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Competition factors of edificator tree stand: Quantitative analysis and synthesis
Abstract
To analyze and quantitatively estimate the contribution of different factors of competition from the edificator tree stand to its effect on plants comprising the lower forest vegetation layer, a set of ecophysiologically based indices of root, light, and integrated competition has been proposed and tested. The results obtained in pine and spruce forests forests of Western Siberia and the Urals show that the growth of the conifer undergrowth is more closely correlated with the index of root competition, and that of heather (Calluna vulgaris (L.) Hull.), with the index of light competition from the edificator tree stand. Moreover, the correlation of their growth with the integrated competition index is 15–25% stronger than the correlation with the indices of root and light competition, irrespective of forest type.
... The comparative analysis of such parameters as projective cover, growth, vitality, and ecologic range of heather populations was performed for different types of geographically vicarious pine forests in Pritobolye (Zavodouspenskoe) and Baltic (Luga). As a result of "coenopopulation-based and microecosystemic" regression analysis, the same objects were used to compare the regional peculiarities of heather response to the tree stand, root, and light competition indexes [27]. ...
The article presents the results of the study of fifty populations of common heather (Calluna vulgaris (L.) Hull) collected throughout its range. A phased comparative analysis (genetic, biochemical, anatomical, morphological, and ecological) was carried out with the estimation of indicators that included two key populations—Zavodouspenskoe (Pritobolye, Western Siberia) and Luga (Baltic, Eastern Europe). It was concluded that heather growing in Western Siberia should be identified as a separate taxonomic group, giving it the status of a subspecies. The gene pool of Pritobolye populations (including Zavodouspenskoe) is represented by the completely dominant (100%) monohaplotype S, which is not found anywhere else. The heather plant growing in Zavodouspenskoe has a longer lifespan. It is distinguished by larger linear leaf dimensions (length 2.06 ± 0.09 mm), thicker cuticle (4.77 ± 0.33 μm), increased number of trichomes (18.98 ± 0.56), and a reduced number of stomata (13.60 ± 0.63) than that growing in Luga. The new subspecies differs in biochemical composition: twice less content of epicatechin (average 1.992 ± 0.005 mg g⁻¹), three times more myricetin (average 2.975 ± 0.005 mg g⁻¹), twice as much chlorogenic acid (average 2.763 ± 0.004 mg g⁻¹). An ecological feature is that C. vulgaris does not grow in the swamps of Western Siberia and has a small population. This species has a high horticultural potential and requires protection as its population in Western Siberia continues to decline rapidly.
... The results of comparative studies on C. vulgaris in geographically vicarious types of pine forests in the Tobol region and Russian Plain have shed light on cenoecogeographic features of its ecological range, the structure of cenopopulations, and their heliophilia and competitive relationships with the edificator (Pinus sylvestris) tree stand (Petrova et al., 2009(Petrova et al., , 2010Sannikova et al., 2012). It has been shown that the growth rate, abundance, and coverage of C. vulgaris under the canopy of similar pine forest types are higher in the Russian Plain than in Western Siberia. ...
Analysis of geographic variation of chloroplast DNA in the eastern European and western Siberian (Tobol) groups of Calluna vulgaris populations isolated from each other for a long period of time has revealed distinct differentiation between them. It has been shown that three populations growing in the Russian Plain and Karelia share two haplotypes characteristic of central Europe, while three Tobol populations contain one haplotype F occurring in Sweden. These results confirm the hypothesis of genetic divergence between the Tobol and eastern European groups of C. vulgaris populations.
