may take more than 120 years. In the mineral horizons of soils, “traces of fires”, in the form of a change in chemical composition and depletion of elements of mineral nutrition are found over 100 years after the fires. No complete recovery of all components of the soil biota has been revealed in the first few decades after the fires, whereas results of longer observations are lacking. Vegetation restoration requires considerable time (tens and hundreds of years), if there are not enough diaspora carriers, i. e. birds and mammals, whose populations are also disrupted by wildfires and other causes. Wildfires are a factor that results in loss of genetic, taxonomic, and functional biodiversity, damage and destruction of habitats for plants, animals and microorganisms, loss of functions of forest ecosystems. Wildfires are a factor in the dynamics of forest ecosystems directed at “erasing of evolution”.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Analysis of literary sources shows that an established opinion expressed in a number of works, that wildfires are, at a certain frequency, essential for the maintenance of forest communities, ignores or misunderstands the role of biotic factors in the functioning of forests. Populations of keystone large mammal species have been lost or drastically reduced in the modern forest ecosystems; consequently, there are no microsites formed by them, including large gaps in the forest canopy (glades) that provide opportunities for maintaining light-demanding flora, insect pollinators and conditions for the development of all-aged polydominant forest ecosystems with high biological diversity in general. Moose, bison, beavers, and other animals create natural barriers to the spread of fire due to formation of gaps, trails, sparse stands, and reservoirs in the forests.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It should be emphasized that an increased number and diversity of individual groups of invertebrates and vertebrates on fire sites is short-term, limited by trophic resources rapidly petering out on fire sites and is due to new open spaces becoming available for settlement by species from neighboring biotopes with high migratory abilities. Often, as a result of large wildfires, huge homogeneous open spaces are formed, which are very far from the sources of diasporas of many plant species and are difficult to be populated by “low-mobility” groups of animals, which results in a steady decline in biodiversity. Wildfires as a powerful factor trigger positive feedback mechanisms leading to the elimination of species, which is why some forest communities have been identified by researchers as fire-dependent.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wildfires of any intensity have an impact on forest ecosystem functions and services of all categories. The short-term increase observed in some cases when providing some non-wood products (berries, mushrooms, medicinal herbs, pollination) resulting from the mosaic pattern of the forest cover created by wildfires does not make up for the loss of other functions and services of forests. The extent of economic damage is difficult to assess, since the long-term effects of wildfires on the climate, soil formation, water regimen regulation, and human health are not taken into account.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is essential to ensure continuous maintenance and restoration of populations of endangered animal species in modern forests, especially large mammals that create zoogenic clearings and gaps in the forest canopy, regulating the density of the stand and the mosaic pattern of the ground cover.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Based on the performed analysis of the impact of wildfires, we can give the following recommendations for the conservation and maintenance of biodiversity and ecosystem functions of forests in the modern forests:</span></p>
<ul style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">take action to prevent wildfires: educate people on how to prevent wildfires; completely ban burning of felling residues during the fire-hazardous season; ban agricultural and any prescribed burning of dry grass vegetation (Postanovlenie&#8230;, 2015; Sosnovchik, 2016; Volchatova, 2019; Vacchiano et al., 2018; Yaroshenko, 2021);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">take action for timely detection and rapid and prompt localizing of fires: abolition of “control zones” where it is allowed not to extinguish fires; increase the staff and funding of road and air forest protection several times; continuous road, air and space monitoring of fire danger in forests (Korovin, Isaev, 1997; Gomes et al., 2006); develop safety barriers that would prevent the spread of wildfires, including channels and water reservoirs to be used for fire extinguishing (Češljar, Stevović, 2015);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">harvest large wood residues in areas of massive blow-downs, provided that the deadfall of individual tree trunks is preserved to maintain the biological diversity of xylobionts (Lust et al., 2001);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">maintain and restore populations of endangered animal species in modern forests, especially large mammals that create zoogenic clearings and gaps in the forest canopy, regulating the density of the stand and mosaic pattern of the ground cover (Van Meerbeek et al., 2019; Van Klink et al., 2020), as well as beavers as the main representatives of “forest firefighters” regulating the groundwater level, creating intra-forest reservoirs that serve as natural barriers to the spread of fire (Evstigneev, Belyakov, 1997; Aleynikov, 2010; Zav’yalov et al., 2016). That is, it is necessary to restore the biotic factor that forms the structural diversity in forest ecosystems (Lukina et al., 2021);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">ensure haymaking and grazing of domestic animals near human settlements. These impacts would, on one hand, prevent the formation of communities with large reserves of dry grass and rags, which create a high fire hazard, and, on the other hand, support biological diversity and productivity of ecosystems (Smirnova et al., 2021; Evstigneev, Gornov, 2021).</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">form mixed stands as more fire stable during forest restoration after wildfires and during plantation management (Korotkov, 2016, 2017; Gomes et al., 2006);</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">if necessary, conduct gap felling with planting or sowing of light-loving tree species in the gaps (Metodicheskie&#8230;, 1989; Korotkov, 2016, 2017).</span></li>
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">fell individual trees and groups of trees to prevent the spread of fire (Allen et al., 2002).</span></li>
</ul>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>ACKNOWLEDGEMENTS</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The study was conducted within the framework of the state CEPF RAS assignment 121121600118-8. The authors express their deep appreciation and gratitude to A. V. Gornov for many valuable comments and additions that have served to improve this article.</span></p>
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<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewers:</strong> Doctor of Biological Sciences K. B. Gongalsky, Candidate of Biological Sciences V. N. Korotkov.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><em><sup> </sup></em></strong></span></p>
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		<title>APPROACHES TO ECOLOGICAL CLASSIFICATION  OF EARTHWORMS: A REVIEW</title>
		<link>https://jfsi.ru/en/5-1-2022-ermolov/</link>
		
		<dc:creator><![CDATA[lena]]></dc:creator>
		<pubDate>Mon, 16 May 2022 08:46:48 +0000</pubDate>
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					<description><![CDATA[Original Russian Text © 2021  S. A. Ermolov published in Forest Science Issues Vol. 4, No. 4,   Article 93  S. A. Ermolov Center for Forest Ecology and Productivity of the Russian Academy of Sciences, Profsoyuznaya st.&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/05/5-1-2022-Ermolov.pdf"><img loading="lazy" class="size-full wp-image-1122 alignright" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></span></p>
<div style="text-align: justify;"><span style="font-size: 10pt; font-family: 'times new roman', times, serif; color: #000000;"><span lang="EN-US">Original Russian Text © 2021  S. A. Ermolov published in Forest Science Issues Vol. 4, No. 4,   </span><a style="color: #000000;" href="https://jfsi.ru/4-4-2021-ermolov/" target="_blank" rel="noopener noreferrer">Article 93</a></span></div>
<div>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> S. </strong><strong>A. Ermolov</strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Center for Forest Ecology and Productivity of the Russian Academy of Sciences, </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Profsoyuznaya st. 84/32 bldg. 14, 117997, Moscow, Russia</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">E-mail: ermserg96@gmail.com</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 07 July 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 01 December 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 07 December 2021</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Earthworms as critical components of natural communities have traditionally attracted the attention of researchers from various fields of biology and agriculture. From the observations of ancient times and up to our time, the research of earthworms has kept its relevance. One of the most interesting tasks has been the creation of an ecological classification of earthworms and its subsequent use. The purpose of this review is to consider various approaches to identifying ecological groups of earthworms (Oligochaeta, Lumbricidae) and the application of these approaches in scientific research. The article reviews the main types of Russian and world ecological classifications of earthworms and some recent additions to these classifications. Particular attention is paid to scientific research with different approaches to the study of ecological groups of earthworms.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Keywords:</strong><em> earthworms, life forms, ecological groups, soil ecology, classification </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In most terrestrial ecosystems, earthworms are one of the main soil-forming organisms. They ensure transformation of soil organic matter through decomposition of plant residues, forming a humus horizon and transferring organic compounds into deep soil layers, and also through consumption of humus, thus providing mineralization and migration of C and N compounds in the soil (Giljarov 1951; Kurcheva, 1971; Holdsworth et al., 2008). Earthworms also contribute to soil aeration and uniform distribution of moisture: plants can reach groundwater with their roots using worm passages, and the mucus released through their skin moistens the walls of passages, prevents cracking and subsequent drying of the soil. This helps to create favorable soil conditions not only for plants, but also for microorganisms involved in soil-forming (Gaponov, Hicova, 2005; Lemtiri et al., 2014). Earthworms’ habitat is not only the mineral horizons of the soil. They also inhabit forest litter, are abundant in forest deadwood and aggregations of animal feces, contributing to their decomposition, and can also live in swamps or at the bottom of shallow reservoirs (Chekanovskaja, 1962; Perel’, 1975). At the same time, in most habitats, for example, in river floodplains and moist forests, earthworms mainly inhabit the soil. In forest ecosystems, however, the species and functional diversity of earthworms can be provided by deadwood to a greater extent (Geraskina, 2016a, 2016b; Salomé et al., 2011; Ashwood et al., 2019; Ermolov, 2020a, 2020b).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Earthworms have become an interesting object of study for a vast array of researchers. For taxonomists, still are a challenging group with very high variability in many morphological features, as several species are characterized by inherent polymorphism, parthenogenetic and polyploid races; therefore, the study of earthworms required using cytology, genetics, and molecular biology (Briones, 1996; Shekhovcov et al., 2016, 2020, 2020a). Ecologists and zoologists consider earthworms as zoological indicators for biological diagnostics of various types of soils (Atlavinite, 1960; Paoletti, 1999; Ivask et al., 2006; Zhang et al., 2015). Many studies have clearly shown the viability of the so-called “lumbricidological method” in monitoring of soil and environmental conditions, assessment of soil moisture and acidity as well as decomposition intensity of plant residues (Boeskorov, 2004; Zhukov, 2004; Uvarov, 2019). Earthworms also have a significant impact on other components of soil biota (Tiunov, 2008). A special place in such studies is occupied by the study of complexes of living forms or ecological groups of earthworms. Representatives of each group in this complex perform certain functions, and the absence of a group indicates a change in soil properties or their disturbance (Perel’, 1975; Geraskina 2016a, 2016b). Therefore, the presence of certain earthworm species and living forms in the biotope, as well as their population density, have become a kind of indicators of soil condition (Chekanovskaja, 1960; Giljarov, 1965; Paoletti, 1999; Akkumuljacija&#8230;, 2018). The study of earthworm ecology has received significant application. For example, earthworms are used by humans to restore affected soils, in particular in abandoned quarries and dumps through introduction (Dunger, Voigtländer, 2002; Geraskina, 2016b, 2019). In agriculture, earthworms are used to increase soil fertility and produce compost, and they are able to maintain soil biological activity for a long time (Igonin, 1995; Titov, 2012; Vorob’eva, Ivanova, 2018). It should be mentioned that certain worm species are sometimes bred for agriculture taking into account their ecological and physiological features, with <em>Eisenia fetida</em> serving as an example, which cannot survive in natural biotopes in most regions and therefore inhabits only anthropogenically transformed areas (Meshherjakova, 2011; Titov, 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Currently, comprehensive studies of earthworm biology are carried out both in many regions of Russia and abroad. One of the most popular aspects is the study of earthworm ecology, in particular, their ecological classification. The purpose of this review is to consider various approaches to the identification of ecological groups of earthworms (Oligochaeta, Lumbricidae) and application of these approaches in scientific research.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>1.APPEARANCE THE SCIENTIFIC INTEREST TO EARTHWORM ECOLOGY</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Since ancient times, earthworms (Oligochaeta, Lumbricidae) have gained the attention of the first researchers of nature. For example, Aristotle highlighted the important role of earthworms in increasing soil fertility and called them “the intestines of the earth”, while in ancient China earthworms were dubbed “angels of the soil” for the same reason (Geraskina, 2016b). During the Middle Ages and Modern times, earthworms were often described in the works of philosophers and natural scientists who tried to put together a system of the animal world. Back then, earthworms had been considered to be insects with special anatomical features for a long time, until С. Linnaeus set them apart as a separate class in his system of animals (Class 6 — Worms) in 1735 (Chesnova, Striganova, 1999).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Charles Darwin was the first to give scientific evidence of the lifestyle of earthworms and their soil-forming activity in his famous book, <em>The Formation of Vegetable Mould, Through the Action of Worms, with Observations on Their Habits,</em> published in 1881. Prior to that, for 50 years, he had conducted many laboratory experiments and observations in nature, which revealed the features of the burrowing activity, behavior, physiology, food preferences of earthworms and demonstrated their ecological function as decomposers and humus-forming organisms (Charles Darwin&#8230;, 1936). It was Darwin who pointed out that earthworms, in fact, change the natural conditions of their habitat, since they transform plant litter not only mechanically, but also chemically, creating humus substances. On the other hand, soil could have passed through the intestines of earthworms many times during its existence, which proves the role and importance of earthworms in soil formation (Charles Darwin&#8230;, 1936).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A similar study was conducted by Darwin’s contemporary, the German zoologist V. Hensen, who described in detail the processes of leaf litter decomposition performed by earthworms and studied the structure of their passages. He also discovered that one worm (<em>Lumbricus terrestris</em>) releases on average 0.5 g of nitrogen-rich casts per day, ensuring an even distribution of organic substances at different depths of the soil (Chesnova, Striganova, 1999).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">These studies paved the way for further research on the role of earthworms in decomposition of plant remains and humification, including in Russia. One of the first Russian works on this topic was an essay by the soil scientist A. I. Polimpsestov (1882), who also argued, however, that, in addition to earthworms, other invertebrates like woodlice and insect larvae also play an important role in soil-forming. This point of view was later developed by other Russian scientists, and new studies of the soil-forming activity of earthworms were reflected in the works of P. E. Muller (1887), P. A. Kostychev (1889), N. A. Dimo (1938) and other researchers. Of great interest are the observations of G. N. Vysockij, who studied the intensive activity and spread of earthworms in the chernozems of southern Russia (Vysockij, 1900).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">We would like to mention the works of M. S. Giljarov (1912–1985) who organized and supervised studies of interaction between earthworms and complexes of other soil invertebrates. It was found that mechanical destruction of plant material in the soil is carried out only by animals and not by any other groups of soil organisms (Giljarov, 1951; Giljarov, Striganova, 1978). Through combining morphological and statistical data on earthworms and other invertebrates with the data on soil conditions, Giljarov together with his students created universal methods of zoological diagnostics of soils, which is still used in soil zoological research (Metody&#8230;, 1975).</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>2. FORMATION OF CLASSIFICATION OF EARTHWORMS ECOLOGICAL GROUPS AND LIVING FORMS</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Until the late 1920s, earthworms were considered an ecologically homogeneous group. Previously researchers had been more interested in the taxonomy of the described earthworm species, mainly the creation of a generic system for Lumbricidae and other families.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Briefly speaking, a compilation of a taxonomic classification of earthworms started in the late 19th — early 20th century and keeps being updated now. Initially, when distinguishing genera, signs of the external and internal anatomy of earthworms were used, among those the position of setae, shape (section) and color of the body, location of the clitellum and tuberculae pubertatis, number of seminal vesicles, position of spermathecae, and structure of muscle fibers. At different times, there had been several generic systems based on combinations of those features, which gradually replaced each other. We should mention such authors as G. Eisen, D. Rosa, V. Michaelsen, V. Pop, P. Omodeo, and M. Bouche in this regard (Perel’, Semenova, 1968; Perel’, 1979). Russian researchers who studied earthworm morphology and taxonomy during this period also include the works of P. G. Svetlov and I. I. Malevich. At that time, almost all the proposed classifications had a common principle: genera were identified based on the structure of the reproductive system, whereas species were identified based on external morphological features (Malevich, 1950; Chekanovskaja, 1960). Upon further revision, the generic systems of those authors were abolished. A very good option for the generic earthworm system was created in the 1970s. The American researcher G. Gates was the first to use the shape of nephridial bladders as a taxonomic feature, and the outstanding Soviet and Russian lumbricologist and taxonomist T. S. Vsevolodova-Perel’ supplemented his classification with data on the orientation of nephridial bladders relative to the head end of the worm and the change in their shape in different body segments in some species (Perel’, 1979; Vsevolodova-Perel’, 1997). We should also mention the significant contribution to the taxonomy of earthworms made by the Hungarian zoologist C. Csuzdi. On multiple occasions, he revised the taxonomy on the basis of morpho-anatomical features (which, for example, resulted in identification of an endemic monotypic genus <em>Rhiphaeodrilus</em> separated from the genus <em>Perelia</em>) and used data from molecular biology studies in taxonomy (as a result, the genus <em>Dendrodrilus</em> was included in the genus<em> Bimastos</em>) (Csuzdi, Pavlíček, 2005; Csuzdi et al., 2017). Despite the universal character of the proposed generic system, the taxonomy of earthworms continues to change at the present time as well.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Over time, collected data on the peculiarities of earthworms’ lifestyle and the heterogeneity of their role in soil-forming processes led to the conclusion that there are several ecological groups of earthworms.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The first, not very rewarding attempts to define ecological groups of earthworm species were made by V. K. Baluev and D. Wilcke, who mainly took into account the vertical distribution of earthworms in the soil, as well as their pigmentation and ability to diapause (Baluev, 1950; Wilcke, 1953). Further studies have shown that earthworms also differ in their feeding habits: there are the so-called “humus-forming” earthworms feeding on poorly decomposed plant material on the soil surface, and “humus-consuming” ones, which feed on soil humus (Franz, 1950, cit. by: Perel’, 1975).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The most complete morpho-ecological classification of the Lumbricidae family, which is still used in the world literature, was proposed by M. Bouche in 1972. He identified three groups of earthworms according to their ecological strategies: <em>epigeiс</em> earthworms that live in litter and feed on it; large <em>anecic </em>earthworms that go deep into the soil using vertical passages but feed on litter on the surface; and <em>endogeic</em> earthworms that live directly in the soil and feed on humus in the humus horizon (Bouche, 1972; Fründ et al., 2010; Fierer, 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the Russian literature, the morpho-ecological classification of earthworms of the Lumbricidae family is used that was developed by T. S. Perel’ in 1975 after extensive laboratory research and field observations. This classification is based on the comparison of anatomical, morphological and physiological features (thyphlosole structure, shape of prostomium, body cross-section, etc.) with some environmental features. Besides, the morpho-ecological group of earthworms formed by species from different genera that live in the same environment and have signs of deep convergence, was designated as a <em>living form</em> in that classification (Perel’, 1975). All representatives of the family were divided into two large morpho-ecological types depending on their feeding habits (on the surface or in the humus horizon), and each of those types included several morpho-ecological groups, distinguished according to their vertical distribution in the soil (epigeiс, epi-endogeic earthworms and endogeic earthworms of different soil layers), as well as subgroups of amphibiotic forms, including species whose life cycle is associated with the aquatic environment (Perel’, 1975, 1979).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">As compared to foreign options, the morpho-ecological classification by T. S. Perel’ is more detailed. For the first time, the group of epi-endogeic earthworms was identified, which had previously been combined with epigeiс earthworms. Unlike the latter, the epi-endogeic earthworms, although they feed on the surface, live mainly in the upper layers of the soil, rarely going to a depth of more than 15–20 cm. According to their ecological function, epi-endogeic forms of earthworms are sometimes considered equivalent to anecic earthworms, but they differ in the depths of habitat in the soil and the degree of adaptation to different humidity regimes: epi-endogeic species are more moisture-loving and found even in swampy soils, whereas anecic earthworms are better adapted to endure periodic drought (Perel’, 1979; Lemtiri et al., 2014; Akkumuljacija&#8230;, 2018). Also, this classification found its application in assessing the zonality of earthworms (tundra and northern taiga are inhabited only by epigeic and epi-endogeic earthworms, while steppes are inhabited by endogeic earthworms, and mixed and broad-leaved forests are inhabited by almost all earthworms living forms) and made it possible to identify the main directions of evolution of the Lumbricidae family (Perel’, 1975). In 2016, A. P. Geraskina introduced the concept of a <em>full-fledged complex of earthworms living forms</em>, implying the presence of all earthworms living forms in a particular biotope (Geraskina, 2016b).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Over time, world classifications attempted to “split” large ecological groups of earthworms into highly specialized ones. In 1977, based on his own observations and experiments, M. Bouche identified intermediate ecological groups of earthworms: <em>epi-endogeic</em>, <em>epi-anecic</em>, <em>endo-anecic</em>, and <em>intermediate</em> (Bouche, 1977). In the late 1990s and early 2000s, an idea started up to identify “subcategories” in the three main ecological groups; for example, <em>polyhumic</em>, <em>mesohumic</em>, <em>oligohumic</em>, and <em>endo-anecic </em>subcategories were identified in the endogeic group (Barois et al., 1999; Chan, 2001). In this case, the allocation of subcategories is based on the ability of earthworms to inhabit the soil horizon that is, to some extent, enriched with organic matter which they feed on. In 2020, all variants of the Bouche’s classification were revised using an original method (Bottinelli et al., 2020): first, a review of publications was conducted, which mentioned earthworms ecological groups proposed by Bouche, both basic and intermediate. It turned out that different researchers could classify one and the same species of earthworms as belonging to different ecological groups: for example, <em>Lumbricus terrestris</em> was described both as <em>anecic</em> and <em>epi-anecic</em>, while <em>Lumbricus rubellus</em> was described as <em>epigeic</em>, <em>epi-endogeic</em> and even <em>epi-anecic</em>. Subsequently, using mathematical modeling, a scheme based on 13 morpho-anatomical features was drawn up, which distributed earthworms into ecological groups. A new classification version of earthworms ecological groups followed: for example, <em>Octolasion lacteum</em> (commonly considered to be <em>endogeic</em>) was assigned to the <em>endo-anecic </em>group<em>, Lumbricus terrestris</em> (<em>anecic</em>) was described as <em>epi-anecic</em>, and <em>Allolobophora chlorotica</em> was described as <em>epi-endo-anecic</em> (Bottinelli et al., 2020). That study has, once again, shown that the question of ecological groups and earthworms living forms is yet open to be discussed.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>3. APPLICATION OF ECOLOGICAL CLASSIFICATION OF EARTHWORMS IN RESEARCH</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>3.1 Earthworms living forms in ecological research </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">After taxonomic and morpho-ecological classifications of earthworms had been created, new approaches to their study appeared. Early environmental studies mainly described the influence of earthworms on soil properties. For example, G. F. Kurcheva experimented with the rate of plant litter neutralization by earthworms at control sites (Kurcheva, 1971); P. U. Bahtin and M. N. Pol’skij investigated the activity of earthworms in sod-podzolic soils (Bahtin, Pol’skij, 1950); K. I. Gavrilov studied the role of earthworms in enriching the soil with biologically active substances (Gavrilov, 1963). At the same time, the participation of certain species and living forms of earthworms in these processes was not evaluated separately.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In further studies, data on the proportions of earthworms living forms started to be used to characterize ecological conditions of biotopes. One of the followers of T. S. Vsevolodova-Perel’, I. B. Rapoport, shows in her works focusing on the landscape distribution of earthworms of the Caucasus how the diversity of earthworms living forms in different biotopes varies depending on the altitudinal zonality (Rapoport, 2010, 2015). She also provides a comparison of the chorological and morpho-ecological groups of earthworms (Rapoport, 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Studies conducted in the Komi Republic in the 1970s–2000s have shown how the species composition and complexes of earthworms living forms change with changing zones from the southern to extreme northern taiga. Approximately nine species of three living forms were found in the southern and middle taiga subzones, while only two species representing one living form live in the subzone of the extreme northern taiga (Krylova et al., 2011). In the taiga zone, M. Ja. Vojtehov investigated the soil-forming activity of earthworms through a series of experiments with food preferences. It was found that, in acidic soils and litter formed by taiga vegetation, different earthworms living forms support each other’s existence: for example, endogeic earthworms can consume decomposition products of coniferous plant litter only when they are enriched with casts of epi-endogeic earthworms (Vojtehov, 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Despite the mutually beneficial existence, a number of experiments have shown that competition between earthworms of different species is possible within the same living form. For example, when epi-endogeic species were kept in mesocosms, competition for food resources was observed between <em>L. rubellus</em> and <em>Eisenia nordenskioldi nordenskioldi</em> (Golovanova et al., 2018). In endogeic species, competition occurs mainly in limited spaces with high population density. It has been shown that once a certain population density level is reached, <em>Al. chlorotica</em> and <em>Aporrectodea caliginosa</em> stop reproducing and lose weight (Uvarov, 2019). However, this is true only for experiments under artificial conditions, and the probability of actual competition among earthworms in the natural environment may be extremely low.