Под пологом кедровника травяно-зеленомошного в южной части Северного Урала с использованием ранее предложенного комплекса индексов световой, корневой и интегральной конкуренции древостоя проведен сравнительный анализ параметров подроста кедра 20-30-летнего возраста, произрастающего непосредственно на почвенном субстрате с моховым покровом и на микроповышениях, образованных лежащими на поверхности почвы полуразложившимися древесными стволами (валеже). На рост подроста кедра, произрастающего на почвенном субстрате, оказывают отрицательное комплексное влияние два основных определяющих фактора: световая конкуренция древостоя (уровень перехвата его пологом фотосинтетически активной радиации) и, почти равная ей, но несколько ниже, корневая конкуренция за почвенное питание. Теснота и достоверность отрицательной связи (R = 0,72) средних за последние пять лет приростов терминальных побегов с комплексным индексом интегральной конкуренции древостоя превышает связи (R = 0,47-0,54) с частными индексами этих факторов в 1,3-1,5 раза. На древесном валеже, где подрост кедра до определенной стадии своего развития находится вне корневой конкуренции прослеживается отрицательная тесная связь (R = 0,71) приростов терминальных побегов с влиянием световой конкуренции древостоя, близкая по своему уровню с комплексным влиянием световой и корневой конкуренции на рост подроста на почве. На обоих типах субстрата максимальные средние приросты терминальных побегов подроста кедра (10-15 см) наблюдаются при значении индекса световой конкуренции древостоя не более 20. С увеличением его значения в диапазоне 20-80 величина средних приростов сокращается в 10-15 раз до 0,5-1 см. Но при одинаковом значении индекса световой конкуренции параметры подроста кедра, произрастающего на валеже, примерно в 1,5 раза выше, чем подроста на почвенном субстрате, что также иллюстрирует отрицательное влияние на него корневой конкуренции материнского древостоя.
Under the canopy of the grass-greenmoss Siberian stone pine forest in the southern part of the Northern Urals, using the previously proposed complex of light, root, and integral competition indices of the stand, a comparative analysis of the parameters of the Siberian stone pine undergrowth 20-30 years old, growing directly on the soil substrate with moss cover and on micro-elevations, formed by semi-decomposed tree steams, lying on the surface of the soil (dead fallen tree steams). Two main determining factors have a negative complex effect on the growth of the Siberian stone pine undergrowth, growing on soil substrate: the light competition of the stand (the level of photosynthetically active radiation interception by its canopy) and, almost equal to it, but slightly lower, the root competition for soil nutrition. The tightness and reliability of the negative relationship (R = 0.72) of the average over the past five years growth of terminal shoots with a complex index of integral competition of the stand exceeds the relationship (R = 0.47-0.54) with private indices of these factors of 1.3-1.5 times. On the dead fallen tree stems, where the Siberian stone pine undergrowth, until a certain stage of its development, is outside the root competition, a negative close relationship (R = 0.71) of the growth of terminal shoots with the effect of light competition of the stand, close in level with the complex effect of light and root competition on the growth of its undergrowth on soil, is traced. On both types of substrate, the maximum average growths of the terminal shoots of Siberian stone pine undergrowth (10-15 cm) are observed when the index of light competition of the stand is no more than 20. With an increase in its value in the range of 20-80, the average growth rate decreases by 10-15 times to 0.5-1 cm. But with the same value of the light competition index, the parameters of the undergrowth, growing on dead fallen tree steams, are about 1.5 times higher than the undergrowth on the soil substrate, which also illustrates the negative effect of the maternal stand root competition on it.
In the forests near Perm, 184-year-old stands of common pine of class I Bonita were studied with a fullness of 0.93 and a reserve of 620 m3/ha. On an area of 2.2 hectares, live and dead trees over 60 years were mapped to the plan with an accuracy of ±10-35 cm. Then, using phyto-and bioindication (biolocation), more than 1200 pieces were chart small geoactive zones (hereinafter referred to as zones) with geo-linking to the centers of the bases of the trunks of nearby trees. They were classified into 7 types: pathogenic (Hartman and Kurri networks), favorable in size 1.0 and 3.0 m, and neutral in size 8.0, 16.0, and 32.0 m. The interaction of trees was divided into six levels, for which 5 map layers were formed. The Windows included places with a distance between trees of 8.76 m or more, where the interaction of trees decreased to 0.16 from the average level. It was found that Windows occupy 41% of the area and the reasons for their formation in 100% of cases were four factors. The first is the absence of favorable zones of 1.0 and 3.0 m in size: 94% of cases of window formation where there were no such zones. The second is the effect of pathogenic zones and depression zones of other zones: 39% of cases; they inhibited the growth of trees and led to their fall, but the proportion of their area in the windows was the same as in the whole stand. The third is the settlement of undergrowth on favorable zones, which may have prevented pine trees from settling on them in the past: 6% of cases. The fourth is the decline due to competition, and the trees were in neutral places without the energy from favorable zones that their neighbors received: 3% of the cases of Windows. That is, the "competitive struggle" was not equal, and a small or large area of nutrition in trees did not play a role in this struggle.