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Earthworms can provide for the existence of other representatives of the soil fauna as well. In a number of experiments, it has been shown that casts of anecic and endogeic earthworms are an available food source for Enchytraeidae, which release C and N compounds and ensure their transport within the soil. Also, they are able to significantly affect soil fertility and increase microbial biomass in the soil only in combination with earthworms (Sandor, Schrader, 2012). The gastrointestinal tract of an earthworm with its complex chemical and microbiological processes is actually similar to a bioreactor (Brown et al., 2000). In each part of it, specific stages of the nitrogen and carbon cycle take place. Some soil bacteria, protozoa and fungal spores are digested, while others pass through earthworm’s intestines undamaged and are dispersed in the soil, and some others are activated only after passing through earthworm’s intestines, eventually reaching favorable conditions for further development (Moody et al., 1995; Lemtiri et al., 2014).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Along with the transfer of microorganisms and decomposition of organic residues, earthworms can accumulate various chemical elements when feeding, in particular heavy metals (Usmani, Kumar, 2015). This made it possible to use earthworms as bioindicator organisms to assess soil pollution. For example, the epi-endogeic earthworm <em>E. nordenskioldi</em> and the endogeic <em>O. lacteum</em> proved to be suitable indicators in the study of the content of heavy metals, especially Pb, in soils near roadsides, clearly showing the detrimental effect heavy metals have on soil biota (Golovanova, 2003). Similar work was carried out to assess the impact of emissions from iron and steel plants: the Ural epi-endogeic endemic <em>Rhiphaeodrilus diplotetratheca</em> (formerly <em>Perelia diplotetratheca</em>) showed significant differences in the size and weight of earthworms depending on the degree of soil pollution (Reznichenko, 2017). It was also found that the anecic earthworm <em>L. terrestris</em> can not only accumulate compounds of As, Cu, Pb, and Zn in its body but also include them in its casts ejected on the soil surface, thereby ensuring the removal of heavy metals from the soil (Sizmur et al., 2011). Some researchers plan to use <em>L. terrestris</em> and <em>E. fetida</em> for detecting and eliminating oil pollution due to the ability of earthworms to accumulate and remove various pollutants (Hanna, Weaver, 2002).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Isotope analysis is one of the modern methods used in the study of the ecological functions of animals. This method is often used when studying trophic relationships of various invertebrates and enables the identification of their feeding features and them being a part of a certain ecological group (Tiunov, 2007; Goncharov, 2016). Using isotope analysis in studying the ecology of earthworms made it possible to assess the features of their feeding and food preferences. For example, when a pasture is converted into a cornfield, the same species of epigeiс and epi-endogeic earthworms prefer to use “fresh” organic residues of C4 plants rather than “old” organic matter of the soil formed mainly by C3 plants, which they fed on earlier (Briones et al., 1999). Isotope analysis helps us understand the trophic features of earthworms during the decomposition of organic matter: the content of accumulated nitrogen in the tissues of earthworms proved that, when feeding, epigeic and anecic earthworms prefer to use organic material less susceptible to microbial decomposition than endogeic earthworms. Adding crushed oat flakes with isotopic labels to feed substrates showed that the endogeic earthworms <em>A. caliginosa</em> are more inclined to absorb small food particles in large quantities, unlike the anecic <em>L. terrestris</em> (Heiner et al., 2011). Isotope analysis methods are also applicable to the study of the chemical composition of earthworm casts, which are complex stable sets of organomineral matter and microbial communities. Using the example of anecic <em>L. terrestris</em>, such studies help us trace the “path of the casts”: what kind of consumed litter they consist of, which microbial communities developed there over time, and how further consumption of casts by plants or other soil animals occurs (Vidal et al., 2019). It is believed that isotope analysis can enable further revision of the ecological classification of earthworms (Briones et al., 1999).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A large number of studies showed that in some biotopes the earthworms species diversity is represented by a certain living form. For example, northern dark coniferous forests had been considered a virtually unsuitable habitat for earthworms and extremely poor in terms of their species for a long time (Perel’, 1958, 1979). Later it was found that in the dark coniferous forests (especially green moss and blueberry-green moss) of the middle and northern taiga, most of the earthworm population may be found in deadwood, not soil (Geraskina, 2016c, 2016d). The main inhabitants of deadwood are epigeiс and epi-endogeic earthworms with a relatively high species diversity; at the same time, deadwood is sometimes also inhabited by endogeic earthworms that use it as a temporary habitat during unfavorable conditions. Similar conclusions were made for other types of forest. Initially, these studies were limited only to the sorting of deadwood accidentally discovered in forest habitats. Later, calculations of the worm population density per unit volume began, and deadwood has been recognized as a specific microsite inhabited by earthworms (Kooch, 2012; Geraskina, 2016d; Ermolov, 2018a, 2018b, 2020a; Vorobejchik et al., 2020). In habitats with disturbed or heavily polluted soil, deadwood often becomes the only habitat for earthworms (Vorobejchik et al., 2018, 2020). In 2019, a new method of site sampling for earthworms in forest communities was developed, which made it possible to give the most accurate assessment when comparing the population of earthworms in soil and deadwood (Ashwood et al., 2019). Conversely, in anthropogenic habitats, in particular agricultural land and fallows, the major part of earthworms population consists of endogeic earthworms, especially middle-soil-layer ones, which can make up to 100% of the entire population (Geraskina, 2009; Shashkov et al., 2016). As the overgrowth progresses, abandoned fields are gradually populated first by epi-endogeic, and then by epigeiс earthworms, while endogeic earthworms, for example, <em>A. caliginosa</em>, live even in fields actively used in agriculture (Geraskina, 2009, 2016a). Therefore, the introduction of earthworms to various anthropogenic areas where earthworms are completely absent is started with representatives of this living form because they are able to survive and show ecological plasticity in relation to various environmental factors (Ansari, Ismail, 2012; Geraskina, 2019).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>3.2. Polymorphism and molecular-biological studies of earthworms living forms </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It’s worth noting that representatives of various earthworms living forms can be found not only within the same genus, but also within a species or subspecies. Currently, the study of polymorphism in earthworms started to use methods of molecular biology, one of them being variability analysis of the gene of cytochrome c oxidase subunit 1 (<em>cox1</em>) along with morphometric analysis (Voronova et al., 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">G. N. Ganin (1959–2019) had studied the Far Eastern endemic <em>Drawida ghilarovi</em> Gates, 1969 (fam. Moniligastridae) and revealed that individuals of this species form two morpho‑ecological groups that differ in their color and ecological features. Earthworms inhabiting meadows and swamps are epi-endogeic, black-colored and have an optional diapause; forest earthworms are anecic, have a brownish color and an obligate diapause (Ganin, 2013a, 2013b; Ganin, 2014). However, upon further study of the identified forms, especially the study of their phylogeny using molecular biology methods, it was found that forest anecic earthworms <em>Drawida ghilarovi</em> consist of ten separate genetic lineages, which hypothetically can be different species, whereas the black epi-endogeic meadow and swamp morph represents a new species (Zhang et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It has been repeatedly suggested that there are two living forms in the Asian subspecies <em>E. n. nordenskioldi</em>, which is characterized by pronounced polymorphism (Perel’, Grafodatskij, 1983). V. S. Boeskorov, who studied the ecology of <em>E. n. nordenskioldi</em> in permafrost soils of Yakutia, identified two morpho-ecological groups of these earthworms (epi-endogeic and anecic) and defined their range (Boeskorov, 2004). Large individuals of <em>E. n. nordenskioldi</em> were also classified as anecic earthworms by T. S. Vsevolodova-Perel’ during the study of earthworms in the forests of the Western Sayan (Perel’, 1994). Ju. B. Byzova, who experimented on the intensity of Oligochaeta respiration in the soil, when describing the collected samples from different regions of Russia, often reported differences in size and weight as well as in physiological features in individuals of this subspecies, classifying them as different living forms (Byzova, 1965, 2007). She classified large individuals of <em>E. n. nordenskioldi</em> collected in Western Siberia (in particular, in the Novosibirsk area) as anecic earthworms (Byzova, 2007). This statement is confirmed by a recent study conducted in the forests of the forest-steppe Ob region in the Novosibirsk area, where morphometric analysis revealed size groups of <em>E. n. nordenskioldi</em> with earthworms having different habitat conditions (Ermolov, 2020b). An attempt was made to confirm that large-sized earthworms <em>E. n. nordenskioldi</em> are anecic earthworms on the basis of their morphological similarity with a typical representative of anecic earthworms <em>L. terrestris</em> (Ermolov, 2020b). Most of the works on molecular biology of <em>E. n. nordenskioldi</em> were performed by S. V. Shekhovcov. Within this subspecies on the territory of Russia, he managed to identify nine different genetic lineages (Shekhovcov et al., 2016, 2018). However, no analysis of the relationship between morpho-anatomical and molecular-genetic differences of individuals of this subspecies was performed; this is planned for future works (Shekhovcov, Berman, 2018). Nevertheless, it was found that earthworms of some genetic lineages differ in cold resistance: there are moderately resistant lineages (–10&#8230; –12 °C) and lineages that tolerate low temperatures (–28&#8230; –34 °C) (Berman et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The Caucasian species <em>Dendrobaena schmidti</em> Michaelsen, 1907 is also polymorphic. There is an assumption that epigeiс, epi-endogeiс and endogeic earthworms living forms can be identified within the species (Rapoport, 2009), since individuals of this species collected in different parts of the Caucasus significantly differ in body size, pigmentation intensity, development of glandular fields and vertical distribution in the soil (Shekhovcov et al., 2020b). Two genetic lineages have also been identified for <em>D. schmidti</em>, individuals in which significantly differ in size and degree of pigmentation. However, these differences may overlap in some cases, and the main taxonomic features of the species within the lineages do not differ (Shekhovcov et al., 2020b).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Polymorphism is also revealed in the endogeic <em>O. lacteum</em> (synonym of <em>O. tyrtaeum</em>). Studies conducted in Belarus and Western Siberia have shown that within this species there are one small and two large-sized earthworms that have different ecological conditions of their habitat. Large-sized earthworms are more common in wetter soils with a well-developed humus horizon, whereas smaller ones predominate in dry soils with low humus content (Shekhovcov et al., 2020; Ermolov, unpublished data). It is noteworthy that large-sized earthworms have not previously been found in Western Siberia (Shekhovtsov et al., 2014). In Belarus and in the Novosibirsk area, a relationship was found between a certain genetic lineage of individuals and their dimensional characteristics within the species (Shekhovcov et al., 2020a).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Pronounced polymorphism discovered in the endogeic cosmopolitan <em>A. caliginosa</em> when studying the populations of this species in Ukraine and Belarus was unexpected. It turned out that earthworms from different populations have significant differences in size of adult individuals, as well as various variations in body and clitellum pigmentation, from light gray and pink to brown and yellow-orange (Mezhzherin et al., 2018). It is noteworthy that polymorphism in <em>A. caliginosa</em> was not observed in Siberia and the Urals; only sometimes it was reported in some regions of Central Russia (Shekhovcov et al., 2016a; Ermolov, unpublished data). S. V. Shekhovcov and colleagues also studied the genetic diversity of <em>A. caliginosa</em> in Russia and the Republic of Belarus (Shekhovtsov et al., 2016; Shekhovcov et al., 2017). In the course of this study, several genetic lineages of this species were identified in Russia, and the morphological diversity of Belarusian earthworms is partly explained by them being part of a certain genetic lineage. However, it is impossible to clearly distinguish genetic lineages based on differences in external morphology, since some signs overlap and may be associated with some ecological features of the habitat of the species (Shekhovtsov et al., 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Sometimes it turns out that there may be several genetic lineages within one species that have few if any, morphological differences. For example, when the genetics of the European cosmopolitans <em>Aporrectodea longa</em>, <em>Aporrectodea rosea</em>,<em> Al. chlorotica</em>, and <em>L. rubellus</em> was studied in the UK, a high divergence (more than 14%) of the nucleotide sequences of the mitochondrial gene <em>cox1</em> was found in some of them. In the species <em>Al. chlorotica</em>, represented by two forms differing in color, 35 haplotypes were identified for the form with pink pigmentation and 20 haplotypes for the form with green pigmentation (King et al., 2008). Later it became necessary to use not only mitochondrial, but also nuclear markers, since only five strongly divergent lineages were identified within this species for the 16S rRNA gene (King et al., 2008). Similar results in difference of data on different genes were also found by Polish researchers in <em>L. rubellus</em> (Giska et al., 2015). At the same time, in the former USSR countries, these species are represented by only one line (Shekhovcov, spoken communication).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>3.3. Classification of earthworms by habitat conditions</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In addition to the classification of the living forms of Lumbricidae mentioned above, researchers have also suggested other options for the identification of earthworm ecological groups based on their relationship with some abiotic factors.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">One of the good examples is the classification of earthworms by cold resistance developed by D. I. Berman, A. N. Lejrih and E. N. Meshherjakova (Meshherjakova, 2011; Lejrih, 2012). The earthworms under study (sampled in different regions of Russia) were divided into three groups: species resistant to below-freezing temperatures in the worm and cocoon phase; species resistant to below-freezing temperatures only in the cocoon phase; and species not resistant to temperatures below –1 °C at any of the phases of ontogenesis. However, each group includes representatives of different living forms, and no connection was found between the cold resistance of earthworms and they are belonging to a particular living form since everything depends on physiological characteristics of a particular species (Berman, Lejrih, 1985; Meshherjakova, 2011).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A very interesting classification of earthworms based on their relation to soil moisture was first proposed by O. V. Zhukov et al., which identifies groups of mesophiles, hygrophiles and ultragygrophiles (Zhukov et al., 2007; Kunah et al., 2010). However, this classification is also only partially consistent with the system of living forms by Vsevolodova-Perel’ or Bouche: for example, all subgroups of amphibiotic earthworms can be attributed to ultrahygrophiles, whereas species belonging to other living forms are classified as either hygrophiles or mesophiles. Nevertheless, Zhukov showed that the system he proposed can be used in zooindication to assess the degree of soil moisture with the help of a complex of earthworms (Zhukov, 2004).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Earthworms are most often used as indicators in the studies of soil acidity (Giljarov, 1965). For example, the experiments of A. I. Zrazhevskij proved the effect of an anion of a certain acid that forms the pH of the soil on earthworms (Zrazhevskij, 1957). The works of O. P. Atlavinite looked into correlations between the population and occurrence of individual earthworm species, on the one hand, and soil pH, on the other hand (Atlavinite, 1960). It was shown that some earthworm species are very resistant to a wide pH range, for example, <em>A. caliginosa</em>, while others are most commonly found in acidic soils (<em>D. octaedra</em>) or prefer neutral and slightly alkaline soils (<em>E. fetida</em>). Earlier studies by R. Baltzer showed that soils of different types with different pH values are inhabited by certain species and complexes of earthworms living forms (Baltzer, 1955). This study implied that, based on the predominance of a particular earthworm species, soil pH can be determined: for example, <em>L. rubellus</em> inhabits acidic and slightly acidic soils, <em>A. caliginosa</em> and <em>A. rosea</em> prefer slightly acidic and neutral soils, whereas <em>O. lacteum</em> inhabit neutral and alkaline soils. However, pH value is by no means the only feature that determines the suitability of soil as a habitat of earthworms. In addition to pH, there are many other physical and chemical soil properties that significantly affect earthworms: for example, a study of earthworms in Western Siberia revealed <em>L. rubellus</em> in acidic (pH = 5.42), slightly acidic (pH = 5.72) and neutral (pH = 7.49) soils, but its greatest population was found in neutral floodplain soils (149 ± 31 individuals/m<sup>2</sup>) due to the highest moisture content (Ermolov, 2020b). Therefore, when studying the relationship of earthworms with soil acidity, it is important to take into account other soil factors as well, such as humidity, content of organic matter, nitrogen, calcium and other macro- and micronutrients (Ivask et al., 2006). However, soil pH, changed by anthropogenic impact, often becomes a limiting factor for earthworms. For example, experiments conducted in China showed that in areas with frequent acid rains, earthworm populations are at risk of complete extinction, since earthworms are not able to survive in soil with pH of 2 and below (Zhang et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is known that earthworms are calciphilic organisms that have special organs for alkalizing acidic food, i. e. <em>calciferous glands</em> (Chekanovskaja, 1960; Vsevolodova-Perel’, 1997; Gaponov, Hicova, 2005). Previously, it was believed that calcareous glands can be of only three types and have no significance as a taxonomic feature (Vsevolodova-Perel’, 1997). Recent studies have shown, however, that the anatomy of calcareous glands is very diverse: a detailed analysis of 13 genera of earthworms identified seven groups of species with different structure of calcareous glands (Briones, Piearce, 2011; cit. by: Biology of Earthworms, 2011). Moreover, the same types of gland structure were often found among different genera. This suggests that the taxonomic position of some species still remains ambiguous. The structure of calcareous glands also makes it possible to indirectly assess the acidity of the habitat of earthworms and their food preferences (Briones, Piearce, 2011; cit. by: Biology of Earthworms, 2011).</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Studies of the ecology of earthworms originate from ancient times and still remain relevant. First simple observations of ancient thinkers and medieval naturalists gradually began to be generalized by naturalists of modern times, who gave them a scientific justification and thereby proved the significant role of earthworms in soil formation.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Later, earthworm classification became a relevant issue. For many decades, various taxonomic systems of earthworms have been developed, but the question of taxonomy remains open even today. In addition to taxonomy, there has been great interest to ecological classification of earthworms, the “classic variants” of which were created in the 1970s in France and Russia. Since the end of the last century, various researchers have been trying to expand the proposed classifications, supplementing the main ecological groups with intermediate groups and subcategories.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The ecological classification of earthworms has found ample applications in various studies. By analyzing the structure and species composition of complexes of earthworms living forms and the ability of individual representatives to live under certain environmental conditions, researchers obtained a unique tool to diagnose soils in different biotopes. Between earthworms of different living forms and other soil organisms trophic and functional connections have been identified that ensure the flow of substances and the maintenance of biodiversity in ecosystems. When studying the polymorphism of earthworms, it was found that there can be different living forms even within the same species, which, based on the evidence from molecular biology, are later often classified as new species.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">However, there are still a lot of unanswered questions that arise when studying the ecology of earthworms. For example, can all different genetic lineages be considered different species and how can this be proven in terms of ecological and genetic concepts? Are some earthworm species invasive to a number of regions or have they always lived there? What are the limiting factors for endemic species and cosmopolitans; is their cohabitation possible? What environmental factors can cause polymorphism in earthworms and what is the reason for the transition to parthenogenesis?</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">These and other issues require further development of methods to study the features of earthworm ecology, and mainly enhancing the experimental component. It is especially important to conduct comprehensive studies at the confluence of ecology and genetics, taxonomy and zoogeography, climatology and soil science. This will help us find answers to the questions posed and lay the foundation for further research.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FINANCING</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The study was performed within the framework of the state CEPF RAS assignment 121121600118-8. </span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
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<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Voronova N. V., Buga S. V., Kurchenko V. P., Posledovatel’nost’ gena subedinicy I citohromoksidazy <em>C</em> v molekuljarnoj taksonomii zhivotnyh: principy, rezul’taty i problemy ispol’zovanija (COI-5 region as a marker for molecular taxonomy of animals: approaches, outcomes and constraints), <em>Trudy BGU</em>, 2012, Vol. 7, part 1–2, pp. 22–42.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Vsevolodova-Perel’ T. S., <em>Dozhdevye chervi fauny Rossii: Kadastr i opredelitel’</em> (The earthworms of the fauna of Russia: Cadaster and key), Moscow: Nauka, 1997, 102 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Vysockij G. N., Dozhdevoj cherv’ (Earthworm),<em> Polnaja enciklopedija russkogo sel’skogo hozjajstva</em>, 1900, Vol. 2, pp. 12–39.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Wilcke D. E. Über die vertikale Verteilung der Lumbriciden im Boden, <em>Z. Morphol. und Ökol. Tiere</em>, 1953, No 41, pp. 372–385.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhang J.-E., Yu J., Ouyang Y. Activity of Earthworm in Latosol Under Simulated Acid Rain Stress, <em>Bulletin of environmental contamination and toxicology</em>, 2015, Vol. 94, No 1, pp. 108–111.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhang Y. F., Ganin G. N., Atopkin D. M., Wu D. H., Earthworm <em>Drawida </em>(Moniligastridae) Molecular phylogeny and diversity in Far East Russia and Northeast China, <em>The European Zoological Journal</em>, 2020, Vol. 87, No 1, pp. 180–191.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhukov O. V., Dozhdevye chervi kak komponent biogeocenoza i ih rol’ v zooindikacii (Earthworms as a component of biogeocenosis and their role in zooindication), <em>Gruntoznavstvo</em>, 2004, Vol. 5, No 1–2, pp. 44–57.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhukov O. V., Pahomov O. E., Kunah O. M., <em>Biologicheskoe raznoobrazie Ukrainy. Dnepropetrovskaja oblast’. Dozhdevye chervi </em>(Lumbricidae) (Biological diversity of Ukraine. Dnipropetrovsk region. Earthworms (Lumbricidae)), Dnepropetrovsk: Izd-vo Dnepropetr. nac. un-ta, 2007, 371 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zrazhevskij A. I., <em>Dozhdevye chervi kak faktor plodorodija lesnyh pochv</em> (Earthworms as a factor of forest soils fertility), Kiev: Izd-vo AN SSSR, 1957, 271 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewers:</strong> Candidate of Biological Sciences E. V. Golovanova, Candidate of Biological Sciences I. I. Lyubechansky</span></p>
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		<title>I. S. MELEKHOV’S SCALE OF NATURAL FIRE DANGER OF FOREST ECOSYSTEMS: OVERVIEW OF MODERN RUSSIAN METHODOLOGICAL APPROACHES</title>
		<link>https://jfsi.ru/en/5-1-2022-plotnikova/</link>
		
		<dc:creator><![CDATA[lena]]></dc:creator>
		<pubDate>Mon, 25 Apr 2022 07:56:47 +0000</pubDate>
				<category><![CDATA[№1 2022]]></category>
		<guid isPermaLink="false">https://jfsi.ru/?p=5018</guid>

					<description><![CDATA[Original Russian Text © 2021 A. S. Plotnikova published in Forest Science Issues Vol. 4, No. 2,   Article 83 A. S. Plotnikova Center for Forest Ecology and Productivity of the Russian Academy of Sciences Profsoyuznaya&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/04/5-1-2022-Plotnikova.pdf"><img loading="lazy" class="size-full wp-image-1122 alignright" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></span></p>
<p style="font-weight: 400;"><span style="font-family: 'times new roman', times, serif; color: #000000; font-size: 10pt;"><span lang="EN-US">Original Russian Text © 2021 A. S. Plotnikova published in Forest Science Issues Vol. 4, No. 2,   </span><a style="color: #000000;" href="https://jfsi.ru/4-2-2021-plotnikova/" target="_blank" rel="noopener noreferrer">Article 83</a></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>A. S. Plotnikova</strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Center for Forest Ecology and Productivity of the Russian Academy of Sciences </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Profsoyuznaya st. 84/32 bldg. 14, Moscow 117997, Russian Federation</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">E-mail: <a style="color: #000000;" href="mailto:plotnikova-as-cepl@yandex.ru">plotnikova-as-cepl@yandex.ru</a></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 10.05.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 08.06.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 29.06.2021</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The article reviews various methodological approaches to assessing natural fire danger (NFD) and the creation, updating, and application of NFD maps, which modern domestic researchers develop. Herein we introduce and analyse the currently accepted natural fire danger assessment scale by I. S. Melekhov. As pointed out by modern researchers, the methodological drawbacks of the scale are listed. The paper reviews the development of a new methodological approach to the compilation of regional scales for assessing the natural fire danger of forests, considering the links between forest growth conditions and seasonal and climatic conditions in the regions of the Russian Federation. The methodology of mapping natural fire danger based on vegetation fuel maps proposed by a scientific group of Sukachev Forest Institute of SВ RAS was studied. The works of the Mytishchi branch of Bauman Moscow State Technical University were reviewed to investigate the possibility of applying mathematical modelling methods for long-term forecasting of changes in NFD under different scenarios of forest management. The method of annual mapping of NFD classes, as proposed in CEPF RAS, is described. An example of NFD maps to assess the probability of forest fires in the Institute for Complex Analysis of the FEB RAS Regional Problems is considered. Future research areas are identified, namely, a cartographic representation of the created regional scales of NFD and the results of mathematical modelling of long-term changes in NFD.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><em>Keywords:</em></strong><em> natural fire danger, </em><em>vegetation fuel maps, forest fires</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Fires are a crucial factor in the development of forest ecosystems. The probability of occurrence and further spread of a forest fire depends on the degree of fire danger. There are two types of fire danger in forests — natural and depending on weather conditions. In this paper, we focus on the natural fire danger (NFD) to study various methodological approaches to its assessment and the creation, updating, and application of NFD maps offered by modern domestic researchers.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In Russia, the natural fire danger of a forest district is assessed using the scale of I. S. Melekhov (Melekhov et al., 2007). For forest fire protection services, the use of the NFD classification is regulated by Order of the Federal Forestry Agency No. 287 dated 05 July 2011 “On approval of the classification of natural fire danger of forests and classification of fire danger in forests depending on weather conditions”. Depending on the object of ignition (typical types of forests, logging areas, other categories of stands and treeless spaces), as well as the conditions for the occurrence and spread of fire, five classes of NFD are distinguished. The first class corresponds to the most fire-dangerous areas with the highest probability of fire occurrence and spread. In contrast, the fifth class corresponds to the areas with the least or no probability thereof.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Classification according to the classes of natural fire danger of forests is one of the critical tasks of monitoring fire danger in forests (Shur et al., 2020). Strategic and operational management of forest fire units implies the use of an NFD concept. Classes of natural fire hazard (CNFDs) as part of strategic management are necessary for planning forest fire prevention measures, a compilation of regional consolidated fire extinguishing plans and forest development projects, forest plans of the subjects of the Russian Federation, as well as forest management regulations of forestries and forest parks. Operational management includes the application of CNFDs in regulating fire monitoring and extinguishing by ground-based forest fire units.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CLASSIFICATION OF NATURAL FIRE DANGER</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Let us take a look into the accepted classification of natural fire danger. The classification introduces the concept of a fire maximum, i. e. a period when the number of forest fires or the area covered by fire exceeds the average long-term values for a given area. Thus, Class I has a very high NFD, and ground and crown fires are possible during the entire fire season. Based on the object of ignition, this class includes young coniferous stands, lichen and heather pine forests, lichen, heather, reedgrass and other types of clear-cuts along dry valleys. Significant spring and autumn fire danger is reported on reedgrass and other herb types of logging areas along dry valleys. Besides, areas of selective logging of high and very high intensity, continuous logging sparing individual trees, as well as debris-strewn (cluttered) fire sites and disturbed, suppressed and severely damaged stands in the form of deadwood, areas of wind snaps and windthrows, residual stands are prone to ignition.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">With a high NFD, which is typical of Class II, ground fires can occur during the entire fire season, whereas crown fires occur during periods of fire maxima. The objects of ignition are cowberry pine forests, especially with pine undergrowth or juniper undergrowth above-average density, and larch forest with cedar elfin. Cedar forests with dense undergrowth or different ages with vertical canopy closure are also considered Class II forests. In areas with a medium NFD (Class III), ground and crown fires are possible during the summer fire maximum. This class includes the following types of forest: sorrel and bilberry pine forests, cowberry larch forests, cowberry and sorrel spruce forests, cedar forests of all types, except for streamside and sphagnum ones. It is reported that in cedar forests, fires can also occur during spring and autumn maxima.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In forests with low NFD (class IV), fires occur only during the summer maximum in all types of forests and haircap-moss logging areas. The exceptions are areas with herb forest types and meadowsweet logging areas, where ground fires may also occur during the spring and autumn fire maxima. The objects of ignition are pine forests, larch forests and forest stands of deciduous tree species in herb forest types. The classification details the objects of ignition: complex pine forests and spruce forests — linden, hazel, oak; bilberry spruce forests; sphagnum and haircap-moss pine forests; streamside and sphagnum cedar forests; cowberry, sorrel, bilberry and sphagnum birch forests; sorrel and bilberry aspen forests; larch peat moss bog forests. Class IV is also characterised by cluttered parts of solid meadowsweet and haircap-moss logging areas. With class V, there is no NFD — a fire is possible only with particularly unfavourable conditions in the form of a prolonged drought. This class includes sphagnum and streamside spruce forests, haircap-moss birch and aspen forests, and alder forests of all types.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The classification defines cases when a natural fire danger of a higher class is identified. Firstly, this is the case of coniferous forest stands, where the structure or other features contribute to the transition of ground fire to a crown fire. Such features may include a dense high undergrowth of coniferous tree species, vertical closure of the crown canopy of trees and shrubs, and significant cluttering. Secondly, for small forest plots on dry valleys surrounded by the forest stands with increased natural fire danger. Thirdly, for forest areas adjacent to railways and public roads.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It should be mentioned that the natural fire danger of a forest area can be controlled through forest fire prevention measures within the territory. It includes, among others, regulation of the stand composition, sanitary logging; prevention of forest cluttering; creation of a network of fire barriers and reservoirs, forest roads, and recreational areas (Chumachenko, Mayuk, 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to A. V. Sofronova’s and A. V. Volokitina’s (2017) work, the above scale was compiled by an expert method, which explains the lack of quantitative assessments of NFD classes. The scale reflects generalised natural fire danger, which takes into account readiness for ignition, seasonal duration of stay in a ready-to-ignite state, the possibility of crown fire occurrence, as well as the difficulty of extinguishing fires in cluttered areas. The authors highlighted such a drawback of the scale as the lack of characteristics of vegetation fuel (VF).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to Yu. Z. Shur et al. (2020), the scale of I. S. Melekhov has several significant methodological shortcomings; for example, regional forest typology is not taken into account, the relationship between the periods of the fire season, predominant types of vegetation fuel and the most likely types of forest fires is not clearly defined. The scale offers no logically rigorous definition of the objects of ignition, and their CNFDs are not established. In addition, the categories of forest and non-forest lands are not fully described.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>REGIONAL ASSESSMENT SCALES OF NATURAL FIRE DANGER</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">We will consider a new methodological approach to the compilation of regional scales for assessing the natural fire danger of forests, considering the interrelations between forest, seasonal and climatic conditions in the entities of the Russian Federation offered by the St. Petersburg Research Institute of Forestry (Shur et al., 2020). The authors introduce the concept of a “regional scale of assessment of natural fire danger of forests” to classify forest NFD for a Russian Federation entity. For each object of ignition and the period of the fire season, the prevailing types of vegetation fuels and the most likely types of forest fires are determined, e. g. running ground fire, independent ground fire, subsurface fire, crown fire. Thus, the approach makes it possible to predict the occurrence of forest fires of various types. It is proposed to consider fuels that make possible the occurrence of ground forest fires as predominant types of vegetation fuels. The proposed scale considers the categories of forest lands not covered by forest vegetation and non-forest lands in more detail.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to the methodology, the main spatial unit for determining the CNFDs is the forest inventory allotment (Table). However, it is possible to determine the weighted average CNFD for a forest block, district forestry, and forestry territory. The authors tested the new methodological approach in several entities of the Russian Federation, in particular the republics of Karelia, Adygea, North Ossetia-Alania; Primorsky and Krasnodar Krai; Tomsk, Rostov, Leningrad and Volgograd Oblast. Regional scales for assessing the natural fire danger of forests were compiled based on schemes of forest types and forest growth conditions of the <em>Roslesinforg</em> Federal State Budgetary Institution.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The authors developed a regional scale for assessing NFD in the area of the Republic of Karelia, shown below. The objects of ignition for very high NFD (class I) are rocky and white moss pine forests, including incomplete stands; lichen logging areas; dead forest stands, except for fire sites, old fire sites, clearings, waste ground in rocky and white moss forest types (FTs); meadows, hayfields, dry alley pastures. The predominant types of VFs and the most likely types of forest fires are determined for the spring, summer and autumn periods of the fire season. For example, in the spring period, lichen-type VF and independent ground fires prevail in all the objects of ignition, except meadows, hayfields and pastures. The following objects of ignition are characteristic of high NFD (class II): heather and cowberry pine forests, including incomplete stands; heather logging areas; dead forest stands, except for fire sites, old fire sites, clearings, waste ground in heather and cowberry FTs. In heather pine forests, the heather or dwarf shrub type of VF prevails, as well as litterfall. In cowberry pine forests, a green moss type of VF or litterfall is predominant. Subsurface, running ground, and independent ground fires are seen in all objects of ignition during any period of the fire season.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>MAPS OF NATURAL FIRE DANGER</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Researchers from the V. N. Sukachev Institute of Forest SB RAS state that the most optimal form for the representation and use of the natural fire danger is in the form of maps (Abroskina et al., 2012). The Institute has developed a methodology for mapping natural fire danger based on maps of vegetation fuels (Volokitina, Sofronova, 2014). NFD mapping for long-term monitoring and the monitoring of the current NFD depending on the fire danger classes according to weather conditions is also an option.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to the definition by N. P. Kurbatskii, vegetable fuels are plants and their residues of various degrees of decomposition that can burn during fires (Kurbatskii, 1970). VF maps are compiled based on their classification, which includes seven groups: prime conductors of burning; litterfall, humus and peat horizons; herb-dwarf shrub layers; large woody debris; a layer of shrubs and undergrowth; needles, foliage, bearing twigs and dry branches in tree crowns, tree trunks and branches (Volokitin, Sofronova, 2014). Only the prime conductors of burning in the ground cover of forest inventory allotments are displayed directly on a vegetation fuel map. Characteristics of all VF groups can be found in the pyrological description, which is attached to the map. The pyrological description can predict the spread and create an operational plan for extinguishing an active fire.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A prime conductor of burning (PCB) is a continuous layer of vegetation fuel on the soil surface, conducting flaming combustion under certain conditions (Volokitina, Sofronova, 2014). The layer includes small plant residues, including twigs up to 2 cm in diameter; mosses and lichens; vascular plants and their parts — stems of grasses and dwarf shrubs as well as small plants. The authors identify two subgroups of PCBs — «mossy», i. e. layers with a predominance of living fuel (mosses and lichens), and «litterfall» — layers with a predominance of dead fuel (fallen needles and foliage, dried herbs). The authors describe several possible ways to determine the type of PCB directly in the field, based on the inventory description of forest management materials, thematic maps of vegetation cover, and decrypting the Earth remote sensing data. In all methods, the identification guide of the PCB type by A. V. Volokitina is used (Red’kin, Volokitina, 2014). When using inventory descriptions of forest management materials, the type of PCB is established in accordance with the description of forest types, which includes a typical location on the relief; the name of the soil and its moisture regime; typical composition of the stand; bonitet; description of the undergrowth, herb-dwarf shrub layer, and moss-lichen cover; as well as the characteristics of forest regeneration.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">If up-to-date forest management materials are lacking, high- and ultra-high-resolution satellite images can be used, as well as vector data of the hydrographic network and relief (Sofronova, Volokitina, 2017a). It is proposed to interpret the PCBs closed by the forest canopy by identifying pyrological categories of sites.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The spatial unit for assessing the natural fire danger based on maps of vegetation fuels is the inventory allotment (Table). This assessment seems more accurate than methods using forest fire maps at the block level. Since the same forest area in spring, summer and autumn can have different types of PCBs, NFD mapping by periods of fire season is possible.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The method of mapping natural fire danger developed at the V. N. Sukachev Institute of Forest SB RAS was tested at the local level in the territories of the experimental farm Pogorelsky Bor, Yemelyanovsky Forestry, the Krasnoyarsk Krai (Abroskina et al., 2012) and the Yurubcheno-Tokhomskoye oil and gas condensate field (Sofronova, Volokitina, 2017b). In addition, maps of the natural fire danger of the Stolby, Sayano-Shushensky, Kuznetsky Alatau and Ubsunurskaya hollow reserves were created (Volokitina, 2017).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>MODELING THE DYNAMICS OF NATURAL FIRE DANGER</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The Mytishchi branch of the N. E. Bauman Moscow State Technical University (formerly Moscow State University of Forestry) is investigating the possibility of using mathematical modelling methods for long-term forecasting of changes in natural fire danger under different scenarios of forest management (Chumachenko, 2012; Chumachenko, Mayuk, 2012; Chumachenko, Mukhin, 2013). A conceptual, mathematical and simulation model of the dynamics of natural fire danger with a simulation step of 5 years has been developed.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">S. I. Chumachenko and D. N. Mayuk (2012) identify factors that determine the natural fire danger — that is, land categories, age of stand, type of forest, predominant and admixed tree species, completeness of stands, the presence of fire-dangerous undergrowth and underbrush, as well as deadwood and cluttering, distance from public roads. The factors mentioned earlier are the main parameters of the natural fire danger dynamics model, and similar data can be found in inventory descriptions and plans of forest stand at the allotment level (Table). Modelling includes a forecast of changes in the species and age composition of the allotment, the average taxation characteristics of the stand by layers, such as height, diameter, age, stock, etc. Due to the lack of a methodology for determining forest types when forecasting the dynamics of stands for a long period under different forest management scenarios, this parameter was replaced by the type of forest conditions.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>UPDATING MAPS OF NATURAL FIRE DANGER</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is commonly known that the subjects of the Russian Federation are developing forest plans, i. e. a document defining the main directions of forest use and regeneration for the next ten years (Forest Code of the Russian Federation, 2006). A forest plan contains forest fire maps of forest districts, which usually provide information about the NFD class at the forest block level. During the ten-year inter-revision period, the type of ground cover and woody vegetation changes under the influence of various destructive factors, e. g. fires, logging, and plagues. These changes render the NFD assessment irrelevant.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The Centre for Forest Ecology and Productivity RAS has proposed a method for updating CNFD maps annually (Plotnikova, Ershov, 2015). This method makes it possible to determine the class of natural fire danger through a comprehensive analysis of thematic satellite maps of vegetation cover, long-term data on fires and meteorological observations, and CNFD data from the forest plan of the entity Russian Federation. The method uses data from meteorological observations (average daily air temperature) to determine the time limits of the spring, summer and autumn periods of the fire season. Long-term fire data are used to determine fire maxima for each year under study based on the analysis of the long-time average annual number of fires per day. Based on thematic satellite maps of vegetation cover, areas of vegetation classes within the boundaries of forest blocks are estimated (Table).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Evaluation of the method of updating CNFD maps was carried out on the territory of the Irkutsk Oblast. Long-term archives of fire data from 1987 to 2011 and meteorological observations from 2006 to 2011 were used. In the study of A. S. Plotnikova and D. V. Ershov (2015), a map of vegetation cover on the territory of Russia with a spatial resolution of 250 meters was used (Bartalev et al., 2011). Further research (Plotnikova, Ershov, 2016) was carried out using a vegetation map of the territory of the Irkutsk Oblast, which had been created based on high spatial resolution satellite data Landsat-TM\\ETM+.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Table.</strong> Key information about modern scientific research of NFD in Russia</span></p>
<div style="overflow-x: auto;">
<table style="border: 1px #f1f1f1 solid; background-color: #ffffff;">
<tbody>
<tr>
<td width="177"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Scientific group</strong></span></td>
<td width="161"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Spatial level</strong></span></td>
<td width="170"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Spatial unit</strong></span></td>
<td width="246"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Initial data</strong></span></td>
<td width="217"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Method</strong></span></td>
</tr>
<tr>
<td width="177"><span style="font-family: 'times new roman', times, serif; color: #000000;">V. N. Sukachev Institute of Forest SB RAS</span></td>
<td width="161"><span style="font-family: 'times new roman', times, serif; color: #000000;">Local</span></td>
<td width="170"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest inventory allotment</span></td>
<td width="246"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest management materials,</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">VF maps, high and ultra-high-resolution Earth remote sensing data, vector data of the hydrographic network and relief</span></td>
<td width="217"><span style="font-family: 'times new roman', times, serif; color: #000000;">NFD mapping based on VF maps</span></td>
</tr>
<tr>
<td width="177"><span style="font-family: 'times new roman', times, serif; color: #000000;">St. Petersburg Scientific Research Institute of Forestry</span></td>
<td width="161"><span style="font-family: 'times new roman', times, serif; color: #000000;">Regional</span></td>
<td width="170"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest inventory allotment,</span><br />
<span style="font-family: 'times new roman', times, serif; color: #000000;">forest block,</span><br />
<span style="font-family: 'times new roman', times, serif; color: #000000;">district forestry,</span><br />
<span style="font-family: 'times new roman', times, serif; color: #000000;">forestry</span></td>
<td width="246"><span style="font-family: 'times new roman', times, serif; color: #000000;">Schemes of forest types and forest growth conditions of <em>Roslesinforg</em> FSBI</span></td>
<td width="217"><span style="font-family: 'times new roman', times, serif; color: #000000;">Compilation of regional scales for NFD evaluation</span></td>
</tr>
<tr>
<td width="177"><span style="font-family: 'times new roman', times, serif; color: #000000;">Mytishchi branch</span><br />
<span style="font-family: 'times new roman', times, serif; color: #000000;">of N. E. Bauman MSTU</span></td>
<td width="161"><span style="font-family: 'times new roman', times, serif; color: #000000;">Local</span></td>
<td width="170"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest inventory allotment</span></td>
<td width="246"><span style="font-family: 'times new roman', times, serif; color: #000000;">Inventory descriptions,</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">plans of forest stands</span></td>
<td width="217"><span style="font-family: 'times new roman', times, serif; color: #000000;">Mathematical modelling for long-term forecasting of changes in NFD under different scenarios of forest management</span></td>
</tr>
<tr>
<td width="177"><span style="font-family: 'times new roman', times, serif; color: #000000;">Centre for Forest Ecology and Productivity of the RAS</span></td>
<td width="161"><span style="font-family: 'times new roman', times, serif; color: #000000;">Regional</span></td>
<td width="170"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest block</span></td>
<td width="246"><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest plan of the entity of the Russian Federation, forest fire maps, medium and high-resolution Earth remote sensing data,</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">satellite thematic maps of vegetation cover, a long-term archive of data on fires and meteorological observations</span></td>
<td width="217"><span style="font-family: 'times new roman', times, serif; color: #000000;">Annual updating of CNFD maps</span></td>
</tr>
</tbody>
</table>
</div>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>APPLICATION OF MAPS OF NATURAL FIRE DANGER </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Maps of natural fire danger are used to assess the likelihood of forest fires. An example is the work of the Institute for Integrated Analysis of Regional Problems of the FEB RAS on the creation of a system for spatial forecasting of vegetation fires depending on weather and forest conditions in the south of the Russian Far East (Kogan, Glagolev, 2013). As the authors note, the possibility of fire occurrence depends on the processes of drying and moistening of vegetation, which determine its transition to a state of “fire maturity” (Kogan, Glagolev, 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to the method developed by R. M. Kogan and V. A. Glagolev, one of the values necessary to determine the probability of fires is the degree of natural pyrologic fire danger of vegetation in the spring/summer/autumn periods of the fire season. The authors use a scale (Starodumov, 1964; Telitsyn, 1988) that classifies vegetation on the territory of the Middle Amur region as one of five danger classes. A very high degree (class I) of natural pyrologic fire danger is typical of areas not covered with forest, open stands, logging areas with grass cover or reindeer lichen, as well as larch-spruce and spruce-larch mountain forests. As forests of high degree (class II) are classified broad-leaved-spruce cedar forests (northern cedar forests); small-leaved forests and sparse forests on mountain slopes; broad-leaved shrub forests on slopes; broad-leaved oak forests on eastern and western slopes. The moderate degree (class III) includes fir-spruce and spruce-fir forests with cedar, broad-leaved species (nemoral spruce forests), i. e. green moss; dwarf shrub-small herbs-green moss; forests of the middle mountain belt, as well as aspen and mixed forests on northern slopes. Moderate degree (class IV) is typical of dwarf shrub-moss larch forests with dwarf birch and sparse forests of intermountain valleys. Low degree (class V) includes sedge-sphagnum larch forests, sphagnum bog spruce forests and constantly moistened sphagnum swamps.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the work of R. M. Kogan and V. A. Glagolev (2015а), a description of the forest fund of the Far Eastern Federal District is given based on natural fire danger. Due to the region’s climatic, forest growth, and geomorphological features, the forest fund has high fire danger and frequency of fire occurrence. The natural fire danger of the forests of the Khabarovsk Krai and the Jewish Autonomous Oblast is one of the highest in Russia: the area of plant formations with classes I–III is more than 80% and 44% of the entity territory, respectively.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The performed review showed the current state of domestic scientific research on natural fire danger. Various scientific groups agree that the currently accepted scale of NFD assessment has several methodological drawbacks, such as the lack of characteristics of vegetation fuels and the relationship between the periods of the fire season, etc. To improve the assessment of natural fire danger, researchers are developing new methodological approaches, e. g. compile regional scales for assessing forest NFD, considering the relationship of forest growth, seasonal and climatic conditions in the entities of the Russian Federation, as well as create NFD maps based on vegetation fuel maps.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">One of the modern lines of research of natural fire danger is mathematical modelling methods for long-term forecasting of changes in NFD under different forest management scenarios. The researchers also highlight the need for annual updating of information about the classes of NFD stated on forest fire maps in the forest plans of the entities of the Russian Federation.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The created scales and maps of NFD are used for tackling not only production tasks of strategic and operational management of forest fire formations but also scientific issues. In particular, the use of NFD maps allows increasing the accuracy of assessment of the probability of forest fires.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">As noted above, maps are the most optimal representation and natural fire danger information. Therefore, the cartographic representation of the created regional NFD scales and the results of mathematical modelling of long-term NFD changes seems to be a promising line of research. One of the possible ways to improve the updating of the NFD maps is the transition to a more detailed spatial unit, which is the forest inventory allotment.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FINANCING</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The study was conducted within the framework of the state CEPF RAS assignment 121121600118-8. </span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>REFERENCES</strong></span></p>
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<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Forest code of the Russian Federation</em>, No 200-FZ of December 4, 2006, URL: https://clck.ru/dWRPn (2021, 12, July).</span></p>