Larix chinensis Beissn forms a subalpine timberline on Taibai mountain, Shaanxi, China. Seed emergence and survival of seedlings were studied by seed sowing experiments across an altitudinal gradient (3150–3400 m a.s.l). Three main factors were considered: seed addition, shrub shadowing and disturbance. In three different habitats (from low altitude to high: the Abies fargesii forest, the Larix chinensis forest and the shrub meadow), seeds emergence was examined and the seedling number and growth in every plot were recorded continuously over 4 years. Kruskal-Wallis ANOVA was used to analyze the relationship between seedling emergence/recruitment and the explanatory variables in every stand. The results showed that seed addition in disturbed plots always caused significantly increased seedling emergence and recruitment in the A. fargesii forest and the L. chinensis forest, but not in undisturbed plots under the canopy of forests. It demonstrated that recruitment of L. chinensis was limited by seed and microsite availability under closed forest, and litter is one of the factors leading to microsite limitation. However, in the shrub meadow, no treatments improved recruitment, indicating that the shrub meadow was not suitable for seedling establishment because of more intense light and lower temperature. In the L. chinensis forest and the A. fargesii forest, more recruited seedlings indicated that these two forests are safer for L. chinensis seedling establishment, though seeds limitation and the litter obstacle had seriously constrained the regeneration of L. chinensis.
A new method for determining the upper forest border (UFB) as the border of the “root-closed” stand and regrowth as its potential vanguard is proposed. The mass dispersal of Pinus sibirica populations by the nutcracker (Nucifraga caryocatactes) in the UFB ecotone in the Northern Urals (Pavdinskii Kamen’ and Tretii Bugor Mountain) from the middle-mountain taiga to the mountain forest tundra and tundra at a distance reaching 1 km is found. It is done based on the original reconstruction method of the dynamics of population number in P. sibirica annual seedlings and on an analysis of their relations with the dynamics of the seed bearing and population number of the nutcracker. The trustworthy consortive relations of the number of Pinus sibirica generations in the mountain forested tundra and tundra zones, with its seed bearing and nutcracker number in the previous year in the middle-mountain taiga zone, are found. A significant increase, 3.7 times, in the number of seedling generations during the last 49 years is found in connection with the increase in the summer air temperature in the subarctic to 2.0°C. A hypothesis of the formation of the united genetic metapopulation of P. sibirica in the forest–tundra mountain ecotone zone as a consequence of its seed ornitochory from the highly different populations is formulated. A forecast of the mosaic formation of the vanguard “thin forests” in 20–25 years and the root-closed curtains of the forest in 40–50 years in the mountain tundra of the Northern Urals at the current rate of the rise in temperature is made.
On the basis of the system of ordination for the geographic zonal and provincial replacement of types of pine forests, gradients, and some geographical trends of the main parameters of the morphostructure, the growth and viability of the cenopopulations of common heather Calluna vulgaris (L.) Hull between various subzones of the Russian Plain and the western part of Western Siberia (Tobolsk area) have been revealed.
The study discusses two approaches to predict e effect the effect of neighboring trees on the 5-year basal area growth of Scots pine. The first approach relies on empirical competition indices and the second on an ecological field theory. The growth models account for 82-84% of the within-stand variation in basal area growth. The directional distribution of competitors has a slight influence on the tree growth, besides the number, size and distance of the neighbors. To demonstrate the use of spatial growth models, three simulation studies are presented. The first one examined the effect of the spatial pattern of trees on the volume increment of the stand, the second one the usability of variation in the stand basal area as a measure of grouping, and the third one the effects of harvest roads.