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<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Kogan R. M., Glagolev V. A., Sistema prostranstvennogo prognoza vozniknoveniya pozharov po pogodnym i lesorastitel’nym usloviyam (System of the spatial forecast of fires emergence on weather and forest vegetation conditions), <em>Bezopasnost’ v tekhnosfere</em>, 2013, Vol. 2, No 5, pp. 11–20.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Kurbatskii N. P., Issledovanie kolichestva i svoistv lesnykh goryuchikh materialov (Investigation of the quantity and properties of forest fuel), In: <em>Voprosy lesnoi pirologii</em> (Forest pyrology issues), Krasnoyarsk: ILiD SO AN SSSR, 1970, pp. 5–58.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Melekhov I. S., Dusha-Gudym S. I., Sergeeva E. P., <em>Lesnaya pirologiya</em> (Forest pyrology), Moscow: Izd-vo GOU VPO MGUL, 2007, 296 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Order of the Federal Forestry Agency</em> “On approval of the classification of natural fire danger of forests and classification of fire danger in forests depending on weather conditions” No 287, 05 July 2011, URL: https://base.garant.ru/12189021/ (2021, 12, July). </span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Plotnikova A. S., Ershov D. V., Aktualizatsiya kart klassov prirodnoi pozharnoi opasnosti lesnykh ekosistem s ispol’zovaniem sputnikovykh tematicheskikh produktov vysokogo razresheniya (Updating natural fire danger maps using high-resolution satellite thematic products), <em>Chetyrnadtsataya Vserossiiskaya otkrytaya konferentsiya “Sovremennye problemy distantsionnogo zondirovaniya Zemli iz kosmosa</em>” (Current problems in remote sensing of the Earth from space), Moscow, 14–18 November 2016, Space Research Institute of the RAS, p. 104.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Plotnikova A. S., Ershov D. V., Metod aktualizatsii kart klassov prirodnoi pozharnoi opasnosti lesnoi territorii s pomoshch’yu sputnikovykh tematicheskikh produktov (The method to update maps of forest natural fire danger levels using satellite-derived thematic products), <em>Sovremennye problemy distantsionnogo zondirovaniya Zemli iz kosmosa</em>, 2015, Vol. 12, No 1, pp. 181–189.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Red’kin A. Ju., Volokitina A. V., Opredelenie tipov osnovnyh provodnikov gorenija v processe lesoustrojstva (Determination of the prime conductors of burning types in forest management), <em>Khvoinye boreal’noi zony</em>, 2014, Vol. 32, No 3–4, pp. 47–52.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Shur Yu. Z., Neshataev V. Yu., Stepchenko A. A., Shapoval N. V., Regional’nye shkaly otsenki prirodnoi pozharnoi opasnosti lesov (Regional natural forest fire danger scales), <em>Trudy Sankt-Peterburgskogo nauchno</em><em>&#8211;</em><em>issledovatel’skogo instituta lesnogo khozyaistva</em>, 2020, No 2, pp. 59–69.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Sofronova A. V., Volokitina A. V., Kartografirovanie rastite’nykh goryuchikh materialov metodom visual’no-instrumental’nogo deshifrirovaniya kosmicheskikh snimkov (Vegetation fuel mapping by the method of visual-instrumental interpretation of space images), <em>Geografiya i prirodnye resursy</em>, 2017a, No 4, pp. 189–196.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Sofronova A. V., Volokitina A. V., Otsenka prirodnoi pozharnoi opasnosti lesnykh uchastkov na territorii neftegazovykh kompleksov s ispol’zovaniem dannykh distantsionnogo zondirovaniya Zemli (Assessment of fire hazard for forest sites at the territory of oil and gas complexes using Earth remote sensing data), <em>Sibirskij Lesnoj Zurnal</em>, 2017b, No 5, pp. 84–94.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Starodumov A. M., <em>Shkala pozharnoi opasnosti nasazhdenii i drugikh kategorii ploshchadei dlya uslovii Dal’nego Vostoka</em> (Fire danger classes for the conditions of the Far East), Khabarovsk: Dal’NIILKh, 1965, 1 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Telisyn G. P., <em>Lesnye pozhary i ikh preduprezhdenie v Khabarovskom krae</em> (Forest fires and their prevention in the Khabarovsk region), Khabarovsk: Dal’NIILKh, 1988, 94 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Volokitina A. V. Sovershenstvovanie otsenki prirodnoi pozharnoi opasnosti v zapovednikakh (Improving the assessment of the natural fire hazard in nature reserves), <em>Geografiya i prirodnye resursy</em>, 2017, No 1, pp. 55–61.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Volokitina A. V., Sofronova T. M., Kartografirovanie rastitel&#8217;nykh goryuchikh materialov (Vegetation Fuel Mapping), <em>Sibirskii lesnoi zhurnal</em>, 2014, No. 6, pp. 8–28.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewer: </strong>Candidate of Geography Sciences V. A. Glagolev</span></p>
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		<title>TREE LITTER PRODUCTION AND DECOMPOSITION IN FOREST ECOSYSTEMS UNDER BACKGROUND CONDITIONS AND INDUSTRIAL AIR POLLUTION</title>
		<link>https://jfsi.ru/en/5-1-2022-ivanova/</link>
		
		<dc:creator><![CDATA[lena]]></dc:creator>
		<pubDate>Mon, 18 Apr 2022 14:01:28 +0000</pubDate>
				<category><![CDATA[№1 2022]]></category>
		<guid isPermaLink="false">https://jfsi.ru/?p=5000</guid>

					<description><![CDATA[Original Russian Text © 2021 E. A. Ivanova published in Forest Science Issues Vol. 4, No. 3,  Article 87 E. A. Ivanova Institute of North Industrial Ecology Problems KSC RAS Akademgorodok st. 14a, Apatity, Murmansk&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/04/5-1-2022-Ivanova.pdf"><img loading="lazy" class="size-full wp-image-1122 alignright" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></p>
<div style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; font-size: 10pt; color: #000000;"><span lang="EN-US">Original Russian Text © 2021 E. A. Ivanova published in Forest Science Issues Vol. 4, No. 3,  </span><a style="color: #000000;" href="https://jfsi.ru/4-3-2021-ivanova/" target="_blank" rel="noopener noreferrer">Article 87</a></span></div>
<div style="text-align: justify;">
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>E. A. Ivanova</strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Institute of North Industrial Ecology Problems KSC RAS</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Akademgorodok st. 14a, Apatity, Murmansk region, 184209, Russia </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">E-mail: <u>ea.ivanova@ksc.ru</u></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 30 June 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 04 August 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 16 August 2021</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The paper provides an overview of Russian and foreign articles devoted to the study of the tree litter production and decomposition in forest ecosystems subjected to natural and anthropogenic factors. The spatial variability (below crown and between crown spaces) and the seasonal features of the tree litter production, its chemical composition, and decomposition processes are poorly studied. In addition, most of the works, both in Russia and foreign countries science, highlight the influence of natural factors on the litter production and the processes of its decomposition, while the impact of local sources of industrial air pollution is rarely considered. The study of the variability of the size, fractional and chemical composition and processes of decomposition of tree litter under conditions of industrial air pollution is important for predicting the dynamics of forest ecosystems subjected to the combined action of natural and anthropogenic factors and reducing the negative impact of production processes on forests.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Keywords: </strong><em>forest ecosystems, tree litter, industrial air pollution, fractional composition, chemical composition, litter decomposition, litter production and decomposition seasonal variability, litter production and decomposition spatial variability</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Tree litter in forest ecosystems acts as a link between plants of the upper tiers and the soil, as a source of soil organic matter and nutrients for biota and is one of the key components of biogeochemical cycles in forest biogeocenoses. Due to its chemical composition, tree litter participates in the formation of phytogenic zones of influence of trees, suppresses or accelerates the growth of herbaceous plants, affects microbial activity, soil composition (Aponte et al, 2013; Chavez-Vergara et al., 2014; Ufimtsev, Egorova, 2016; Kolmogorova, Ufimtsev, 2018; Pomogaibin E., Pomogaibin A., 2018), contributes to changes in the composition and abundance of soil microorganisms and invertebrates during the decomposition (Rakhleeva et al., 2011). Removal of leaf litter reduces the biological activity of the upper soil horizons, depletes the forest ecosystem of mineral nutrients, slows tree growth, and reduces soil respiration (Sayer, 2005; Xu et al., 2013; Ivanova et al., 2015), whereas addition of leaf litter reduces the temperature amplitude in the soil, increases nitrogen and aluminium availability (Loydi et al., 2014) and promotes higher methane production rates (Yavitt, Williams, 2015). The adding of litter in tropical forests increased the input of nitrogen and phosphorus into the soil (Wood et al., 2009), increased the concentration of nitrates and the stocks of inorganic nitrogen in the soil (Sayer, Tanner, 2010). The forest floor formed from undecomposed plant litter acts as a physical barrier to the emergence of shoots in species with small seeds, promotes the emergence and establishment of large-seeded species, maintains a microclimate favourable for herbivores and pathogens, acting as their habitat (Sayer, 2005; Dupuy, Chazdon, 2008).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Quantitative and qualitative characteristics of tree litter are of practical interest for assessing the level of radiation pollution (Bondareva, Rubailo, 2016; Komissarov, Ogura, 2017), fire danger based on the accumulation of combustible materials (Arkhipov, 2014; Sobachkin et al., 2017), accumulation of heavy metals by tree plants in urbanized areas (Kopylova, 2012). The use of tree litter debris, mostly coniferous or foliar, as a source of cellulose (Danilova, Stepanova, 2017), sorption material (Alekseeva, Stepanova, 2015; Silaicheva, Stepanova, 2016; Shaimardanova et al., 2017; Sverguzova et al., 2017), calcium fertilizer (Petrochenko et al., 2015) is being actively investigated. In mathematical studies, data on the input and decomposition of plant litter are used to form models for estimating the participation of litter in the biological cycle, the relationship with atmospheric CO<sub>2</sub> and climate (Brovkin et al., 2012; Mironenko, 2017). In particular, data on the foliar litter dynamics of evergreen tropical forests in Panama, French Guiana, and Brazil were used to modify a global terrestrial ecosystem model to allow a more accurate estimate of gross primary productivity (De Weirdt et al., 2012). The elemental composition of tree litter is of interest for understanding the patterns of element cycles and soil formation (Meier et al., 2005; Wood et al., 2006; Wood et al., 2009; Vesterdal et al., 2012; Osipov, 2017).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The parameters of tree litter are studied mainly in background conditions, unaffected by industrial air pollution from large industrial plants or thermal power plants. Air pollution is known to cause degradation of forest ecosystems, changes in forest stand structure: death of conifers and their replacement by small-leaved species (Chernenkova et al., 2016), reduction of species diversity and adaptability of plant communities, death of mosses and lichens (Salemaa et al., 2004; Hale, Robertson, 2016). Acid-forming substances and heavy metals — components of emissions — cause damage to assimilating organs of coniferous woody plants (Lukina, Nikonov, 1998; Yarmishko, Lyanguzova, 2013), reduced life span of fir-needles (Lamppu, Huttunen, 2003), almost no seed production of trees and shrubs (Tsvetkov V., Tsvetkov I. 2012). At the same time, a decrease in the activity of soil microorganisms and changes in the number of micromycetes and soil invertebrates is observed in forest ecosystems, resulting in slower decomposition of organic matter and increased thickness of forest floor (Nieminen et al., 1999; Zenkova, 2000; Polyanskaya et al., 2001; Nikonov et al., 2001; Fomicheva et al., 2006; Lukina et al., 2008; Vorobeichik and Pishchulin, 2009, 2016). The accumulation of Cu and Ni in forest soils near metallurgical plants leads to a deficit of basic cations (exchangeable Ca, Mg, K) in the organic layer (Derome, Lindroos, 1998). Even when the technogenic load is reduced, the death of the stand continues (Vorobeichik et al., 2014), the value of radial growth of trees in the pollution zone remains significantly lower than the control and background values (Chernenkova et al., 2012). In the vicinity of the Severonickel Combine, coniferous forests remain in a critical condition despite a reduction in emissions (Chernenkova et al., 2011; Lyanguzova et al., 2018). In this regard, the study of the processes of formation and decomposition of tree litter, as one of the key links in biogeochemical cycles, is of particular interest for understanding the dynamics of the functioning of forest ecosystems under changing man-induced loads.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The aim of this paper is to review the current state of research on the formation and decomposition of tree litter in forest ecosystems.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>METHODOLOGICAL APPROACHES TO THE STUDY OF THE CHARACTERISTICS OF TREE LITTER AND ITS DECOMPOSITION PROCESSES</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) has developed a detailed toolkit for participants on tree litter sampling and analysis (Ukonmaanaho et al., 2016). The translated guidelines for integrated monitoring, including the methods of the ICP Forests programme, provide recommendations for both the selection of litter and the assessment of decomposition rate (Guide…, 2013). Major works analyse the methods and results of various experiments in boreal and low-disturbed forests (Berg, McClaugherty, 2008) and describe the most common and specific methods (ecological, chemical, microbiological, etc.) for studying plant litter decomposition processes comprehensively (Methods&#8230;, 2005). Study of the literature showed that field research methods vary considerably, depending on the climatic features of the area, the composition of the stand and the research objectives.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">A number of modern papers present already known and proven methods for the sampling of tree litter. The most commonly used litter traps are box (preferably with a mesh bottom for water drainage) and collecting funnel located 1–1.5 m above the ground. For sampling the material directly from the ground surface, templates of different sizes are used (Table). In addition, litter is sometimes collected from the surface and without the use of a template from survey plots (Boldeskul et al., 2015; Ufimtsev, Egorova, 2016; Kolmogorova, Ufimtsev, 2018). Preston et al. (2006) simultaneously used different designs for collecting litter: plastic containers 27.3 cm in diameter and 30 cm high with a mesh bottom and 1 m<sup>2</sup> square nets laid on the forest floor to capture the branches.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Table. Examples of the most commonly used types of tree litter collection equipment</span></p>
<div style="overflow-x: auto;">
<table style="border: 1px #f1f1f1 solid; background-color: #ffffff;">
<tbody>
<tr>
<td width="121"><span style="font-family: 'times new roman', times, serif; color: #000000;">Design</span></td>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">Size</span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Samples</span></td>
</tr>
<tr>
<td rowspan="4" width="121"><span style="font-family: 'times new roman', times, serif; color: #000000;">Box</span></td>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">0.98 m<sup>2</sup></span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Bazilevich et al., 1978</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Bryanin, Abramova, 2017</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Abramova et al., 2018</span></td>
</tr>
<tr>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">1 m<sup>2</sup></span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Rodin et al., 1967</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Ermakova, 2009</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Boev et al., 2018</span></td>
</tr>
<tr>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">50 × 50 cm</span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Likhanova, 2014</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Osipov, 2017</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Yusupov et al., 1995</span></td>
</tr>
<tr>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">80 × 80 cm</span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Kopáček et al., 2010</span></td>
</tr>
<tr>
<td width="121"><span style="font-family: 'times new roman', times, serif; color: #000000;">Funnel</span></td>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">0.2-0.5 m<sup>2</sup></span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Ukonmaanaho et al., 2008</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Kouki, Hokkanen, 1992</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Jonczak, Parzych, 2014</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Berg et al., 1999</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Stojnić et al., 2019</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Ivanova, Lukina, 2017</span></td>
</tr>
<tr>
<td rowspan="3" width="121"><span style="font-family: 'times new roman', times, serif; color: #000000;">Template</span></td>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">0.031 m<sup>2</sup></span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Reshetnikova, 2011</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Vedrova, Reshetnikova, 2014</span></td>
</tr>
<tr>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">100 × 100 cm</span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Bessonova et al., 2017</span></td>
</tr>
<tr>
<td width="123"><span style="font-family: 'times new roman', times, serif; color: #000000;">0.25 m<sup>2</sup></span></td>
<td width="223"><span style="font-family: 'times new roman', times, serif; color: #000000;">Nakazato et al., 2021</span></td>
</tr>
</tbody>
</table>
</div>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The spatial variability of tree litter inflow to the soil surface is rarely considered. There are options for arranging the equipment in a non-random manner, but without specifying details (Ukonmaanaho et al., 2008); randomly (Dearden et al., 2006; Novák et al., 2014; Boev et al., 2018); evenly across the site (Yusupov et al., 1995; Stojnić et al, 2019); diagonally in sites (Albrektson, 1988); in a straight line 10 m apart (Michopoulos et al., 2020); in two lines (Meier et al., 2005; Ermakova, 2009); in a uniform grid on the site (Jonczak and Parzych, 2014); in different parts of the slope (Wood et al., 2006; Bessonova et al., 2017). A detailed study of the influence of stand structure is expressed in the distribution of equipment by parcel type (Lukina, Nikonov, 1996), below the crowns/between the crowns (Ivanova, Lukina, 2017). Tsandekova O. L. (2018), investigating the dynamics of ash accumulation in <em>Acer negundo</em> litter, conducted sampling at sampling sites in different conditions of crown density, taking into account the influence zones of trees: in sparse stands and in stands with crown closure of 50–60%, litter was collected in the subcrown and near-front zones, and in stands with crown density of 100% — in the near-trunk and inter-crown zones. Similarly, in the study of the chemical composition of Scots pine litter, samples were taken in the under-crown, near-front (inter-crown), and outer zones on sampling sites in sparse (open stand), thin and dense forest stands (Kolmogorova and Ufimtsev, 2018).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The frequency of material sampling depends largely on the climatic zone in which the study is carried out, and can be done monthly/every 2 weeks during the warm period and once during the winter (Yusupov et al., 1995; Ukonmaanaho et al., 2008; Lenthonen et al., 2008; Ťupek et al., 2015; Bryanin and Abramova, 2017); monthly during the growing season without sampling for the winter period (Shpakovskaya and Rozhak, 2014); every two weeks (Wood et al., 2009); only in spring and autumn (Likhanova, 2014; Novák et al., 2014; Ivanova and Lukina, 2017); three times a year: in spring, late summer and autumn (Kopáček et al., 2010); or once in autumn (Boldeskul et al., 2015; Boev et al., 2018). In tropical forests, sampling periods counted in days (De Weirdt et al., 2012). In large-scale studies covering sites in different climates, sampling frequencies can vary from 3 to 12 times per year depending on site location (Berg et al., 1999; Berg, Meentemeyer, 2001).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">In order to study the tree litter fraction composition, the plant material is sorted after sampling. Depending on the purpose of the study, only two parts could be distinguished in the litter: the fir-needles and a mixed fraction consisting of all other collected components (seeds, cones, bark etc.) (Berg et al., 1999; Berg, Meentemeyer, 2001). The litter was also divided into fractions: green needles, perennial needles and remaining fractions, while increasing the number of sampling periods (Ukonmaanaho et al., 2008). When sampling was from the ground surface, in addition to tree litter fractions (needles/leaves, bark, twigs and cones), dwarf shrub litter, moss litter, lichens, grasses could be identified (Yusupov et al., 1995; Preston et al., 2006; Reshetnikova, 2011; Hilli, 2013; Sobachkin et al., 2017; etc.). There are also papers describing an even more thorough accounting of the tree litter fractions, distinguishing fruit, buds, seeds and ament and other fractions (Ermakova, 2009; Shpakovskaya, Rozhak, 2014; Ivanova, Lukina, 2017). In terms of the study of tree litter as the L horizon of the forest floor, it is also possible to divide the plant material into active (leaves, needles, chaff, seeds) and inactive (cones, small branches, bark) fractions (Karpachevskii et al., 1980).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Studies of litter characteristics are most often devoted to a rather short observation period — up to 4-5 years, but there are also long-term studies. Ťupek et al. (2015) used data from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) from 1996 to 2011 and studies by the Finnish Forest Research Institute (Metla) from 1960 to 2010. Lenthonen et al. (2008) studied time series for needle litter, tree growth, microstrobila (male pollinic cone) litter and daily weather data for 43 years from 1961 to 2004. In south-eastern Finland, Scots pine needle litter had been sampled over a period of 24 years (1962–1986) (Kouki and Hokkanen, 1992). Long-term observations of tree litter parameters are valuable for understanding the functioning of forest ecosystems in response to climate change.</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Litter decomposition</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The study of litter decomposition processes is mainly based on field experiments on incubating plant samples of active litter fractions (needles/leaves) directly in stands for periods ranging from 2-3 (Rakhleeva et al., 2011; Likhanova, 2014) to 4-6 years (Moore et al., 2006; Symonds et al., 2013). There are also studies covering just one season (Abramova et al., 2018). The frequency of sampling varies depending on the objectives of the study, climatic features of the area. When the initial decomposition stages were examined in more detail, the intervals ranged from a few days to a month (De Marco et al., 2007; Wood et al., 2009; Abramova et al., 2018). For the overall assessment of litter mass loss and changes in its chemical parameters, the periods ranged from a few months (Rakhleeva et al., 2011) and half a year (Aponte et al., 2013) to a year.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">In the described incubation experiments litter samples were deposited for decomposition in mesh bags made of inert material: nylon (Rakhleeva et al., 2011), plastic (Wood et al, 2009), kapron (Likhanova, 2014), glass fibre (Ogden and Schmidt, 1997), polypropylene (Moore et al., 2006), terilene (polyethylene terephtalate) (Berg et al., 1993) with different hole sizes, which depended on the size of plant residues. For example, leaf litter from oak, birch, robinia, aspen and pine were placed in 1.5 × 1.5 mm mesh bags, while for douglas-fir litter the bags had 1 × 0.6 mm mesh size to avoid sample loss (Van Nevel et al., 2014). In studies of invertebrate participation in litter decomposition, it is possible to use a combined option where the lower part of the bag uses a mesh with smaller cells (0.5–1 mm) and the upper part with larger cells (0.2–1 cm) for biota access (Wood et al., 2009; Rakhleeva et al., 2011; Slade, Riutta, 2012). In addition to cloth bags, there are mentions of the use of 100 × 100 × 5 cm containers (De Marco et al., 2007).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The spatial patterns of litter decomposition and changes in its chemical composition are as little studied as its quantitative characteristics. An example of a study of tree influence zones can be found in the work of E. L. Vorobeichik and R. G. Pishchulin (2011), which studied the decomposition of pure cellulose in the near-trunk areas (at 0.2–0.4 m from the trunk), in the middle of crown projection (1.2–1.8 m), in the canopy gaps (3.8–5.3 m) and on the opposite side from the gap under a closed forest canopy (2–3 m from the trunk). The influence of the stand structure was also assessed when the samples were located below the crowns/between the crowns of spruce and pine forests (Lukina et al., 2017, Ivanova et al., 2019).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>AMOUNT AND FRACTIONAL COMPOSITION OF TREE LITTER:</strong></span><br />
<span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>NATURAL FACTORS AND INDUSTRIAL AIR POLLUTION</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The quantity and quality of tree litter regulates carbon accumulation, element cycles in forests. Within the framework of the International Biological Program, work has been carried out to assess the weight and fractional composition of litter in taiga forests of European Russia (Kazimirov &#038; Morozova 1973; Zaboeva 1975; Manakov &#038; Nikonov 1981). Numerous long-term observations of plant litter volumes in the same years were carried out abroad (Bray and Gorham, 1964; Flower-Ellis, 1985; Kouki and Hokkanen, 1992).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Influence of natural factors on the formation of tree litter</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Among the natural factors affecting tree litter formation, there is a dependence of coniferous litter mass on geographic latitude: among sites of similar fertility, the amount of coniferous litter is lower for sites located in the north (Albrektson, 1988; Berg et al., 1999). The size and composition of the litter depends on the composition of the forest stand (Shpakovskaya and Rozhak, 2014), the annual growth of trees, and their age (Pedersen and Bille-Hansen, 1999). In particular, litter mass (needles, cones and bark plus dead grass) in a mature pine forest exceeded its for a middle-aged plantation (Sobachkin et al., 2017). In recent decades, relationships between above-ground tree biomass and litter have been assessed based on long-term monitoring data (Lenthonen et al., 2008; Ukonmaanaho et al., 2008; Ilvesniemi et al., 2009; Novák et al., 2014). Thus, in the pine forests of northern Finland, the Scots pine needles litter depends on the production and mass development of needles, which occurs 4-6 years earlier (Lenthonen et al., 2008). The amount of litter may depend on weather conditions: unfavorable climatic factors, cooling, and lack of precipitation inhibit the development of plant leaf apparatus (Likhanova, 2014). A long-term study in south-eastern Finland showed a positive association of Scots pine needle litter with mean July temperature and high temperatures between March and April: the high temperature in July coincided with an increase in litter amount in the same year and the following year (Kouki and Hokkanen, 1992). In young oak stands, a positive relationship was found between annual litter and precipitation and a negative relationship with summer temperature (Novák et al., 2014). A high yield of pine seeds and cones is associated with the warm weather of previous years (Nekrasova, 1957).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Litter size exhibits species specificity, and an increase in species diversity leads to an increase in litter production (Scherer-Lorenzen et al., 2007). The average annual litter in the spruce forests of Finland was higher than that of pine forests (Ukonmaanaho et al., 2008); in the area with Siberian stone pine cultivation, there was almost three times more litter than in the spruce forest (Reshetnikova, 2011). Differences in the amount of litter fall of Hudson Bay pine (<em>Pinus banksiana</em>) and black spruce (<em>Picea mariana</em>) growing along the Boreal forest transect in northern Canada have been attributed to site conditions (soil structure and drainage) and forest floor thickness (Preston et al., 2006).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The activity of insect pests leads to changes in the litter production: in mature spruce stands in the Lake Plešné catchment (Czech Republic), the amount of litter increased after bark beetle infestation (Kopáček et al., 2015).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The fractional composition of tree litter, as well as its total mass, may depend on the age or species composition of the stand. In forb-green-moss pine forest of various ages in the forest-steppe zone cones and needles predominated in the fractional composition of litter. In the middle-aged stand, the needles accounted for most of the litter (52.2%), while in the mature stand the share of needles decreased to 36.7%. The participation of cones in litter is also explained by age differences in stands: in the litter of a mature stand, it is higher than in a mid-age stand (Sobachkin et al., 2017). In 40-year-old stands of the main forest-forming species of Siberia under cedar about 90% of the litter mass is needles, under pine, larch and spruce 40–50% are needles and 20–45% are branches (d ≤ 10 mm). In birch and aspen forests 70–74% of the litter mass is represented by leaves and 21–29% — by branches (Reshetnikova, 2011). High values of the mass of needle, bark, and branch litter can also be caused by the action of dangerous weather phenomena — strong winds and snowstorms (Report&#8230;, 2015).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Seasonal and spatial variability of tree litter formation</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Tree litter formation processes (its mass and fractional composition) in seasonal dynamics and depending on forest canopy pattern structure are poorly studied. It is known that in spruce-beech and fir-spruce-beech forests of the Ukrainian Carpathians, most of the annual amount of litter occurs in October and November due to an increase in needles and leaves part in its composition. At the same time, the dynamics of needle supply has the form of a curve with two peaks at the beginning and at the end of the growing season and with a minimum at the beginning of autumn (Shpakovskaya and Rozhak, 2014). In a burnt forest and in control forest, the maximum inflow of litter occurs in October and the minimum — in July (Bryanin, Abramova, 2017). The bulk of the leaves of the flat-leaved birch in the forest stands of the Khamar-Daban ridge (Southern Baikal region) begins to fall at the end of the second decade of September, with a maximum rate during the third decade of September — the first pentad of October, and the bulk of the leaves of the Siberian mountain ash fall within 3–4 weeks — from the beginning of the second decade of September until the end of the first decade of October (Ermakova, 2009). In medium taiga spruce forests, the winter-spring period accounts for 52–58%, the summer period for 20–23%, and the autumn period for 22–25% of the total mass of litter (Likhanova, 2014). In 40-year-old coniferous stands of Siberia, the mass of litter in the summer-autumn period is higher than in winter (Reshetnikova, 2011). As part of a study of the role of litter in the formation of phytogenic tree fields in coal mine dumps, differences in the fractional composition of the litter horizon L were found in different zones: needles and cones dominated under the crowns, while in the outer zone the litter consisted almost entirely of meadow vegetation litter (Ufimtsev, Egorova, 2016).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Anthropogenic factors affecting the formation of litter</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Fires as an anthropogenic factor lead to significant changes in the functioning of forest ecosystems. In the post-fire larch forest in the foothills of the Tukuringa Ridge (Upper Amur Region), the input of litter from the aboveground part of the vegetation was reduced by 2.8 times compared with the control forest. In addition, the fractional composition was also characterized by differences: in the control forest, there was a gradual decrease in the fractional composition of the total amount of litter in the series leaves — fir — needles — branches — grass — other fractions (33%, 26%, 21%, 11%, 9% respectively), whereas in the post-fire stand, grass litter was dominating and the fractional decrease occurred in reverse order: grass — other fractions — fir — needles — leaves — branches (28%, 23%, 22%, 20%, 7%, respectively) (Bryanin, Abramova, 2017). With a prolonged non-fire period, the branch/tree ratio in the litter increases due to reduced needle litter, resulting in a reduced decomposition rate (Dearden et al., 2006).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Air pollution with heavy metals and acid-forming substances leads to damage to the assimilating organs of coniferous woody plants and a decrease in the life span of needles — defoliation of trees is not only in phenological terms, contributing to an increase in the amount of litter (Nieminen, Helmisaari, 1996; Rautio et al., 1998; Lukina, Nikonov, 1998; Lamppu and Huttunen, 2004; Nikonov et al., 2004; Yarmishko and Lyanguzova, 2013). As pollution levels increase, the number of female cones per tree decreases (Stavrova, 1990), the proportion of large cones decreases, the number of damaged and diseased cones increases, cone diameter and average raw weight decrease (Tsvetkov V., Tsvetkov I., 2003). The proportion of epiphytic lichens in litter as an element of biogeocenosis sensitive to air pollution decreases in the area affected by Severonickel Combine emissions. At the same time, there are clear trends towards an increase in the total mass of litter due to pine needles and bark, despite a decrease in emissions over 20 years, which may be due to weakening of trees and premature die-off of individual organs (Ivanova, Lukina, 2017).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FEATURES OF THE CHEMICAL COMPOSITION OF TREE LITTER</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The chemical composition of fresh tree litter determines its quality for the decomposing organisms and consequently influences the rate of decomposition and the change in the chemical composition of the plant residues during the mineralisation process. Thus, the work of N. V. Likhanova (2014) showed that birch leaves litter decomposition was the most intensive, where the С : N ratio was 35–38, while this ratio varied from 38 to 43 for spruce and pine needles, from 43 to 60 for tree branches, and from 105 to 142 for bark. The low content of nitrogen and phosphorus in the needles leads to an increase in the С : N ratio, which increases the chance of nitrogen immobilization during the early stages of decomposition (Symonds et al., 2013).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Natural factors determining the chemical composition of tree litter</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Both the quantitative characteristics and the chemical composition of plant residues depend on various factors. Concentrations of Mg, N and K were found to decrease with increasing age of beech stands (Trap et al., 2013). In the artificial plantation of black locust, the N, K, Mg, P input with litter was higher in the lower third of the studied slope of Voyskovoye gully (Bessonova et al., 2017). Ca concentration in needle and leaf litter was negatively associated with annual precipitation, probably due to washout by rain and melting snow (Berg et al., 2017).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Numerous works have shown that the content of elements in the litter depends on the tree species (Preston et al., 2006; Ukonmaanaho et al., 2008; Aponte et al., 2013; Jonczak and Parzych, 2014; Boev et al., 2018; Neumann et al., 2018; Becker et al., 2018). Norway spruce (<em>Picea abies</em>) and lodgepole pine (<em>Pinus contorta</em>) needles litter contain more calcium than Scots pine needles litter (<em>Pinus sylvestris</em>). In addition, Ca concentrations in fresh litter are positively related to P, K, and Mg concentrations: for pine species (<em>Pinus contorta</em> and<em> Pinus sylvestris</em>) Ca content was positively related to Mg and Mn concentrations, for Scots pine (<em>Pinus sylvestris</em>) — with Mg content (Berg et al., 2017). In undisturbed 40-year-old Siberian stands, the carbon to nitrogen ratio in cedar litter was 101, in pine was 98, in larch and spruce — 87, in birch — 76 and in aspen — 118 (Reshetnikova, 2011).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The fauna also introduces changes in the chemical composition of plant residues. In the needles falling off after the forest desiccation due to infection of the forest with bark beetles, the concentration of N increased for 1–3 years, and the C : N and C : P ration decreased, indicating decomposition by endophytes already on the trees. At the same time, the concentrations of Mg, K, and P increased in the total litter due to an increase in the proportion of rowan litter (Kopáček et al., 2015).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Seasonal and spatial variability of the chemical composition of tree litter</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Seasonal and spatial features of the chemical composition of the litter, both in Russia and abroad, have been studied rather poorly. The larch litter taken in spring was 10% enriched in N and 40% depleted in Ca compared to the litter taken in autumn. Changes in lignin : N, C : N and C : P ratios after the winter season indicated the beginning of litter decomposition (Chuldiene, 2017). According to other data, the nitrogen content in the total pine litter in the conditions of rockfall increased uniformly during the growing season (Kolmogorova, Ufimtsev, 2018). In Finnish forests, there were two main periods when C and N were deposited in the ground: May–October and November–April, with higher depositions in the first period, peaking in September (Portillo-Estrada et al., 2013). In pine forests in Poland, the Mn, Zn and Ni content in pine needles in 2007 was shown to be higher in autumn, whereas in 2009 it was higher in spring (Jonczak and Parzych, 2014). Under the conditions of the rock dump (in the reclaimed areas of opencast coal mine overburden), the content of total phosphorus in the Scots pine litter reached a maximum in the sub-crown and near-front zones of the dense stands (Kolmogorova, Ufimtsev, 2018). In the<em> Acer negundo</em> litter, greatest accumulation of the ash component occurs in the under-crown and near-trunk zones of single trees in sparse stands compared to other groups of trees and with control forest (Tsandekova, 2018).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Changes in the chemical composition of tree litter caused by anthropogenic factors</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Sharp changes in the functioning of forest ecosystems caused by anthropogenic factors significantly affect the chemical composition of tree litter. In the post-pyrogenic larch forest in the foothills of the Tukuringa Range, the litter is dominated by organic remains enriched in nitrogen but poor in carbon. In larch needles on the control sample plot, the C : N ration approaches 170; in a stand damaged by fire, its does not exceed 110 (Bryanin, Abramova, 2017).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Atmospheric pollution leads to disruption of the processes of retranslocation of elements within trees (Lukina and Nikonov 1996, 1998; Nieminen and Helmisaari, 1996; Rautio et al, 1998; Steinnes et al, 2000; Kiikkilä, 2003; Tarkhanov, 2009; Yarmishko and Lyanguzova, 2013; Sukhareva and Lukina, 2014; Vacek et al., 2016). In the impact zone of the Middle Ural copper smelter compared to the control zone, more Ca was supplied with pine needle litter (Yusupov et al., 1995). The long-term effect of acid precipitation and nitrogen saturation in Czech spruce forests has caused a decrease in Ca, Mg, and Mn concentrations and Ca : Al and Mg : Al ratios, increase in N content and N : Mg ratio in the litter (Kopáček et al., 2010). In defoliating forests and pollution-induced sparse forests in the area of the Severonickel Combine, a deterioration in the quality of plant material was recorded: an increase in heavy metals Ni and Cu and a decrease in Ca, Mn, K, Mg (Lukina et al., 2017; Ivanova et al., 2019), and lignin content increased in the birch leaf litter when approaching the combine (Artemkina, 2018).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>DECOMPOSITION OF TREE LITTER IN FOREST ECOSYSTEMS</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The evaluation of litter decomposition processes is reflected in numerous works from all over the world. The rate of plant residue mass loss and changes in chemical composition are influenced by various environmental factors: stand composition, soil conditions, weather, microbial activity, etc. (Fig. 1). The current concept is that litter quality is the dominant factor at large spatial scales, and the activity of the decomposing organisms is regulated by climate and litter quality (Bradford et al., 2016).</span></p>
<div id="attachment_5001" style="width: 1034px" class="wp-caption aligncenter"><img aria-describedby="caption-attachment-5001" loading="lazy" class="size-large wp-image-5001" src="https://jfsi.ru/wp-content/uploads/2022/04/Figure-1.-1024x597.jpg" alt="" width="1024" height="597" srcset="https://jfsi.ru/wp-content/uploads/2022/04/Figure-1.-1024x597.jpg 1024w, https://jfsi.ru/wp-content/uploads/2022/04/Figure-1.-300x175.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/04/Figure-1.-150x87.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/04/Figure-1.-768x448.jpg 768w, https://jfsi.ru/wp-content/uploads/2022/04/Figure-1..jpg 1184w" sizes="(max-width: 1024px) 100vw, 1024px" /><p id="caption-attachment-5001" class="wp-caption-text"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Figure 1.</strong> Factors influencing the litter decomposition processes</span><br /><span style="font-family: 'times new roman', times, serif; color: #000000;">(acc. to Krishna, Mohan, 2017)</span></p></div>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Influence of natural factors on tree litter decomposition processes</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">One of the main factors affecting the rate of decomposition is the activity of soil biota: invertebrates, microorganisms, and fungi (Vorob’eva, Naumova, 2009). Litter decomposition in the most surface soil horizon is attributed to the predominance of saprotrophic fungi and the absence of mycorrhizal fungi (Högberg et al., 2017). However, the larger soil fauna is also influential. Slade, Riutta (2012) showed that macrofauna accounted for 22–41% of the total mass loss of leaf litter. Earthworms increased mass loss of litter with lower C : N (Belote and Jones, 2009). In a laboratory experiment, high concentrations of Cd, affecting earthworm activity, inhibit leaf litter decomposition and lead to a decrease in soil fertility (Liu et al., 2020).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The mineralisation of tree litter depends on the hydrothermal conditions of soils (Kuznetsov, 2010; Kuznetsov, Osipov, 2011) and is positively related to mean annual temperature and annual precipitation (Albrektson, 1988; Pausas, 1997; Portillo-Estrada et al., 2016). In Mediterranean sites, the decomposition of senescent pine needles was faster than in continental forests of the Pyrenees, and a sharper reduction in decomposition rates was observed there when stands were thinned (Blanco et al., 2011). A study on the effect of altitude on the decomposition of plant residues revealed that decomposition processes are mainly influenced by the quality of the litter. These processes do not depend so much on altitude but rather on a combination of specific conditions such as temperature, precipitation, different types of forest floor and different trophic interactions between the plants and the microbial community (Marian et al., 2017). The decomposition rate of thin woody residues (branches of different diameters) increased from north to south in a large-scale study with plots along a climatic gradient from Northern Finland to Central Estonia (Vavrova et al., 2009).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Since the fractional and chemical composition of the litter depends on the species composition of the forest stand, the process of its decomposition has corresponding features. It has been demonstrated that the dynamics of the content of elements during the decomposition of litter on Mount Vesuvius in four different pine species (<em>Pinus pinea, P. laricio, P. sylvestris</em> and<em> P. nigra</em>) is mainly governed by their original content. For example,<em> P. nigra </em>litter, the richest in nitrogen, released N during decomposition. Potassium was accumulating in <em>P. sylvestris</em> litter, while Mn was accumulating in<em> P. nigra</em> and<em> P. pinea</em> litter, which had the lowest initial concentrations of K and Mn, respectively (De Marco et al., 2007). Spruce needle litter, characterized by a higher content of nutrients and narrower C : N and lignin : N ratios, within two years was decomposing noticeably faster than pine needle litter. Wherein the litter of birch leaf (<em>Betula pendula</em>), growing in pine forests and characterized by a lower N : P ratio, decomposes faster than downy birch litter (<em>B. pubescens</em>) in spruce forests (Ivanova et al., 2019). In coastal forests of British Columbia, grape maple litter with higher concentrations of N, P, Ca, Mg, K, Fe and Zn was decomposing significantly faster than conifer litter (Ogden and Schmidt, 1997). However, the rate of degradation of pure cellulose is higher in spruce-fir forests than in birch forests (Vorobeichik and Pishchulin, 2011). Subordinate foliage plants with sharply contrasting feeding and water yield characteristics compared to the dominant evergreen plants significantly influenced litter decomposition at the community level, despite their low abundance (Guo et al., 2020).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">A number of experiments in different types of terrestrial ecosystems have shown: the decomposition process depends on the fractional composition of the incoming litter (Bobkova, 2000; Fang et al., 2015). In undisturbed of 40-year stands in Siberia, woody species that annually shed their leaves (needles): larch, aspen and birch (Reshetnikova, 2011; Vedrova, Reshetnikova, 2014) are characterized by the maximum mass loss in the annual cycle of decomposition.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Many studies have shown the influence of the initial quality of the litter, determined by concentrations of nutrient, heavy metals, element ratios, on the decomposition rate (Berg, 2000; Wardle et al., 2003; De Marco et al., 2007; Zhang et al.,2008; Berg, McClaugherty, 2008; Rahman et al., 2013; Tu et al., 2014; Lukina et al., 2017; Ivanova et al., 2019). Litter with a higher nitrogen content decomposes faster than those with low nitrogen and high lignin concentrations (Wardle et al., 2003). Accordingly, the stoichiometric C : N and lignin : N ratios in plant residues have a significant effect on decomposition: the narrower these ratios, the higher the rate of decomposition (Berg and McClaugherty, 2008; Lukina et al., 2017; Ivanova et al., 2019). At the early stages of decomposition, nitrogen has a stimulating effect, while at later stages, on the contrary, it inhibits the decomposition rate, while Ca and Mn have a significant positive effect (Berg, 2000; Berg and Meentemeyer, 2001; Davey et al., 2007; Berg, 2014 ). Some authors have studied an excessive intake of one element, most commonly nitrogen. Tu et al. (2014) found that high nitrogen input reduced the rate of decomposition in forests, and the mass of undecomposed litter was closely related to residual lignin during the decomposition process. During the early stages of decomposition, nutrients such as nitrogen and phosphorus as well as water-soluble organic compounds have the greatest effects, whereas in the later stages lignin is the main determinant of decomposition dynamics (Rahman et al., 2013). Mineral N application or mixing of litter of different quality, expressed in C : N ratio and N content, increased the intensity of mineralization of N-poor litter fractions and inhibited the release of CO<sub>2</sub> during the decomposition of N-rich litter (Bonanomi et al., 2014; Larionova et al., 2017). Under the conditions of the incubation experiment, sodium chloride and sodium sulphate exhibited an inhibitory effect on the biota involved in the decomposition of birch litter, while, in contrast, when the litter was treated with solutions of iron salts, an increase in the mineralising activity of the biota was observed (Smirnova et al., 2017).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Changes in the chemical composition of plant residues during decomposition</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">In the process of mineralization of plant material, changes in the chemical composition are observed. Pine and spruce needle litter at the initial stages of decomposition (up to 165 days) releases monoterpene hydrocarbons in the gas phase at a rate comparable to emissions from living needles of these trees (Isidorov et al., 2010). In undisturbed 40-year-old Siberian stands, as plant residues decompose, their carbon, P and K contents decrease and Mg concentration increases (Reshetnikova, 2011). Changes in the content of elements can be interrelated. In the subarctic to cool-temperate highlands of Canada, the decomposition of assimilating tree organs litter usually retained N in the decomposing litter until about 50% of the initial C remained. Peak N content in litter was observed to be between 72% and 99% of the original remaining C with C : N ratios ranging from 37 to 71. The rate of phosphorus loss inversely correlated with the initial concentration of phosphorus in the litter, which varied from 0.02% to 0.13%. There was a trend toward higher nitrogen and phosphorus retention during litter decomposition at sites with lower C : N and N : P ratios, respectively (Moore et al., 2006). As the leaf/coniferous litter decomposes, an increase in Ca concentration is shown, often followed by a decrease. The maximum calcium concentrations are positively related to manganese and negatively related to nitrogen, which can have a direct influence on the decomposition rate (Berg et al., 2017).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Seasonal and spatial variability of tree litter decomposition</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Seasonal and spatial patterns of decomposition, in turn, depend largely on the activity of soil destructors and the influence of trees: in winter the process slows down considerably (Vorob’eva, Naumova, 2009). Litter mass losses in spruce and pine forests were higher between tree crowns compared to undercrown spaces (Lukina et al., 2017; Ivanova et al., 2019), but net cellulose degradation rates were higher in spruce-fir and birch forests under tree crowns compared to canopy gaps (Vorobeichik and Pishchulin, 2011).</span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Anthropogenic factors influencing the processes of decomposition of tree litter</em></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Forest management can change the rate of decomposition and cycling of elements. In beech and spruce forests with high intensity forest management, higher rates of litter decomposition and release of most nutrients were observed than in unmanaged deciduous forests (Purahong et al., 2014). In a plantation of Chinese pine (<em>Pinus tabulaeformis </em>Carriere), N during litter decomposition was accumulating until the ratio of acid-non-hydrolysable residues to nitrogen was reached 57–69. At the same time, thinning accelerated the decomposition of nitrogen-poor litter and also increased nitrogen accumulation (Chen et al., 2014). In 4–6-year old cutover patches after clear-cutting in medium taiga spruce forests, the highest decomposition rate was observed for birch leaves in the first year, while for spruce and pine needles an increase in decomposition rate was observed in the second year of the experiment. The tree litter components belonging to the inactive fraction (branches, bark, cones) were decomposing very slowly (Likhanova, 2014). In the post-fire larch forest (12 years after the fire), in the initial stages of decomposition, as well as in the control larch forest, the maximum losses were observed in the first 75 days of the experiment, and the decomposition rate in the studied forest ecosystems decreased in the series: grass–leaves–needle–branches (Abramova et al., 2018).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The extremely change in the litter decomposition processes is brought about by airborne industrial pollution. In the area affected by airborne emissions from the smelter, the proportion of poorly decomposed dead wood was higher than in the background area in the southern taiga, indicating a strong inhibition of tree residues degradation (Bergman, Vorobeichik, 2017). Soil contamination with heavy metals (Cu, Pb, Cd, Zn) reduced the rate of cellulose degradation in spruce-fir forests and birch forests by 2.7–5.4 times (Vorobeichik, Pishchulin, 2011). The use of tree-ring dating and an exponential decomposition model made it possible to determine that pollution had led to a decrease in the rate constant of wood decomposition by 16–60% (Dulya et al., 2019). In the Sudbury (Ontario) copper-nickel smelter impact area, a decrease in the rate of litter decomposition was observed (Freedman, Hutchinson, 1980). There, during the period of significant emission reductions, a decrease in the decomposition rate of white birch (<em>Betula papyrifera </em>Marshall) leaf litter was still observed and an increase in Cu and Ni in the litter was recorded, indicating that atmospheric inputs of Cu and Ni from Sudbury smelters remained high enough at the time of the 1999–2001 experiment to have a negative impact on degradation processes (Johnson, Hale, 2004). Scots pine needle litter 0.5 km from the Outukumpu copper smelter in the Harjavalta region in southwestern Finland had the lowest mass loss rate — 28.1%, while in the background it was 37.9% for the entire time. In addition, copper and nickel accumulation and a decrease in the carbon/nitrogen ratio have been observed in the impact zone over time (McEnroe and Helmisaari, 2001). In Belgium, in sandy soils contaminated with metals, a change in chemical composition during decomposition was observed: samples with initially low metal content were enriched in Cd and Zn, while metal losses were observed for samples with high content (Van Nevel et al., 2014).</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">In the vicinity of the Severonickel Combine near Monchegorsk, a decrease in the rate of decomposition of birch leaves was observed (Kozlov, Zvereva, 2015); in spruce and pine forests, a decrease in the rate of litter decomposition was noted, associated with a decrease in litter quality: increased initial content of heavy metals Ni and Cu, low content of nutrients and an increase in the lignin : N, C : N ration. In addition, during decomposition, plant residues in spruce and pine forests lost Ca, Mn, K, and Mg more intensively compared to the background and accumulated lignin, Al, Fe, Ni, and Cu (Lukina et al., 2017; Ivanova et al., 2019).</span></p>
<p>&nbsp;</p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">Tree litter acts as a link between tree vegetation and soil. Data on the content of elements in the litter allows us to estimate the amount of elements entering the soil and predict the rate of decomposition, during which the elements are released and re-engaged in biogeochemical cycles. In recent decades, factors affecting the litter formation and decomposition, taking into account the producing species, have been extensively investigated. Although the quantitative and qualitative characteristics of tree litter and its decomposition and mineralisation processes have been studied, the spatial and seasonal variability of these parameters and processes have been studied insufficiently. There are not enough studies devoted to the influence of local sources of air pollution — metallurgical complex enterprises, thermal power plants, nuclear power plants and others — on the tree litter. Understanding the processes of adaptation of forest ecosystems to climate change, the variability of ecosystem functions of forests requires research into the variability of size, fractional composition, chemical composition and decomposition processes of tree litter, taking into account seasonal and spatial variability (forest canopy pattern structure) under conditions of combined natural and anthropogenic factors, including atmospheric pollution. This will improve forecasts of further changes in forest ecosystems and develop recommendations for optimizing production processes to reduce the impact on forest ecosystems.</span></p>
<p>&nbsp;</p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FINANCING</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">The research was carried out under the State Assignment of the Institute of North Industrial Ecology Problems FRC KSC RAS No. 0226-2018-0111 (AAA-A18-118021490070-5) and partially under the Contract No D-1087.2021 “Integrated research of Russian Arctic forests to improve their productivity and preserve ecosystem functions”.</span></p>
<p>&nbsp;</p>
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<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p>&nbsp;</p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewer: </strong>Candidate of Biological Sciences F. I. Zemskov</span></p>
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		<title>INFLUENCE OF VEGETATION ON SOIL CARBON STOCKS IN FORESTS (REVIEW)</title>
		<link>https://jfsi.ru/en/5-1-2022-kuznetsov%d0%b0/</link>
		
		<dc:creator><![CDATA[lena]]></dc:creator>
		<pubDate>Thu, 31 Mar 2022 06:27:35 +0000</pubDate>
				<category><![CDATA[№1 2022]]></category>
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					<description><![CDATA[Original Russian Text © 2021 А. I. Kuznetsovа published in Forest Science Issues Vol. 4, No. 4, Article 95 A. I. Kuznetsovа Center for Forest Ecology and Productivity of the Russian Academy of Sciences, Profsoyuznaya st. 84/32 bldg.&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/04/2022-5-1-Kuznetsovа.pdf"><img loading="lazy" class="alignright wp-image-1122 size-full" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></span></p>
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<p><span style="font-family: 'times new roman', times, serif; font-size: 10pt; color: #000000;"><span lang="EN-US">Original Russian Text © 2021 </span>А<span lang="EN-US">. I. </span><span lang="EN-US">Kuznetsovа</span><span lang="EN-US"> published in Forest Science Issues Vol. 4, No. 4, <a style="color: #000000;" href="https://jfsi.ru/4-4-2021-kuznetsov%D0%B0/" target="_blank" rel="noopener noreferrer">Article 95</a></span></span></p>
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<div style="text-align: center;"><span style="color: #000000;"><strong style="font-family: 'times new roman', times, serif;">A. I. Kuznetsovа</strong></span></div>