Theoretically possible forms of the dependence of phytogenic field strength on the distance between the measurement point and the effecting tree were examined. Four possible dependence types were described: a hyperbolic curve, a logistic curve, a curve with a maximum at some distance from the trunk, and graduated changes. Parameters describing the strength of tree fields in lichen-green moss pine forests of Karelia and the St. Petersburg district were obtained on the basis of the hypothesis of the additivity of phytogenic field interference. Phytogenic fields of pine trees can be described by two strengths-shadow and trunk ones. Both of these strengths correspond to a hyperbolic dependence curve and increase upon an increase in the dimensions of the effecting tree. For interfering tree phytogenic fields, strengths are summed. The abundance of species of the herbaceous-dwarf shrub and moss-lichen layers depends linearly on strength logarithms. Numerical values of the parameters of these dependences were obtained, making it possible to predict the status of the ground vegetation based on the dimension and location of surrounding pine trees.
Zusammenfassung Die Abhängigkeitsbeziehung zwischen drei Konkurrenzvariablen und dem Volumenzuwachs vonP. radiata wurde untersucht, nämlichProdan-Ehrlenspiels n-Baum-Stichprobe (Bantre
i), eine Abwandlung vonSpurrs Modifikation der Winkelzählprobe (bangtree
i) undKennels Abstandsvariable (distree
i). Die Ergebnisse waren folgende:
1.
In den undurchforsteten A-Flächen ergabbangtree
i die höchste Regressionsquadratsumme, in den stark durchforsteten C-Flächen dagegendistree
i.
2.
Die Regressionsquadratsumme ist vom Wert des Index i abhängig.
3.
Für dieselbe Baumhöhe und denselben Wert der Konkurrenzvariablenbangtree
4 ist der Zuwachs in stark durchforsteten Beständen größer als in schwach durchforsteten.
A distance-dependent individual tree based growth model (FOREST) was compared with a diameter-class growth model (SHAF) for describing changes in stand density and structure. Projections of Lake States' northern hardwood stand development were made by each model for 5–26 years over a range of stand conditions and harvest treatments. Results from numerous performance tests and comparisons of actual and predicted diameter distributions, basal areas, and numbers of trees, indicate the individual tree model was considerably more sensitive to harvest treatments and reproduction response than the diameter-class model. Conversely, the latter was much less expensive to operate. Prediction of species and individual tree growth with the individual tree model appeared to provide sensitivity nearly equal to that observed for predictions of the stand as a whole. Long-term projections (120 years) for reserve (no cut) and clear-cut stand conditions further suggest the potential and limitations of the models for management analyses.
Competition indices, based on the distance weighted ratio of competitor and subject tree sizes, were correlated with annual diameter and height growth. Correlations with growth were greater when competitors were chosen by angle gauge techniques than by a fixed competition radius. Some of the tested indices were as highly correlated with tree growth as the complex area overlap and weighted area overlap competition indices used in the past. Correlation with growth and computational ease indicate that simple competition indices may be well suited to tree growth simulation and field applications. Forest Sci. 22:454-456.
The microecosystem approach and an adequate system of quantitative methods have been used to study the relationship between the chorologic changes in the structure and functions of edificator tree stand and other components of the pine (Pinus sylvestris L.) forest cenoecosystems of the Transural and Northern Turgai regions. Significant correlations of the density of pine undergrowth with the litter thickness; of its growth with root competition, stand closeness, and, to a lower degree, photosynthetically active radiation (PAR); and of the numbers of small mammals and groups of soil mesofauna with herbaceous phytomass have been found. The possibility of the synthesis of population-ecological, synecological, and biogeocenological methods has been demonstrated.
A graphical method is discussed which allows a specification of the optimal diet of a predator in terms of the net amount of energy gained from a capture of prey as compared to the energy expended in searching for the prey. The method allows several predictions about changes in the degree of specialization of the diet as the numbers of different prey organisms change. For example, a more productive environment should lead to more restricted diet in numbers of different species eaten. In a patchy environment, however, this will not apply to predators that spend most of their time searching. Moreover, larger patches are used in a more specialized way than smaller patches.
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Basic Concepts of Forest Biogeocenology
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Microecosystem Approach in Population Ecology and Biogeocenology
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Basic Ecology, Philadelphia: Saunders, 1983. Translated under the title Ekologiya
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