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<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Center for Forest Ecology and Productivity of the Russian Academy of Sciences, </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Profsoyuznaya st. 84/32 bldg. 14, 117997, Moscow, Russia</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">E-mail: nasta472288813@yandex.ru</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 15 November 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 17 December 2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 18 December 2021</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Existing estimates of carbon stocks in taiga and coniferous-broadleaf forests show that almost half of the total organic carbon in these ecosystems is accumulated in forest soils. Vegetation as the main source of organic matter in the soil interacts with soil biota, which processes plant litter, and, with abiotic environmental factors, determines the processes of formation and accumulation of soil organic matter. Changes in the composition of vegetation are the driver of the dynamics of soil carbon stocks; however, insufficient attention has been paid to the analysis of this issue. This review analyzes the main ways of transferring carbon from the vegetation pool to the soil pool and the influence of three main predictors of vegetation that affect the carbon stock in soils: the amount and quality of litter of individual plant species (species identity) and the structural diversity of the plant community; it identifies gaps in knowledge and proposes some ways of developing this field of research.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><em>Keywords</em></strong><em>:</em> <em>vegetation, soil organic horizon, soil, carbon stock, litter quality, litter amount, interbiogeocenotic heterogeneity, intrabiogeocenotic heterogeneity</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Climate change is one of the current global challenges. Forests play a huge role in climate regulation due to their ability to absorb greenhouse gases and store carbon in both biomass and soils. The proportion of soil carbon in the total carbon stocks of forests is 40% or more (Framstad et al., 2013). Over the past 10 years alone, several meta-analyses have been conducted on a regional and global scale to identify the main factors of carbon accumulation in the soil.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">When assessing soil carbon stocks on a regional scale, the leading role of climate is highlighted (Wiesmeier et al., 2019). Local abiotic factors that have an impact on the level of carbon accumulation in soils include granulometric and chemical composition of soil-forming rocks, as well as topographic features of an area. Special focus is on the study of anthropogenic factors, that is, the influence of forestry regimes in the past and present, the influence of forest fires, and the history of forest development (Framstad et al., 2013; Mayer et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Vegetation, fauna and microbiota are the main biotic factors of soil carbon accumulation. The influence of biota on soil carbon stocks can be considered at different spatial levels, taking into account natural borders (forest catchments of different orders, forest type, etc.) and administrative (national, regional and local) boundaries.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">At high spatial (e. g., regional) levels, the relation between soil carbon stocks and climatic conditions was revealed: there are positive correlations with average annual temperature, average annual precipitation and, accordingly, net primary productivity (Amundson, 2001). Comparing different climatic zones showed a decrease in carbon stocks of the litter pool and an increase in carbon stocks in the mineral layers of the soil from north to south (Wiesmeier et al., 2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">At all spatial levels, the correlation with vegetation was found. Special emphasis was put on the influence vegetation has on organic matter decomposition in forest ecosystems (Prescott, 2010, Krishna, Mohan, 2017; Berg, McClaugherty, 2020, Ivanova, 2021). Some studies analyze stabilization processes of soil organic matter of microbial and plant origin (Angst et al., 2021) and dissolved organic carbon in forest soils (Karavanova, 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Vegetation composition determines the quantity and quality of incoming plant litterfall, its transformation and transition to soil pools with the active participation of decomposers, as well as migration processes of carbon compounds within the soil profile (Gleixner, 2013; Krishna, Mohan, 2017). For boreal and subboreal forests, differences in carbon stocks in forests of different types were shown (Jandl et al., 2007; Oostra et al., 2006; Schulp et al., 2008; Akkumuljacija&#8230;, 2018; Kuznetsova et al., 2019, 2020), including the contribution of not only the tree layer, but also the ground cover (Lukina et al., 2020; Kuznetsova et al., 2021). The question of the combined effect of different plant species on carbon stocks remains open.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A better understanding and assessment of the combined influence of various factors on carbon accumulation in forest soils is crucial for tackling tasks like development of measures for climate change mitigation and forecasting possible changes in ecosystem functions and services. Such estimates are few, although they are of vital scientific and applied importance, especially more recently, due to the necessity to achieve carbon neutrality.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The purpose of this review is to summarize existing knowledge about the influence of vegetation-related factors on the dynamics of soil carbon pools.</span></p>
<ol style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> Major mechanisms of accumulation of soil organic matter</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>1.1. Major mechanisms of carbon transfer from the vegetation pool to the soil pool</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Soil carbon stocks result from organic matter entering the soil and carbon compounds being lost due to decomposition, washing out and leaching (Fig. 1). The source of organic matter is the plant material of both aboveground and underground organs. Other sources of carbon compounds are root secretions, including exudates of plant roots and associated symbionts (for example, mycorrhizal fungal mycelium), as well as fecal material and bodies of soil biota. Once in or on the soil, organic matter is gradually transformed by soil fauna and microorganisms (fungi, bacteria and archaea). Extracellular enzymes secreted by the microorganisms break down the organic matter into simpler compounds that can be assimilated. Part of the plant organic matter is mineralized, and the other part is accumulated in the biomass of consumers and decomposers or in their metabolites, which can partly be excreted from the cells and stabilized. The rest of some stable plant biomolecules, including lipids, lignins and sugars, can also be stabilized by minerals and aggregates.</span></p>
<div id="attachment_4935" style="width: 1034px" class="wp-caption aligncenter"><img aria-describedby="caption-attachment-4935" loading="lazy" class="size-large wp-image-4935" src="https://jfsi.ru/wp-content/uploads/2022/03/kuz-1024x631.png" alt="" width="1024" height="631" srcset="https://jfsi.ru/wp-content/uploads/2022/03/kuz-1024x631.png 1024w, https://jfsi.ru/wp-content/uploads/2022/03/kuz-300x185.png 300w, https://jfsi.ru/wp-content/uploads/2022/03/kuz-150x93.png 150w, https://jfsi.ru/wp-content/uploads/2022/03/kuz-768x474.png 768w, https://jfsi.ru/wp-content/uploads/2022/03/kuz.png 1200w" sizes="(max-width: 1024px) 100vw, 1024px" /><p id="caption-attachment-4935" class="wp-caption-text"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Figure 1.</strong> Carbon fluxes in a forest ecosystem acc. to Mayer et al. (2020), modified</span></p></div>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Since vegetation is the main source of organic matter, it is important to discuss the principal mechanisms of carbon transition from the vegetation pool to the soil pool. Three main mechanisms were identified.</span></p>
<ol style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">Processing of aboveground and underground litterfall by soil fauna and microbiota. Soil fauna mechanically destroys and mixes the litterfall with the soil organic horizon and the mineral part of the soil; therefore, it can contribute to the spatial redistribution and primary destruction of plant material (Brussaard, 1997; Frouz et al., 2013). As decomposers, especially microbiota, further process the soil organic matter (SOM), the soil organic horizon undergoes chemical and structural changes (Frouz, 2018). For example, earthworm activity can both enhance the transformation of soil organic horizon into more stable organomineral aggregates and contribute to the accelerated SOM degradation (Wolters, 2000; Fox et al., 2006), which can be taken into account when identifying functional types of macrofauna (Geraskina, 2020).</span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The transformation of litterfall by soil biota differs significantly under different climatic conditions, since temperature and water regimes are the limiting factors for many decomposers. In boreal forests of the Northern Hemisphere with low temperatures and humid water regime, fungal decomposition usually prevails, which contributes to litter accumulation and formation of raw forms of humus such as mor and moder. In southern regions, biological cycle increases in intensity, and, together with fungal decomposition, the proportion of bacterial decomposition and contribution of soil animals increases, thus contributing to a more intensive litter decomposition and humification of the soil organic horizon and upper mineral horizons.</span></p>
<ol style="text-align: justify;" start="2">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">Vertical stratification of tree roots (Brassard et al., 2011), root litter (Brassard et al., 2013) and root turnover (Brassard et al., 2011; Lei et al., 2012), as well as root exudates (Bardgett, 2005), contribute to the successful association of organic matter with soil aggregates or clay minerals. Some researchers suggest that most of the SOM originates from roots, and the deeper they are, the more significant this source becomes (Rasse et al., 2005), since different types of root systems make vertical stratification of tree roots and redistribution of the rhizosphere into deep mineral horizons possible. Root litter usually decomposes slower than foliar litter of the same species (Lauenroth, Gill, 2003; Cusack et al., 2009); therefore, more stable carbon compounds of roots stay in the soil longer than carbon compounds of aboveground shoots (Rasse et al., 2005). The key mechanisms that lead to root carbon stabilization in the soil include: (1) chemical resistance of roots, partly due to the presence of a persistent compound suberin; (2) physical protection in aggregates; and (3) physico-chemical protection of lignin and reactive carbon compounds in the soil (root exudates) associated with clay minerals. However, the opposite effect is also possible: labile C isolated from the roots may stimulate the decomposition of the already stable SOM (Kuzyakov, Domanski, 2000; Dijkstra, Cheng, 2007).</span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">When comparing different biomes, global patterns in the rate of root system renewal between plant groups and depending on climatic gradients were established. Since the rate of root renewal increases exponentially along with an increase in the average annual temperature (Gill, Jackson, 2000), a more significant contribution of root litter to carbon stabilization can be expected in southern regions.</span></p>
<ol style="text-align: justify;" start="3">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">Leaching of dissolved organic carbon (DOC) from living plants and forest soil organic horizon is yet another way for carbon compounds to go into deeper soil horizons (Fröberg et al., 2011). DOC leaching fluxes are usually higher directly under the forest soil organic horizon and are often associated with microbial activity and biomass (Smolander, Kitunen, 2002). Also, high DOC fluxes into the mineral part of the soil are found in forests with high carbon content in the soil organic horizon, for example, those formed by common spruce. Using the example of spruce forests in the European part of Russia, it was shown that, at a soil solution carbon concentration of 35 mg/L and above, water-soluble organics (WSOs) begin to be sorbed in the BF horizon (Karavanova et al., 2020). In dissolved organic substances moving down the soil profile, lignin derivatives (phenols) are mainly preserved on the surface of oxides and hydroxides in shallow soil layers (Kaiser et al., 2004; Kaiser, Zech, 2000). DOC moving from upper to deeper soil horizons can result in stabilization and, consequently, a significant increase in soil carbon deposition (Kalbitz, Kaiser, 2008). This can be pronounced against the background of excessive moisture, when an increase in the flux of plant-derived DOC and its consolidation are possible (Mikutta et al., 2019). However, an inverse effect is also possible: estimates of DOC removal from temperate pine forests in Belgium (about 10 g/m<sup>2</sup> per year) have shown that DOC can account for 11% of ecosystem net productivity (Gielen et. al., 2011).</span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Carbon fluxes in forests can differ significantly in the north and in the south, since it is known that climatic conditions can have a significant impact on both the amount of DOC and intensity of its removal (Gmach et al., 2020). On the one hand, the biological cycle intensity is higher in southern regions than in northern ones, which contributes to active decomposition of the soil organic horizon and DOC being released. On the other hand, during prolonged dry seasons, decomposition rate of the soil organic horizon may decrease, and secondary metabolites may accumulate (Kalbitz et al., 2000).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Therefore, three main ways of transferring carbon from the vegetation pool to the soil pool were identified. At the same time, there are no estimates of the contribution of each of these mechanisms or their combined impact. It was found that, in the northern and southern regions (of the Northern Hemisphere), the intensity of these processes manifests in different ways. It is expected that in northern ecosystems the contribution of DOC to organic matter accumulation will be high, since these areas have a high level of precipitation and highly developed soil organic horizon. The predominance of fungal decomposition will also contribute to the accumulation of the soil organic horizon. At the same time, the influence of root litter can be very significant, especially if we take into account the influence of not only woody plants, but also that of shrubs widespread in taiga forests and other ground cover vegetation. In the southern coniferous-broadleaf forests, DOC fluxes are much smaller than in taiga, because there is less precipitation. However, an increase in biomass and activity of soil fauna increases its contribution to litter processing and subsequent stabilization of SOM in the mineral part of the profile. An increase in overall productivity of roots, along with influence of saprophages, is probably the key mechanism for a more intensive formation of stable carbon in the mineral horizons of soils found in coniferous-broadleaf forests. However, all these assumptions are hypothetical and require further research.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>1.2 Main mechanisms of carbon fixation in the SOM</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Carbon sequestration in soils is hardly a new aspect in the studies of biogeochemical carbon cycle within the framework of the global climate change and terrestrial ecosystems: as early as in the 1990s some studies provided evidence of the importance of soils as a carbon sink under conditions of elevated CO<sub>2</sub> concentration in the atmosphere.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Carbon sequestration capacity of the soil (Carbon Protection Capacity, CPC) reflects its ability to stabilize and retain the carbon that came with organic matter as part of soil organic matter (SOM) (Semenov et al., 2009; Kogut, Semenov, 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is now agreed that there are two main mechanisms of organic matter stabilization in soils: due to the formation of organomineral complexes (Semenov, Kogut, 2015) and due to the formation of soil aggregates (Six et al., 2002, 2004; von Lützow et al., 2006; Gunina et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Organomineral complexes are formed as a result of physicochemical interaction between molecules of soil organic matter (OM) and mineral components, increasing the protection of this SOM (soil organic matter) from decomposition (von Lützow et al., 2006). For example, these include the formation of high-molecular humic substances; binding of hydrophilic components of fresh organic matter by hydrophobic centers of humic substances, and binding of amino acids by polyphenols; formation of organomineral complexes of aluminum and iron, calcium; and formation of organomineral complexes in mineral matrices of clay particles. These processes mainly involve fine clay and mineral particles, with the high contribution of surfaces of chemically reactive oxides and phyllosilicates (Blume et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Soil aggregates are homogeneous mixtures of minerals, organic compounds and organomineral complexes. For example, microaggregates can prevent microorganisms from getting access to SOM inside the aggregates, increasing its stability (Six et al., 2004). Accumulation of dissolved organic carbon in pores that are smaller than bacteria is also possible (Totsche et al., 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">There is also the third, currently widely discussed mechanism for OM stabilization, i. e. biochemical protection due to certain chemical properties of biomolecules. It was established that in forest ecosystems the major part (≥50%) of stabilized carbon is plant biomolecules, including lipids, lignin and sugars, which can make up a significant part of organic matter protected by minerals and aggregates (Angst et al., 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Biochemical resistance is believed to potentially be most relevant at the initial stages of decomposition and possibly play only a minor role in the long-term protection of SOM in mineral soil (Marschner et al., 2008). Some biomolecules can, however, be “selectively preserved” depending on the thermodynamic conditions of the environment and their ability to be stabilized due to physicochemical interactions in the soil matrix: for example, this is true of lignin (Feng et al., 2005), some sugars (Amelung et al., 1999; Kiem, Kögel-Knabner, 2003) and lipids (Angst et al., 2017b; Ludwig et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The intake and subsequent stabilization of aromatic and aliphatic acids, such as lignin monomers and its oxidation products, depend on the dominant vegetation. Guaiacyl (vanillin) phenols dominate in coniferous forests (Kovalev, Kovaleva 2016); they persist longer during the initial decomposition processes and have a higher probability of binding to mineral surfaces (Clemente, Simpson, 2013). Broadleaf and small-leaved forests contain equal proportions of vanillins and syringils (Kovalev, Kovaleva 2016). The main source of aromatic phenolic compounds in soils is lignin from aboveground and underground biomass of higher plants. Underground parts of plants play the main role (Kovalev, Kovaleva, 2016). The number of studies on the extraction of lignin from intact aggregate structures is small. Few studies indicate higher lignin content and a low degree of lignin oxidation in macroaggregates as compared to microaggregates (Xiao et al., 2007; Thevenot et al., 2010). The participation of lignin phenols in aggregation and concretion formation was established depending on the redox situation of soils (Kovalev, Kovaleva, 2016).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Neutral sugars of plant origin (mainly derived from hemicelluloses) are preferred microbial substrates compared to other forms of SOM (Gunina, Kuzyakov, 2015), such as lignin, and may contribute to the accumulation of microbial necromass. Interestingly, the concentration of mostly neutral sugars of plant origin (for example, xylose and arabinose) in organomineral complexes (52–128 mg/g C) may be several times higher than the concentration of lignin (Kiem, Kögel-Knabner, 2003; Córdova et al., 2018). Neutral sugars of plant origin can make a significant contribution (up to 130 mg/g C) (Córdova et al., 2018) to the stabilization of SOM.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Some lipids of plant origin are considered relatively resistant to degradation, including long-chain n-alkanoic acids, certain monomers of cutin and suberin with groups of hydroxylic and/or carboxylic acids (Quenea et al., 2004; Jandl et al., 2005; Angst et al., 2017a; Anohina, 2020). Stabilization of plant-derived lipids through organomineral interactions may depend on the monomeric composition of lipid biopolymers and chemical properties of these monomers, as well as the composition of microbial population, sorption properties and mineral composition of soils (Bull, 2000). The contribution of plant-derived lipids to stable C ranges from ~2% to ~10% (i. e. ~ 20–100 mg of lipids per g of C (Angst et al., 2021). Studies on the extraction of these compounds from aggregates are few and suggest the presence of alkanes in the soil as part of capsules of the independent phase of lipids (Anohina, 2020). Differences in the quantitative and qualitative composition of the lipid organic profile were revealed for different types of forests.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, vegetation as the main source of organic matter entering the soil determines the possibility and speed of formation and stabilization of soil organic matter. The significant direct contribution of vegetation to the accumulation of SOM is emphasized. While the mechanism of carbon stabilization due to the formation of organomineral complexes has been sufficiently studied and has quantitative estimates, the mechanisms associated with the study of the physical protection of organic matter in soil aggregates and the biochemical stability of plant compounds require attention.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<ol style="text-align: justify;" start="2">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> Mechanisms related to the influence of vegetation on soil carbon stocks at the ecosystem level</strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Taking into account the features of the biogeochemical carbon cycle in forest ecosystems (Fig. 1) and the main mechanisms for the intake and stabilization of organic matter in soils, three main mechanisms of vegetation influence on the dynamics of soil carbon pools at the ecosystem level can be distinguished: (1) litterfall quantity and (2) quality of both individual species (species identity) and their combined influence, i. e. (3) biodiversity of the community.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The amount of litterfall is an important predictor of carbon accumulation (Grandy, Neff, 2008; Gentile et al., 2011; Carrington et al., 2012; Dungait et al., 2012), since vegetation is the main supplier of organic matter to the soil. The amount of litterfall produced is proportional to the net primary productivity of forests in natural forests, since the litterfall is part of the NPP (Chen et al., 2017).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Litter quality also depends on the composition of vegetation and the content of nutrients (nitrogen, phosphorus, potassium, etc.) and secondary metabolites (polyphenols, lignins, cellulose, hemicellulose, etc.) (Berg et al., 1993; Cadisch, Giller 1997; Perez-Harguindeguy et al., 2000, Berg, McClaugherty, 2020). The concept of correlation between accumulation of soil carbon and the quality of litter has been proposed, where high-quality litter does not always contribute more effectively to an increase in stable soil organic matter as compared to low-quality litter (Castellano et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">One of the aspects of biodiversity is structural biodiversity characterizing the spatial organization of vegetation (vertical and horizontal structure), which determines the spatial variations of carbon intake, firstly, by regulating the penetration and falling of organic residues into the organic layer, and secondly, by influencing the hydrothermal regime of the organic layer, that is, the dynamics of temperature and water content, which in turn can affect decomposition to a greater extent than differences in macroclimatic conditions on a continental scale (Joly et al., 2017). Other aspects of biodiversity (typological, species, functional, age, ontogenetic structure, etc.) also determine the dynamics of carbon intake and its quantitative and qualitative characteristics.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Let’s look into the influence of these mechanisms at different spatial levels.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>2.1. </strong>Regional level</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Estimates of the relation between carbon stocks and the amount of litterfall are usually mediated and carried out through changes in productivity. A study by Robert Amundson (2001) shows a trend of increasing soil carbon stocks in forests with an increase in average annual temperature, average annual precipitation and, accordingly, net primary productivity.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is known that the productivity of woody plants in the northern hemisphere naturally decreases from south to north (Utkin, 1975). It is also shown that the amount of litterfall increases with decreasing latitude (Albrektson, 1988). Annual intake of litter in the North taiga forests varies within 0.9–2.5 t/ha (Nikonov, 1986), in the middle taiga — 2.1–3.9 t/ha (Kazimirov, 1977; Red’ko, 1984). More highly productive communities contribute to a greater intake of litterfall: according to literature data, the annual intake of litterfall in the forests of the coniferous-broadleaf subzone varied from 2.5 to 4.4 t/ha in the forests of the Moscow region (Karpachevsky, 1977), 3.1–4.4 t/ha in the forests of the Bryansk Polesie (Shabliy, 1990), and 3.9–12.2 t/ha in the forests of the North-Western Caucasus (Zon, 1950).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The litter quality of both coniferous and deciduous trees may vary depending on climatic conditions. The relation between the concentration of N and the average annual temperature and annual precipitation was revealed; it was found that for both coniferous and deciduous tree species, the total concentration of N in the litterfall increases with an increase in hydrothermal values (Berg, McClaugherty, 2020). It is believed that, at the regional level, temperature is the leading factor in determining the rate of litter decomposition (Meentemeyer, 1978; Hobbie, 1996). However, when studying the rate of decomposition of ground litter, taking into account the climatic gradient, it is noted that in boreal forests about 16% of decomposition can be explained by nitrogen concentration (Dyer et al., 1990). When studying the rate of decomposition of underground litterfall, taking into account climatic characteristics, it was shown that the chemical composition of the roots is the main regulator of decomposition processes, while climatic and environmental factors were of secondary importance (Silver, Miya, 2001). It is shown that the quality of litter can be a more significant predictor of soil organic horizon decomposition as compared to hydrothermal characteristics (Swift et al., 1979; Berg, 2000), especially at the initial stages of its decomposition (Canessa et al., 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>2.2. </strong>Local level</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">2.2.1. Interbiogeocenotic heterogeneity</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Studies show that the diversity of tree species increases forest productivity due to greater spatial complementarity of tree crowns, which, in turn, provides a positive correlation between the diversity of tree species and the productivity of litterfall (Zheng et al., 2019). The amount of litterfall increased with increasing species richness. Data on the relation between age and amount of litterfall are contradictory. Some publications have noted a positive relation between age and amount of litterfall in the first 98 years of the development of the pine community with the subsequent flat trend (Chen et al., 2017); in other studies the amount of litterfall decreased with increasing age of the stand (Albrektson, 1988). It is also noted that the amount of litterfall may increase with increasing soil fertility (Albrektson, 1988). In coniferous-broadleaf forests the effect of age on the amount of litterfall was not seen (Huang et al., 2017).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Low-quality litter is characterized by a low content of bases, high acidity, high content of lignin and secondary metabolites, as well as a wide C/N ratio. The litter of coniferous trees has a low nitrogen content: for example, pine litter often contains less than 0.4% nitrogen (Berg, McClaugherty, 2020); the litter of boreal shrubs is rich in polyphenolic compounds (Wardle et al., 2003); and green mosses have a low content of nutrients (Hilli, 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The high-quality litter is characterized by a high content of bases, low acidity, and narrow C/N ratio. Deciduous tree litter is rich in nitrogen: for example, birch litter contains 0.7%, beech — 0.9%, aspen — 1.0%, hornbeam — 1.1%, oak — 1.2%, maple — 1.3%, and lime — 1.5% (Simon et al., 2018). The richer is litter with nutrients, the faster is the soil organic horizon decomposed by soil biota, which leads to a decrease in its stock and, accordingly, carbon stocks in it.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Already in the 1990s it was recognized that the availability of nitrogen is the main determinant controlling the response of soil carbon to climate changes in ecosystems whose nitrogen is a limiting factor (Diaz et al., 1993; Nohrstedt, 1992). A number of studies have shown that the addition of nitrogen stimulates the decomposition of high-quality litter, but slows down or prevents the decomposition of low-quality litter (Knorr et al., 2005), which is due, on the one hand, to the suppression of the activity of lignolytic enzymes (Carreiro et al., 2000), and, on the other hand, to an increase in the number of microorganisms (Córdova et al., 2018). There is also evidence that nitrogen stabilizes organic matter in the soil (Neff et al., 2002; Swanston et al., 2004) and prevents the mineralization of carbon accumulated earlier (Hagedorn et al., 2003). Broadleaf species, in particular elm, oak and poplar, can be considered the land-reclamation species that accelerate the nutrient cycle in pine plantations (Polyakova, Billor, 2007).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">When comparing decomposition rates of different plant groups, the influence of the functional characteristics of plants associated with phylogenetic groups was demonstrated. Faster decomposition of deciduous broadleaf species as compared to coniferous species, as well as a faster decomposition of grass species as compared to cereals, has been found. Slow decomposition rates of ferns and mosses have been noted (Cornwell et al., 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">However, the most common predictors of the decomposition rate of soil organic horizon are relative indicators like the C/N ratio, as well as the content of nutrients in the soil organic horizon (Zhang et al., 2008). Some European (Lovett et al., 2004; Reich et al., 2005; Oostra et al., 2006) and North American studies (Finzi et al., 1998; Neirynck et al., 2000; Dijkstra, Fitzhugh, 2003; Hagen-Thorn et al., 2004) of plants of the genera Fraxinus, Acer, Quercus and Fagus show the differences in carbon pools of the soil organic horizon and the C/N ratio as an indicator of accumulation rate of the soil organic horizon. Ash, maple and lime are combined into a group of plants with high-quality litter, that is, the litter with high nitrogen content, which leads to low accumulation of C in the soil organic horizon due to a high decomposition rate. Oak and beech have a relatively low content of C and N in the litterfall which results in low content of C and N in the forest soil organic horizon, high C/N ratio in the forest soil organic horizon, and low decomposition rate, which leads to an increase in pools of C and N in the forest soil organic horizon. When comparing coniferous species with deciduous ones, it was found that spruce has the highest C/N ratio and carbon stocks in the soil organic horizon (Vesterdal et al., 2008). At the same time, comparison of coniferous species with each other showed that the soil organic horizon of pine forests often has a much wider C/N ratio than that of spruce forests (Lukina et al., 2020). Several studies have confirmed the informative value of the lignin/N ratio of the soil organic horizon in predicting the rate of decomposition of the soil organic horizon between species (Gower, Son, 1992, Heim, Frey, 2004). The following lignin/N ratios were revealed (in descending order): spruce, beech > oak >> maple, lime >>> ash (Melillo et al., 1982; Lovett et al., 2004; Sariyildiz, Anderson, 2005; Kalbitz et al., 2006; Cotrufo et al., 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">High-quality litter, i. e. that enriched with nutrients and with a minimum C/N ratio and lignin content, usually decomposes faster than low-quality litter (with nutrient deficiency and a lot of lignin). However, the contribution of rapidly and slowly decomposing litter fractions to SOM accumulation is currently not fully clear (Castellano et al., 2015).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">It is believed that slowly decomposing material of soil organic horizon contributes to the accumulation of carbon in the soil more than more rapidly decomposing material (Swift et al., 1979), especially in organogenic soil horizons, since low-quality litter is slowly processed by soil biota (Striganova, 1980; Prescott et al., 2000; Huang et al., 2020; etc.) and promotes the growth of fungi and their contribution to carbon stabilization on the mineral matrix (Six et al., 2004; Soares, Rousk, 2019). However, the results of other studies show that mixing slowly decomposing soil organic horizon with low-quality litter and rapidly decomposing soil organic horizon with high-quality litter contributes to higher efficiency of carbon transfer to soil mineral horizons (Cotrufo et al., 2013; Córdova et al., 2018), mainly due to an increase in DOC fluxes from the developed soil organic horizon (Fröberg et al., 2011) and by adding readily available nitrogen of rapidly decomposing litter fractions. It has been shown that the content of DOC, especially in the surface layers of the soil, is positively correlated with the decomposition rate of the soil organic horizon (Zhou et al., 2015). There are regular connections between litter stocks and the flux of DOC: in forests with a high proportion of deciduous trees with less developed soil organic horizon, less intensive removal of DOC was noted (Fröberg et al., 2011). There is also data on the relation between the qualitative and quantitative characteristics of DOC and root distribution.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Another important aspect of the impact of litter quality is the comparison of litter quality in the soil organic horizon of monodominant and mixed forests. It has been shown that forests with higher diversity, that is, high plant species density, have a narrower C/N ratio (Polyakova, Billor 2007; Huang et al., 2017). There are also studies on the positive effect of plant functional diversity on the decomposition of litter (Patoine et al., 2017), since the combined effect of litter of different quality can create special favorable conditions for the activity of soil biota. For example, a number of studies have shown that the poor quality of spruce and fir litterfall results in the accumulation of the soil organic horizon, which is a habitat for saprophages functionally related to it (Kuznetsova et al., 2019; Huang et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The joint influence of species is seen when comparing carbon stocks of different forest types (Framstad et al., 2013; Lukina et. al., 2020; Kuznetsova et al., 2021). The differences may be due to the different ratio of plants of the tree layer. In the soils of broadleaf forests, it was not possible to find significant differences between carbon accumulation in monocultures in common garden experiments (Vesterdal et al., 2008). The question of the combined effect of several species of woody plants on carbon stocks remains open. V. N. Shanin and colleagues (Shanin et al., 2014) studied the influence of the richness of boreal forests on their productivity and carbon dynamics and revealed that mixed forests are more productive than monodominant ones. With regard to coniferous-deciduous forests, it has been shown that a greater variety of tree species causes an increase in organic carbon stocks in the soil (Vesterdal et al., 2013). However, there are other estimates showing that the carbon stocks of the soil are more influenced by the identity of tree species as compared to their diversity (Dawud et al., 2016).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The higher the diversity of woody plant species, the more mixed the litter becomes. At the same time, the litter of conifers is decomposed faster in the presence of deciduous tree litter (Patoine et al., 2017), therefore, the quality of litter in general improves, which contributes to more intensive decomposition and an increase in carbon fluxes into mineral horizons. Manipulations with the ratio of species with different litter quality are common practice in climate-smart forestry (<a style="color: #000000;" href="https://doi.org/10.1016/j.foreco.2020.118127">Mayer</a> et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">There are few studies on the relation between the diversity of vegetation under the canopy with ground cover and carbon dynamics. In boreal forests, there has been a significant increase in carbon stocks in forests where the contribution of grasses to the total projective cover exceeds 10% (Lukina et al., 2020). In coniferous-broadleaf forests, a positive relation between the content of soil carbon and the diversity of trees in the layer of sub-canopy vegetation was noted (Bakhshandeh-Navroud et al., 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Unlike boreal forests, where mosses and dwarf shrubs mainly predominate in the ground cover and the addition of grasses significantly affects the processes of carbon accumulation, in coniferous-broadleaf forests, where grasses mainly predominate in the ground cover, their influence may be less pronounced (Kuznetsova et al., 2021).</span></p>
<ul style="text-align: justify;">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;">Intrabiogeocenotic heterogeneity</span></li>
</ul>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">There are quite a lot of works evaluating carbon cycles in different mosaic elements (Orlova, Lukina, 2016; Lukina et al., 2018; Priputina et al., 2020; etc.). Humification of the leaf-based soil organic horizon in the gaps and below the crowns differed in winter and summer (Ni et al., 2015). Emission was 4 times higher in the gaps of old-aged spruce forests as compared to the space below the canopy (Karelin et al., 2017). The mass of needle litterfall was significantly higher (7.5%) directly under the tree crowns. Whereas the characteristics of the litter quality (acidity, content of N and other nutrients, C/N ratio) did not show any significant differences when comparing the spaces in below and between the crowns in 55-year-old pine forests of northwestern Germany, a number of studies show trends towards higher pH and nutrient content in spaces below the crowns in contrast to spaces between the crowns (Penne et al., 2010).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">There are very few studies assessing the influence of different mosaic elements related to the distribution of underground parts of plants on both cycles and carbon pools (Liang et al., 2017; Sokol et. al., 2019). It is known that different types of trees have different patterns of root distribution in the soil profile. Root systems of the common spruce are of the surface type and are located mainly in the forest soil organic horizon (Puhe, 2003). In the mineral soil, low mass of roots was seen (<5 mm) at a depth of 0–20 cm under the spruce and beech as compared to the oak and ash (Oostra et al., 2006). It has also been reported that ash has relatively thinner roots at a depth of 16–30 cm than oak, whereas at a depth of 0–15 cm the situation is quite opposite (Ponti et al., 2004). Analysis of the qualitative composition of organic carbon taking into account the depth of the soil profile showed an increase in the content of lipids, the source of which was underground litterfall, as compared to the content of lipids, the source of which was ground litterfall (Nierop, 1998; Nierop et al., 2006; Feng, Simpson, 2007; Spielvogel et al., 2014; Angst et al., 2016). Using the example of both coniferous and deciduous forests, it was shown that root litterfall can make a comparable contribution to the soil carbon stock as compared to ground litterfall (Rasse et al., 2005).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Regulation of the migration of carbon compounds by atmospheric precipitation and soil waters within the boundaries of the soil profile may also be associated with the structural organization of biogeocenosis (Ershov, 2021). According to long-term observations, the intake of organic carbon compounds with atmospheric precipitation in northern taiga forests below pine crowns is 5-6 times higher than in spaces between the crowns throughout the vegetation season, which may also explain the differences in carbon concentrations in soil waters (Lukina et al., 2018; Ershov et al., 2019). Significant carbon removal with soil waters is typical of coniferous forests, especially in the spaces between the crowns (Fröberg et al., 2011; Lukina et al., 2018; Akkumuljacija&#8230;, 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Thus, forest biodiversity may influence carbon cycles, on the one hand, through change in quality and quantity of litterfall, and, on the other hand, through change in physical environmental conditions like humidity or temperature. All this affects the change in activity of the soil biota. Vegetation composition determines quantity, quality and rate of decomposition of plant litter, its horizontal distribution and distribution of carbon compounds within the soil profile (Gleixner, 2013), which determines the formation of soil carbon pools.</span></p>
<ol style="text-align: justify;" start="3">
<li><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> Existing estimates of carbon stocks in taiga and coniferous-broadleaf forests </strong></span></li>
</ol>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.1. Regional level</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Russia accounts for more than 20% of the world’s forest cover and more than half of the world’s boreal forest resources. According to existing estimates, the soils of forest ecosystems account for about 46% of the total carbon stocks in the soil cover of Russia (Schepashchenko et al., 2013). At the same time, the contribution of forest lands in the Asian and European parts of the country is different and amounts to 48% and 37%, respectively, which reflects the level of forest cover of these territories.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Soil carbon stocks are closely related to the natural and climatic zone: the highest level of accumulation is characteristic of forests that formed under cool and humid conditions, whereas in warmer and drier climates, stocks decrease both on a global scale (Post et al., 1982; Jobbagy, Jackson, 2000) and on a (sub)regional scale (Burke et al., 1989; Alvarez, Lavado, 1998; Paul et al., 2002; Callesen et al., 2003; Baritz et al., 2010; Badgery et al., 2013; de Brogniez et al., 2014; Rossel et al., 2014; Hobley et al., 2015; Gray et al., 2016; Chestnyh et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Total carbon stocks in automorphic and semi-hydromorphic soils, taking into account the hydromorphic soils of swamps in the forest areas of the European-Ural part with an area of 181.13 × 10<sup>6</sup> ha, amount to 19.3 × 10<sup>9</sup> t C (Chestnyh et al., 2020). Of these, north taiga carbon stocks in the 0–30 cm layer, including the carbon stocks of the soil organic horizon, organogenic and mineral horizons, amount to 4.94 ± 2.01 × 10<sup>9</sup> t C, in the middle taiga forests — 2.92 ± 0.93 × 10<sup>9</sup> t C, and in southern taiga forests — 2.09 ± 1.80 × 10<sup>9</sup> t C. The lowest stocks were revealed in the zone of coniferous-broadleaf forests — 1.02 ± 0.67 × 10<sup>9</sup> t C. The highest average values were found in the soils of the northern taiga forests, if we take swamp forests into account. The minimum average values are typical for the coniferous-broadleaf area, which the authors explain with either climatic features and widely spread breaking new grounds, or lack of data on swamps in this area.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In automorphic soils, soil organic horizon on average accounts for 30% of the total stock of C in the 0–30 cm layer, whereas in layers of 0–50 cm and 0–100 cm, it decreases to 24% and 18%, respectively (Chernova et al., 2020). The average carbon stock of the soil organic horizon depends on the natural and climatic zone: there is a trend to decreasing stocks of the soil organic horizon and carbon stocks therein from the northern taiga subzone to the subzone of coniferous-broadleaf forests (Chestnyh et al., 2007; Kuznetsova et al., 2020). The average carbon stock of the soil organic horizon is 11 t/ha in the forests of the northern taiga, 10 t/ha in the forests of the middle taiga, and 7 t/ha in the more southern regions (Chestnyh et al, 2007). Total carbon stock in the forest floor throughout Russia ranges from 5.3 Pg С (Chestnyh et al., 2007) to 8.4 Pg С (Schepashchenko et al., 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In contrast to carbon stocks of the soil organic horizon, more intensive carbon accumulation was revealed in the mineral horizons of automorphic soils of the southern regions as compared to the northern ones. For example, carbon stocks of the soil organic horizon in blueberry pine forests naturally decreased from 47±8 t/ha in the northern taiga to 8±1 t/ha in coniferous-broadleaf forests, and stocks in the 0–10 cm layer representative of the humus-accumulative horizon, on the contrary, increased from 8±1 to 18± 2 t/ha (Kuznetsova et al., 2020).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.2. Local level</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>3.2.1. Interbiogeocenotic heterogeneity </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the boreal forests of the European part of Russia, the average carbon stock varies from 10.6 to 17.2 t/ha in organic horizons, while in the 30-centimeter layer (mineral layer) it varies from 46.6 to 122.2 t/ha (Rasporjazhenie&#8230;, 2018). The average carbon stock of the soil organic horizon ranges from 0.6 to 28 t/ha in pine forests, from 0.9 to 58 t/ha in spruce forests, from 1 to 29 t/ha in broadleaf forests, from 0.3 to 27.4 t/ha in birch forests, and from 0.7 to 19.9 t/ha in aspen and other soft-wooded broadleaf forests (Chestnyh et al., 2007). In the Komi Republic, in the north-east of the European part of Russia, the carbon stock in the one-meter layer of soil varied from 29 t/ha to 121 t/ha, depending on the type of soil (Dymov, 2018). In the Republic of Karelia, the stocks of soil carbon in the one-meter layer also varied significantly and amounted to 24–434 t/ha in pine forests and 39–402.4 t/ha in spruce forests depending on the type of soil and humidity (Bakhmet, 2018). The results of soil carbon assessments in the boreal forests of the Scandinavian countries indicate that the greatest diversity of soil carbon stocks associated with environmental factors is detected in organic horizons (Framstad et al., 2013). According to field data, the average national value of organic carbon stocks was 92 t/ha in forests with spruce predominance and 57 t/ha in forests with pine predominance (Stendahl, 2010). Modeling has demonstrated that the accumulation of organic carbon stocks is 22% higher in spruce forests than in pine forests under similar environmental conditions. In Norway, soil carbon stocks were higher in more productive forests than in less productive forests, which was believed to be due to different soil thickness (de Wit, 1999). In Finland, carbon stocks in the soil varied depending on the weight of the forest soil organic horizon, weather conditions and logging (State of&#8230;, 2012). It has been shown that a more productive forest accumulates more carbon in the soil of Finnish forests (Leskinen et al., 2020). In Canada, soil carbon stocks in boreal mixed forests on sandy loam (northeast Ontario) was on average 51 t/ha and amounted for up to 30% of the total carbon stocks, whereas the forest soil organic horizon accumulated 22 to 36 t C per ha. In the Canadian boreal biome, soils in forests of black spruce, that have a slow carbon cycle, stored more organic carbon than the soils of aspen forests. (Laganiere, 2013).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the zone of coniferous-broadleaf forests of the European part of Russia, the average carbon stock ranges from 10.6 to 17.2 t/ha in organic horizons, while in the 30-centimeter soil layer it ranges from 46.6 to 122.2 t/ha (Rasporjazhenie&#8230;, 2018). In the secondary post-agrogenic lime-aspen mixed-herbs forests of the Moscow region, the carbon stock in the 0–60 cm layer was 88 t/ha (Baeva et al., 2017). In the forest biogeocenoses of the Middle Volga region in the soil layer of 0–50 cm, the carbon stock ranges from 12.8 to 439.5 t/ha depending on the type of forest, soil type and humidity (Demakov et al., 2018).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In the organic horizon of broadleaf forests of the Central Europe, low carbon stock was found under the beech (0.42 t/ha) and in lime forests (0.20 t/ha) (Langenbruch, 2012), whereas in hornbeam-oak forests it is up to 8.4 t/ha (Bruckman et al., 2016) and is about 6 t/ha in the organic horizon of spruce forests of the Western Black Sea region (Misir et al., 2012). In common garden experiments, carbon stocks of the soil organic horizon ranged from 1.8 t/ha in lime, maple and ash crops to 3.8 t/ha in oak and 4.5 t/ha in birch forests. Up to 14.5 t/ha of carbon accumulated under the spruce (Vesterdal et al., 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Soil carbon stocks in the mineral layer of 0–20 cm amounted to 52 t/ha in beech forests and 45 t/ha in lime forests of Central Europe (Langenbruch, 2012), while in the 0–50 cm soil layer of hornbeam-oak forests it reached 77 t/ha (Bruckman et al., 2016), and in fir forests of the western Black Sea region — 155 t/ha (Misir et al., 2012). In common garden forests, carbon stocks in the 0–30 cm layer ranged from 61 t/ha in spruce and birch stands to 64–67 t/ha in maple and lime and 69–71 t/ha in oak and ash forests (Vesterdal et al., 2008).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Therefore, it was shown that forest soils of the temperate zone have a significant carbon stock of up to 100 Mg C/ha or more. However, the variability is very high for both organogenic and mineral soil horizons. The rate of soil carbon sequestration in these forests depends on the type of soil, previous carbon stock, species composition of vegetation and other natural and anthropogenic factors (Lal, Lorenz, 2012).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>3.2.2. Intrabiogeocenotic heterogeneity </em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">There are very few studies assessing carbon pools in different elements of the mosaic. When comparing spruce parcels, minimal carbon stocks were found in the oxalis and dead-cover parcels, where stocks on average amount to 13 t/ha in the upper mineral layer of 0–5 cm as compared to blueberry, green moss and sedge spruce parcels, where stocks amount to 17–18 t/ha (Podvezennaja, Ryzhova, 2010). An increased pool of the soil organic horizon in the near-trunk spaces was shown as compared to the spaces below and between the crowns (Podvezennaja, Ryzhova, 2010). The example of pine forests in northwestern Germany demonstrated a tendency to increasing carbon stocks of soil organic horizon from 35 ± 9 t/ha in the spaces below the crowns to 38 ± 9 t/ha in the spaces between the crowns (<a style="color: #000000;" href="https://doi.org/10.1016/j.geoderma.2010.05.007">Penne</a> et al., 2010).</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Vegetation as the main source of organic matter entering the soil determines the possibility and rate of soil organic matter formation and accumulation. Three main ways of transferring carbon from the vegetation pool to the soil pool have been identified: processing of aboveground and underground litterfall by soil fauna and microbiota, association of organic matter with soil aggregates due to root litter and root exudates, leaching of dissolved organic carbon from living plants and forest soil organic horizon. At the same time, there are no estimates of the contribution of each of these mechanisms or their combined impact. It was shown that such mechanisms of soil organic matter fixation as physical protection of organic matter in soil aggregates and biochemical stability of plant compounds require attention.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Analysis of the current state makes it possible to identify three main mechanisms that determine the dynamics of soil carbon pools and are associated with vegetation: quantity and quality of litter of both individual species (species identity) and their joint influence (structural diversity of communities). Vegetation diversity affects carbon and nitrogen cycles through changes in biotic conditions (litter quality and quantity), on the one hand, and through changes in physical environmental conditions (humidity, temperature), on the other hand. The influence of all aspects of diversity is significant. There are a number of works showing the influence on carbon of tree species, age of the stand, structure of crowns and mosaic pattern of the biogeocenosis.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Due to a close relation between vegetation diversity and carbon stocks, it is possible to manage soil carbon pools with the introduction of climate-smart forestry. However, in order to properly manage the size of the soil carbon pool, deeper knowledge of the sequestering potential of soils and its main controlling factors is needed, where gaps in knowledge are known. For example, there are singular estimates of the contribution of the lower layers of vegetation to the variation of soil carbon stocks, rare are estimates of the effect of micromosaic pattern (structural diversity) of forests on the soil carbon pool, comparison of the influence of individual woody plants and their combined effect on the soil carbon pool. Estimates of the impact of vegetation regulation of volume and composition of precipitation penetrating through the vegetation canopy on the soil carbon pool are also rare, and there are next to none estimates of the contribution of combinations of various factors, including vegetation-related ones, to the regulation of soil carbon stocks.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Studies are mostly focused not on carbon pools, but on individual components of stock estimates, mainly the carbon content. There is an issue as well that is related to the limited number of estimates of soil carbon pools in the subzone of coniferous-broadleaf forests.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FINANCING</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The study was performed within the framework of the state CEPF RAS assignment 121121600118-8and RFBR project No 20-34-90137.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
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<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Xiao C., Bolton R., Pan W. L., Lignin from rice straw Kraft pulping: Effects on soil aggregation and chemical properties, <em>Bioresource technology</em>, 2007, Vol. 98, No 7, pp. 1482–1488.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhang D. Hui D., Luo Y., Zhou G., Rates of litter decomposition in terrestrial ecosystems: global patterns and controlling factors, <em>Journal of Plant Ecology</em>, 2008, Vol. 1, No 2, pp. 85–93.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zheng L. T., Chen H. Y. H., Yan E. R., Tree species diversity promotes litterfall productivity through crown complementarity in subtropical forests, <em>Journal of Ecology</em>, 2019, Vol. 107, No 4, pp. 1852–1861.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zhou W. J. Sha L. Q., Schaefer D. A., Zhang Y. P., Song Q. H., Tan Z. H., &#8230; &#038; Guan H. L., Direct effects of litter decomposition on soil dissolved organic carbon and nitrogen in a tropical rainforest, <em>Soil Biology and Biochemistry</em>, 2015, Vol. 81, pp. 255–258.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Zonn S. V., <em>Gorno-lesnye pochvy severo-zapadnogo Kavkaza </em>(Mountain forest soils of the northwestern Caucasus)<em>,</em> Leningrad: Izd-vo AN SSSR, 1950, pp. 55–145.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewer: </strong>Candidate of Agricultural Sciences L. G. Isaeva</span></p>
<p style="text-align: justify;">
</div>
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		<item>
		<title>REVIEW OF OPEN SOURCE QGIS FORESTRY PLUGINS</title>
		<link>https://jfsi.ru/en/5-1-2022-podolskaia/</link>
		
		<dc:creator><![CDATA[lena]]></dc:creator>
		<pubDate>Thu, 24 Mar 2022 10:25:19 +0000</pubDate>
				<category><![CDATA[№1 2022]]></category>
		<guid isPermaLink="false">https://jfsi.ru/?p=4899</guid>

					<description><![CDATA[Original Russian Text © 2021 E. S. Podolskaia published in Forest Science Issues Vol. 4, No. 2, Article 86 E. S. Podolskaia Center for Forest Ecology and Productivity of the Russian Academy of Sciences Profsoyuznaya&#46;&#46;&#46;]]></description>
										<content:encoded><![CDATA[<p style="text-align: left;"><span style="color: #000000;"><a style="color: #000000;" href="https://jfsi.ru/wp-content/uploads/2022/03/5-1-2022-Podolskaia.pdf"><img loading="lazy" class="size-full wp-image-1122 alignright" src="https://jfsi.ru/wp-content/uploads/2018/10/pdf.png" alt="" width="32" height="32" /></a></span></p>
<p style="font-weight: 400; text-align: justify;"><span style="font-family: 'times new roman', times, serif; font-size: 10pt; color: #000000;">Original Russian Text © 2021 E. S. Podolskaia published in Forest Science Issues Vol. 4, No. 2, <a style="color: #000000;" href="https://jfsi.ru/4-2-2021-podolskaia/" target="_blank" rel="noopener noreferrer">Article 86</a></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>E. S. Podolskaia</strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Center for Forest Ecology and Productivity of the Russian Academy of Sciences </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>Profsoyuznaya st. 84/32 bldg. 14, Moscow 117997, Russian Federation</em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em> </em></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">E-mail: podols_kate@mail.ru</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Received 13.04.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Revised 12.05.2021</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Accepted 21.06.2021</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">To date, the forestry industry has obtained certain experience in implementing Open Source software. The article describes Open Source QGIS-plugins for the tasks of forest fires and forest resources monitoring and management in research and applications. The functionality analysis performed aimed to simplify the selection of tools for a forest geoinformation project in desktop and web versions. The article presents a brief description of presently downloadable QGIS (version 3.18.1) forestry plugins. The analysis of external QGIS-plugins for working with forest resources and fires has shown the heterogeneity of research. This prevents from identifying trends so far. An option for future research subjects may be development of plugins with available data as cartographic services for territories of different spatial coverage, taking into account that archived data and their availability is a key asset in the forestry. Subject-related forest scope in the present-day repository of QGIS-plugins tends to be relatively limited. Transport and environmental applications implemented in the form of GIS tools are more numerous and can solve individual tasks in the orest project. A review of plugins functionality, compatibility to the core QGIS and their performance should be performed on a regular basis, following new QGIS versions.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><em>Keywords:</em></strong><em> forest resources, forest fires, forestry management, QGIS, plugin</em></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">According to the strategy for the development of the forest complex in the Russian Federation until 2030 (Decree of the Government of the Russian Federation dated September 20, 2018 No 1989-р), forestry is an industry for reproducing forests, protecting them from fires, harmful organisms, and other negative factors; regulation of forest utilization and record of forest resources with the purpose of meeting the needs of the economy in wood and other forest products while preserving the environmental and social functions of the forest.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Awareness of the industry scope implies the need for tools, especially geoinformation-based ones, with the help of which the assigned thematic tasks are to be addressed. By the beginning of the third decade of 21<sup>th</sup> century, forestry has accumulated some experience in using Open Source software. Namely, in 2010, the branches of the Federal State Institution Russian Centre of Forest Health attempted to design blueprints for the state of plantings and the number of pests in Open Source QGIS (Krylov et al., 2012). International examples are presented by such subject-related projects as the construction of forest fire danger maps for local government in QGIS and GRASS (https://clck.ru/bVQE3; https://clck.ru/bVQE8). Paper (Korosov, Zorina, 2016) considers environmental issues of implementing QGIS in the learning guide. Open Source is widely used in a variety of geographical analysis’s instruments. In particular, a work done by Lovelace (2021) presents a large contemporary overview of tools for geographical analysis. It provides a description of tools’ ecosystem, dynamic in time and content and currently consists of around 25 items.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Systematization of currently used geoinformation tools to solve forestry issues could be useful in the form of links list with description. This list could be used when starting a new subject-related forest GIS project. In the present case, the need for such a description arose at the starting point of the interdisciplinary working group “Mapping forestry ecosystem services”, established at the CEPF RAS in February, 2021.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In recent years the number of solutions for QGIS, which occupies a remarkable niche in the modern GIS landscape, has been increasing (https://gisgeography.com/mapping-out-gis-software-landscape/). There are examples of books describing technology for a specific thematic task. For example, QGIS for remote sensing in the forestry industry and agriculture (QGIS and applications in agriculture and forest, 2018) contains a section on the recognition and mapping of continuous logging on the optical satellite images. Basic operations with raster and vector data are performed using standard or core plugins. They are mandatory for any subject-related area, combined in a set of so-called top-5-10 tools (https://digital-geography.com/top-5-qgis-plugins/) or plugins for solving a field-specific task (e. g., https://clck.ru/bVQEN). Often a plugin release in the repository is accompanied by a link to the article published (e. g., in research community ResearchGate, https://www.researchgate.net) with the information on its development, for example, in the works by M. Jung (2013) and L. Duarte et al. (2018). An educational module for the forestry issues is included in the documentation, an example for QGIS version 3.16 (https://clck.ru/bVQEQ).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The article aims to provide a review of plugins for solving forestry challenges to monitor and to manage forest fires and forest resources in the desktop version of Open Source QGIS 3.18.1, which is current as of April, 2021. Selected plugins are aimed to solve the transportation problem of land access and transport modeling at different degree. These topics are in the scope of CEPF RAS Forest ecosystems monitoring laboratory’s activity (<a style="color: #000000;" href="http://cepl.rssi.ru/transport-modeling/">http://cepl.rssi.ru/transport-modeling/</a>), including using Open Source software and Open data Open Street Map (OSM, <a style="color: #000000;" href="https://www.openstreetmap.org/">https://www.openstreetmap.org/</a>), in particular, in the work by Podolskaia et al. (2020). QGIS tools application experience is systematized in a manual by Podolskaia (2020).</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>MATERIALS AND METHODS</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">QGIS (Quantum GIS) with a plugin library (QGIS plugins web portal), located at https://plugins.qgis.org/plugins/, currently holds the leading position among Open Source software used in scientific research. Plugins are software modules used to extend the standard functionality of QGIS application in desktop and web-projects, solving a field-specific task using custom or predefined data sets, services, classifications and formats (a typical example is available at https://clck.ru/bVQES). Plugins can be internal (core) and external (https://clck.ru/bVQEV). Core plugins are written in C++ or Python and included in every new QGIS version released; QGIS core developers maintain them. Python-developers write and support external plugins worldwide.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">In this review we consider upon external plugins. The interface of such plugins can be presented in different languages (English is a priority) depending on the target audience, data used in the plugin and a country of development. Plugins functionality should be compatible to the coming QGIS versions. Plugin performance depends on the technical capacities of hardware which hosts the plugin. Speed and uninterrupted Internet connection is of primary importance when there is a need to connect to the third-party services, including map services. The variety of plugins available in the repository (as of April 8, 2021) for forest infrastructure and forest fires is presented on the Fig. 1.</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">(a)</span></p>
<p style="text-align: justify;"><span style="color: #000000;"><img loading="lazy" class="aligncenter size-full wp-image-4900" src="https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-†.jpg" alt="" width="341" height="369" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-†.jpg 341w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-†-277x300.jpg 277w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-†-139x150.jpg 139w" sizes="(max-width: 341px) 100vw, 341px" /></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">(b)</span></p>
<div id="attachment_4901" style="width: 850px" class="wp-caption aligncenter"><img aria-describedby="caption-attachment-4901" loading="lazy" class="wp-image-4901 size-full" src="https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-b.jpg" alt="" width="840" height="480" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-b.jpg 840w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-b-300x171.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-b-150x86.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-1-b-768x439.jpg 768w" sizes="(max-width: 840px) 100vw, 840px" /><p id="caption-attachment-4901" class="wp-caption-text"></span> <span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Figure 1.</strong> Plugins on forest infrastructure and forest fires (a) and tools to solve forestry tasks (search by the keyword «forestry» (b) https://plugins.qgis.org/plugins/tags/forestry/)</span></p></div>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Plugin Builder (a plugin for plugins), was already available in the first versions of QGIS 1.x and 2.x (https://plugins.qgis.org/plugins/pluginbuilder/) and has been used since 2011 to create the templates for plugins themselves. The tools for working with OSM project data currently prevail in terms of the number of views and downloads. One of the oldest plugins for the forest industry is the plugin for landscape ecology statistics called LecoS (Landscape Ecology Statistics, <a style="color: #000000;" href="https://plugins.qgis.org/plugins/LecoS/">https://plugins.qgis.org/plugins/LecoS/</a>), which is available for use starting from QGIS 1.8. It is developed in Python and uses its SciPy and Numpy libraries to calculate basic and additional metrics of landscape analysis sourced from FRAGSTATS software (https://clck.ru/bVQEe).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Some plugins available are indicated as experimental. An example is a module to create a network of routes for accessing forest resources and forest fires (LCPNetwork, https://plugins.qgis.org/plugins/LCPNetwork/). It implements Dijkstra algorithm, which is considered classic for transport applications and is used to build optimal routes between features of two point vector layers. The result of the plugin is an accumulated cost map for each of the points and a linear layer of traffic routes. Table 1 summarizes the description of plugins for solving forestry issues (repository at <a style="color: #000000;" href="https://plugins.qgis.org/">https://plugins.qgis.org/</a> as of April 8, 2021). The status of all plugins in Table 1 is defined by the authors as non-experimental, with the exception of Manejo.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Table 1. </strong>External QGIS plugins for forest resource management and forest fire monitoring</span></p>
<div style="overflow-x: auto;">
<table style="border: 1px #f1f1f1 solid; background-color: #ffffff;">
<tbody>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Plugin name and link to the QGIS plugins repository</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Description</strong></span></td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Links to additional Internet sources</strong></span></td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></td>
</tr>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Active Fire Data within 24 hours (MODIS)</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">https://clck.ru/bVQGT</span></td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugin loads to QGIS data on the latest wildfires in the world over the past 24 hours, detected according to MODIS data</span></td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://github.com/Themapworkshop/Active-Fire-Plugin">https://github.com/Themapworkshop/Active-Fire-Plugin</a></span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.0</span></p>
<p>&nbsp;</p>
<p>&nbsp;</td>
</tr>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>North Australia &#038; Rangelands Fire Information (NAFI) Map Services</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">https://clck.ru/bVQGZ</span></td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugin loads a set of mapping services that display fire-affected areas and burning fire sites in Australia</span></td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://github.com/nafi-org/nafi-qgis">https://github.com/nafi-org/nafi-qgis</a>;</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://firenorth.org.au/nafi3/">https://firenorth.org.au/nafi3/</a></span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.4.0</span></td>
</tr>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Manejo</strong> (Managing Forest Areas Under The Power Lines Network)</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">https://clck.ru/bVQGe</span></td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugin is for managing the forest areas under power lines; allows projecting lines according to reference points; analyzing the territory using buffers</span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://github.com/romariocarvalhoneto/Manejo">https://github.com/romariocarvalhoneto/Manejo</a></span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.0.0</span></p>
<p>&nbsp;</td>
</tr>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>qgis2fds</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://plugins.qgis.org/plugins/qgis2fds/">https://plugins.qgis.org/plugins/qgis2fds/</a></span></p>
<p>&nbsp;</p>
<p>&nbsp;</td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugin exports the raster surface to NIST FDS modeling system (<a style="color: #000000;" href="https://pages.nist.gov/fds-smv/">https://pages.nist.gov/fds-smv /</a>) for subsequent modeling of fire dynamics, gas and heat flows</span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">https://github.com/firetools/qgis2fds;</span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://pages.nist.gov/fds-smv/">https://pages.nist.gov/fds-smv/</a></span></p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.14–3.16</span></td>
</tr>
<tr>
<td width="141"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FireHunter</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://plugins.qgis.org/plugins/firehunter/">https://plugins.qgis.org/plugins/firehunter/</a></span></p>
<p>&nbsp;</td>
<td width="208"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugin creates a Sentinel-2 mosaic for a rectangular area in a given time interval</span></td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;"><a style="color: #000000;" href="https://github.com/olegruk/firehunter">https://github.com/olegruk/firehunter</a></span></p>
<p>&nbsp;</p>
<p>&nbsp;</td>
<td width="142"><span style="font-family: 'times new roman', times, serif; color: #000000;">3.16</span></p>
<p>&nbsp;</td>
</tr>
</tbody>
</table>
</div>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>RESULTS AND DISCUSSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A group of three external plugins (Active Fire, NAFI Fire Maps and Manejo) was selected for testing in QGIS version 3.18.1.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Two layers of shp-format in geographical coordinates WGS84 “MODIS C6 1 km” (Moderate Resolution Imaging Spectroradiometer) with a spatial pixel resolution of 1000 m and «VIIRS 375 m» (Visible Infrared Imaging Radiometer Suite) with a spatial resolution of 375 m are available for download in the <strong><em>Active Fire </em></strong>plugin. Both sources are well known in scientific and applied research for monitoring wildfires (forest fires). Their descriptions are available at https://clck.ru/bVQGp.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong><em>NAFI</em></strong> <strong><em>Fire Maps</em></strong> plugin allows uploading to the QGIS desktop project up-to-date and archived data (retrospectively up to 2000 inclusively) on wildfires in Australia, one of the continents with a constant long-term wildfire hazard. Fire scars and cleaned up hotspots, distributed by months and years, are included in the sets available. Fire data are accompanied by a block of general geographic layers within the area of interest mainly in northern Australia (NAFI base layers) and Google layers (a standard set of streets, satellite base map and their hybrid) for the whole world. The data are available in the Mercator projection as WMS and WMST services. The set of project data is presented as a web-GIS application hosted at <a style="color: #000000;" href="https://firenorth.org.au/nafi3/">https://firenorth.org.au/nafi3/</a>with zooming, changing the legend, saving and printing options for the selected map part (Fig. 2). Downloading a data file in kml-format for viewing, for example, in Google Earth, is also possible as an option. Archived data 2000–2020 can be downloaded in raster (geotiff) and vector (shp) formats at <a style="color: #000000;" href="https://firenorth.org.au/nafi3/views/data/Download.html">https://firenorth.org.au/nafi3/views/data/Download.html</a>.</span></p>
<div id="attachment_4902" style="width: 1090px" class="wp-caption aligncenter"><img aria-describedby="caption-attachment-4902" loading="lazy" class="size-full wp-image-4902" src="https://jfsi.ru/wp-content/uploads/2022/03/Figure-2.jpg" alt="" width="1080" height="567" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Figure-2.jpg 1080w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-2-300x158.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-2-1024x538.jpg 1024w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-2-150x79.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/03/Figure-2-768x403.jpg 768w" sizes="(max-width: 1080px) 100vw, 1080px" /><p id="caption-attachment-4902" class="wp-caption-text"></span> <span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Figure 2.</strong> View and data operation options in web GIS application of North Australia &#038; Rangelands Fire Information (NAFI) project</span></p></div>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Manejo (Managing Forest Areas Under The Power Lines Network, <a style="color: #000000;" href="https://plugins.qgis.org/plugins/manejo/">https://plugins.qgis.org/plugins/manejo/</a>), is a plugin for managing forest areas located under power lines. It creates a network of lines from points (power towers), then makes buffer and protective zones. Screenshots of test results are in the Table 2 (repository at <a style="color: #000000;" href="https://plugins.qgis.org/">https://plugins.qgis.org/</a> as of April 8, 2021).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Table 2. </strong>Test results for external QGIS plugins for forest resource management and forest fire monitoring</span></p>
<div style="overflow-x: auto;">
<table style="border: 1px #f1f1f1 solid; background-color: #ffffff;" width="652">
<tbody>
<tr>
<td width="75"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Plugin </strong></span></td>
<td width="227"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Plugin interface</strong></span></td>
<td width="350"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>View of the results screen</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></td>
</tr>
<tr>
<td width="75"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Active Fire </strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></td>
<td width="227">&nbsp;</p>
<p>&nbsp;</p>
<p>&nbsp;</p>
<p><span style="color: #000000;"><img loading="lazy" class="aligncenter size-full wp-image-4906" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS.jpg" alt="" width="317" height="184" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS.jpg 317w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS-300x174.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS-150x87.jpg 150w" sizes="(max-width: 317px) 100vw, 317px" /></span></p>
<p>&nbsp;</td>
<td width="350"><span style="color: #000000;"><img class="aligncenter size-full wp-image-4905" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®.jpg" alt="" width="1027" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®.jpg 1027w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®-300x85.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®-1024x290.jpg 1024w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®-150x43.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®-768x218.jpg 768w" sizes="(max-width: 1027px) 100vw, 1027px" /></span></p>
<p><span style="color: #000000;"><img class="aligncenter size-full wp-image-4904" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®_Ігђ.jpg" alt="" width="658" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®_Ігђ.jpg 658w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®_Ігђ-300x211.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_ActiveFire_MODIS_C6_and_VIIRS_бЂЃ®_Ігђ-150x106.jpg 150w" sizes="(max-width: 658px) 100vw, 658px" /></span></td>
</tr>
<tr>
<td width="75"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>NAFI Fire maps</strong></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></td>
<td width="227">&nbsp;</p>
<p><span style="color: #000000;"><img class="aligncenter size-full wp-image-4907" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ѓ°й••.jpg" alt="" width="294" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ѓ°й••.jpg 294w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ѓ°й••-207x300.jpg 207w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ѓ°й••-104x150.jpg 104w" sizes="(max-width: 294px) 100vw, 294px" /></span></p>
<p>&nbsp;</td>
<td width="350"><span style="color: #000000;"><img loading="lazy" class="aligncenter size-full wp-image-4908" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†.jpg" alt="" width="1361" height="705" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†.jpg 1361w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†-300x155.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†-1024x530.jpg 1024w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†-150x78.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_Ґ®§_&#x2122;†ав_®≠в•ад•©б†-768x398.jpg 768w" sizes="(max-width: 1361px) 100vw, 1361px" /></span></p>
<p><span style="color: #000000;"><img loading="lazy" class="aligncenter size-full wp-image-4909" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_з†бвЃв†ПЃ¶†аЃҐ2000_2020.jpg" alt="" width="898" height="484" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_з†бвЃв†ПЃ¶†аЃҐ2000_2020.jpg 898w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_з†бвЃв†ПЃ¶†аЃҐ2000_2020-300x162.jpg 300w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_з†бвЃв†ПЃ¶†аЃҐ2000_2020-150x81.jpg 150w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_NAFIfiremaps_з†бвЃв†ПЃ¶†аЃҐ2000_2020-768x414.jpg 768w" sizes="(max-width: 898px) 100vw, 898px" /></span></td>
</tr>
<tr>
<td width="75"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Manejo</strong></span></td>
<td width="227"><span style="color: #000000;"><img loading="lazy" class="aligncenter size-full wp-image-4910" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б.jpg" alt="" width="312" height="428" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б.jpg 312w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б-219x300.jpg 219w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б-109x150.jpg 109w" sizes="(max-width: 312px) 100vw, 312px" /></span></td>
<td width="350"><span style="color: #000000;"><img loading="lazy" class="size-full wp-image-4903 aligncenter" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO.jpg" alt="" width="244" height="234" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO.jpg 244w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO-150x144.jpg 150w" sizes="(max-width: 244px) 100vw, 244px" /></span></p>
<p><span style="color: #000000;"><img loading="lazy" class="size-full wp-image-4911 aligncenter" src="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б2.jpg" alt="" width="287" height="461" srcset="https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б2.jpg 287w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б2-187x300.jpg 187w, https://jfsi.ru/wp-content/uploads/2022/03/Т†°Ђ®ж†2_PowerLinesNetworkForestAreas_MANEJO_®≠в•ад•©б2-93x150.jpg 93w" sizes="(max-width: 287px) 100vw, 287px" /></span></p>
<p><span style="font-family: 'times new roman', times, serif; color: #000000;">   </span></td>
</tr>
</tbody>
</table>
</div>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The plugins published are very heterogeneous in terms of number and content, as shown in the Tables 1 and 2. It is difficult to make any assumptions about trends in the development and functionality of tools for monitoring wildfires (forestfires) and forest resources management. The shp-file remains as a working format for the desktop QGIS in these plugins. The data are available in WMS format, which could be useful to view the results of plugins implementation in other GIS applications. The most commonly used coordinate system for the world coverage data is WGS84. For the continents and countries some specific systems are used, for instance, an Australian coordinate system. An ability to access archived data is an advantage for solving forestry issues. In this sense, the NAFI Fire Maps project for Australia may serve as example as it provides access to the archives and current data. In this project QGIS plugin has become a continuation and extension of the web-GIS section (https://www.gaiaresources.com.au/fire-mapping-qgis-plugin/).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">At the end of the discussion we have to note that there are still few examples in the QGIS-plugin repository for operating data on the forests, forest fires, resources and infrastructure. Notably, as of April 13, 2021, the «forest» query (https://clck.ru/bVQFY) in the repository resulted in a list of 12 plugins of different status. The «resources» query (<a style="color: #000000;" href="https://plugins.qgis.org/search/?q=resources">https://plugins.qgis.org/search/?q=resources</a>) &#8211; 5 plugins, of which one relates to the forest resources (<a style="color: #000000;" href="https://plugins.qgis.org/plugins/ontario_efri_treelist_generator/">Ontario EFRI Treelist Generator</a>). Numerous transport, infrastructural and environmental applications are related to the present research. The «routing» query (<a style="color: #000000;" href="https://plugins.qgis.org/search/?q=routing">https://plugins.qgis.org/search/?q=routing</a>) shown a list of 31 plugins; the «ecology» query (<a style="color: #000000;" href="https://plugins.qgis.org/search/?q=ecology">https://plugins.qgis.org/search/?q=ecology</a>) — 23 plugins; the «transport» query (<a style="color: #000000;" href="https://plugins.qgis.org/search/?q=transport">https://plugins.qgis.org/search/?q=transport</a>) — 8 plugins; the «forest» query (<a style="color: #000000;" href="https://plugins.qgis.org/search/?q=forestry">https://plugins.qgis.org/search/?q=forestry</a>) — 6 plugins.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>CONCLUSION</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The niche of economic and infrastructural forest applications in the up-to-dated repository of QGIS-plugins is relatively limited. For example, the plugins mentioned in the article are out of the list of first 20 popular and downloadable plugins at <a style="color: #000000;" href="https://plugins.qgis.org/plugins/popular/">https://plugins.qgis.org/plugins/popular/</a> (for the library out of total of 1.380 plugins as of April 08, 2021). Among the actual examples for the forestry, FireHunter plugin can be cited. It was introduced in April 2021 and occupies the 4th line in the list of new plugins (<a style="color: #000000;" href="https://plugins.qgis.org/plugins/fresh/">https://plugins.qgis.org/plugins/fresh/</a>) out of 19 as of April 8, 2021.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">A way for further research may be a plugins development with linked and pre-processed data sets for a continent, region, or a country administrative unit. The plugins promising for transport modeling in the forestry are Forest Road Designer (https://clck.ru/bVQEn), as well as Forest Roads Network (https://clck.ru/bVQFT) and Road Emission Calculator (<a style="color: #000000;" href="https://plugins.qgis.org/plugins/RoadEmissionCalculator/">https://plugins.qgis.org/plugins/RoadEmissionCalculator/</a>). We are planning to devote a separate study to test these plugins using data on the key forest areas in Russia.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Development, publication in the repository and testing of QGIS-plugins by users as well as QGIS-releases currently continue to be a very dynamic area where update intervals vary from a quarter to a half year, so the review of plugins functionality and performance, and their compatibility to the core QGIS application should be done within appropriate time frame.</span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>FINANSING</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">The study was performed within the framework of the state CEPF RAS assignment 121121600118-8. </span></p>
<p style="text-align: center;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>REFERENCES</strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Duarte L., Silva P., Teodoro A. C<em>.,</em> Development of a QGIS Plugin to Obtain Parameters and Elements of Plantation Trees and Vineyards with Aerial Photographs, <em>International Journal of Geo-Information,</em> 2018, Vol. 7, No 3, P. 109. DOI: <a style="color: #000000;" href="http://dx.doi.org/10.3390/ijgi7030109">10.3390/ijgi7030109</a></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Government Order, 20.09.2018, Nо 1989-r (edited 28.02.2019).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Jung M., LecoS — A QGIS plugin for automated landscape ecology analysis<em>, PeerJ PrePrints</em>, 2013, pp. 1–10, DOI: 10.7287/peerj.preprints.116v2</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Korosov A. V., Zorina A. A., <em>Jekologicheskie prilozhenija Quantum GIS: uchebnoe posobie dlja studentov biologicheskih special&#8217;nostej</em> (QGIS environmental applications), Petrozavodsk: PetrGU, 2016, 211 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Krylov A. M., Vladimirova N. A., Malahova E. G., Ispol&#8217;zovanie svobodnyh GIS v sisteme distancionnogo lesopatologicheskogo monitoringa (Open Source GIS in the remote forest pathology monitoring system), <em>Lesnoj vestnik, </em>2012, No 1. pp. 148–152.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Lovelace R., Open source tools for geographic analysis in transport planning, <em>J. Geogr. Syst.</em>, 2021, pp. 1–32, DOI: 10.1007/s10109-020-00342-2 (2021, 12 May).</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Podolskaia E., <em>Osnovy raboty v geoinformacionnom prilozhenii Open Source QGIS: geodannye, koordinaty, bazovaja funkcional&#8217;nost&#8217;, kontrol&#8217; kachestva dannyh, oformlenie i publikacija proekta</em> (Basics of working in Open Source QGIS: geodata, coordinates, basic functionality, data quality control, design and publication of the project), Lab-publisher, 2020, 52 p. </span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;">Podolskaia E., Ershov D., Kovganko K., Comparison of data sources on transport infrastructure for the regional forest fire management. Reyer, C., Bilogub, M., Mahnken, M., Gutsch, M., Kruger, K., Ramming, A., Reineking, B., Seidl, R., Schelhaas, M.-J., Makela, A., Verkerk, H. (Eds.), <em>Managing forests in the 21<sup>st</sup> century: Book of abstracts, Managing forests in the 21<sup>st</sup> century, Conference at the Potsdam Institute for Climate Impact Research,</em> Potsdam, 2020, 59 p., DOI: org/10.2312/pik.2020.002</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><em>QGIS and applications in agriculture and forest</em>, Baghdadi N., Mallet C., Zribi M. (Eds.), Vol. 2, Wiley-ISTE, 2018, 364 p.</span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong> </strong></span></p>
<p style="text-align: justify;"><span style="font-family: 'times new roman', times, serif; color: #000000;"><strong>Reviewer:</strong> Candidate of Technical Sciences O. V. Perfileva</span></p>
<p style="text-align: justify;">
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