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HayKOBMM BicHkiK H/lTyyKpaÏHM Scientific Bulletin of UNFU
http://nv.nltu.edu.ua
https://doi.org/10.15421/40270808
Article received 26.10.2017 p. Article accepted 24.11.2017 p.
UDC 630*22.8(292.451/.454)
ISSN 1994-7836 (print) ISSN 2519-2477 (online)
[^1 Correspondence author V. I. Parpan parpan@il.if.ua
T. V. Parpan, V. I. Parpan
The Vasyl Stefanyk Precarpathian National University, Ivano-Frankivsk, Ukraine
ECOLOGICAL STRUCTURE OF BEECH AND CONIFEROUS/BEECH MOUNTAIN CLIMAX FOREST STANDS OF UKRAINIAN CARPATHIANS
According to the age classes, middle-aged forest stands predominate in the mountan forests of the Carpathians (46.2), overmature stands occupy 6.9% (62 thousand ha). According to the recent inventory, about 85 thousand hectares of climax old-growth forests are identified, including climax virgin forests that occupy about 54 thousand hectares which are uneven-aged. For their identification, a population matrix with a division into age groups is suggested, taking into account the age condition of the edificator. There are 6 age groups: pre-generative - juvenile-immature, virginal; generative - young generative, middle-aged, mature and old generative. In terms of the number of uneven-aged beech and coniferous/beech stands, the adequate cenopopulations of beech is dominated. The age spectra of fir and spruce, sycamore maple, ash-tree are generally non-stabilized or fragmentary. On average, on 1 ha of elementary demographic cenopopulation of beech and coniferous/beech stands on the generative generation there are 174-235 beech individuals, 42-52 - fir, 5-16 - spruce, 5-10 - sycamore and ash-tree, respectively.Half of the timber stock (50-60%) is over-mature, and 25% is in mature age condition. In climax cenosis, it can be recommended to conduct a uniform or non-uniform selection felling system. Introduction of population paradigm in forest synecology opens up a new opportunity to divide stands into age generations and treat them as a set of cenopopulations of key species for old-growth forests and virgin forests.
Keywords: typology; population; structure; beech stand; coniferous/beech stand.
Introduction
About eleven basic formations of edificators have been developed in the phylogenesis process in the mountainous conditions of the Ukrainian Carpathians: Picea abies, Abies alba, Fa-gus sylvatica, Quercus robur, Quercus petraea, Pinus sil-vestris, Alnus glutinosae, Alnus incana, Pinus mughus, Alnus viridis, Juniperus sibirica. In quaternary period their genesis was closely connected with the development of the natural landscapes of Central Europe, and during the last millennium a modern structure of forest vegetation was formed and partially changed depending on the climatic conditions of the region (Kalinovich, 2003; Stoyko, 1985). During this period, subformational and typological-cenotic structure of the forest cover was formed in the different high-altitude climatic zones of the southwestern and northeastern macroslopes of the Carpathians within the locally differentiated soil-water-geochemical conditions of the trophotope. In the last two centuries there have been large-scale changes in the structure of stands as a result of human economic activity, which has disrupted the stability of mountain forests and the performance of their ecological functions (Golubets, 2012; Parpan, Parpan & Hudyma, 2016; Tretiak & Chernevyj, 2015). The changes involved primarily the stand which is an object of manage-
ment and, in the forest ecosystem, ensures its functioning (Golubets, 2007; Krynytskyi, 2012).
The purpose of this publication is to analyze the typological structure of mountain ecosystems at the level of the subformation as an integral evolutionary natural historical process for the formation of forest cover in the mountains, as well as the analysis of the cenopopulation structure of climax beech and coniferous/beech stands of different ages according to silvicultural and taxonomic, and population-ecological methodological principles (Volosyanchuk, Prot-sia & Kahala, 2015; Markiv, 1982).
Material and methods
The principles of silvicultural and ecological typology are used in the research: forest formation and subformation, coordination grid (Golubets, 2007, 2012), type of forest conditions, type of forest and type of stand (Vorobiev, 1967; Gerushinsky, 1996; Parpan, Parpan & Hudyma, 2016; Stoyko, 1985; Cherniavskyi, Krynytskyj & Parpan, 2011).
The age structure of the stands was identified according to the age groups adopted in forest science (Pogrebnyak, 1963) and the proposed population-ecology approach (Smirnova, 2011) which uses the periodization of ontogeny (age status) of the edificator (Rabotnov, 1964). This peri-odization was elaborated in detail for key species such as Quercus robur (Smirnova, 1994, 2004), Fagus sylvatica (Chistyakova & Parpan, 1991), Picea abies (Parpan, 1993),
^TyBaHHfl 3a flCTY: Parpan T. V., Parpan V. I. Ecological structure of Beech and Coniferous/Beech mountain climax forest stands of
Ukrainian Carpathians. HayKOBMM BicHMK HIITy yKpaÏHM. 2017. Bun. 27(8). C. 59-63. Citation APA: Parpan, T. V., & Parpan, V. I. (2017). Ecological structure of Beech and Coniferous/Beech mountain climax forest stands of Ukrainian Carpathians. Scientific Bulletin of UNFU, 27(8), 59-63. https://doi.org/10.15421/40270808
Abies alba (Markiv, 1982; Molotkova, 1968) and implemented in relevant scientific papers (Smirnova, 1994, 1998, 2004; Parpan, 2014; Smirnova & Chistyakova, 1980; Chistyakova & Parpan, 1991).
A spatial area is chosen as elementary ecological (cenopopulation) unit on which all age generations are concentrated within the subcompartment, the type of water catchment site and the type of forest, calculated by the empiric method. This cenopopulation is called the minimal demographic unit (EDO), which covers an area of about 1 hectare (Parpan, 1993).
Based on the correlation between age, height and diameter of beech, fir and spruce, with the predominance of the diameter interval, we proposed a population matrix for uneven-aged coniferous/beech and beech stands according to the Forestry Age Group (Pogrebnyak, 1963) and the biological age status (Parpan, 1993) (Table 1).
13. Spruce/Beech and Spruce/Fir/Beech 69.8 30.8 100.7 9.9
14. Sycamore maple/Beech 0.0 5.9 6.0 0.6
Other forest formations 12.1 19.5 31.6 3.1
Area of forest subformations of the Ukrainian Carpathians 562.9 459.0 1021.8 100.0
Of most ecological-resource and stabilizing importance are the edificator formations: Picea abies, Fagus sylvatica, Abies alba, which account for 97 % of the area of the mountain forests. In their structure, the corresponding forest types and types of stands are highlighted.
The structure of forest stands by age groups. Due to the coupes-type felling system over the last two centuries, modern forest stand successional system has been formed in the forest cover of the Carpathians, these stands being identified by six age groups (Table 3).
Table 1. Population matrix of structure of climax beech and coniferous/beech stands
Generation* Intervals Age group Age state
A, years D, cm H, m
1/jim 10-40 go 6 0,1-2,5 Young growth Juvenile and mature
2/vi 41-80 8-12 4-14 Pole-stage stand Virginal
3/g1 81-120 16-28 16-25 Middle-aged Young generative
4/g2 121-160 32-48 25-40 Ripening Middle-aged generative
5/g3 161-200 52-64 32-40 Mature Mature generative
6/g3S 201-260 (360) 68-120 30-35 Overmature Old generative
Table 3. Age groups of forest-forming species in functional categories of forests of the Ukrainian Carpathians (within the boundaries of the State forestry agency land,
Age groups Functional categories of forests * Total
1 2 3 4 ra %
Seedling stage 10.7 5.5 13.3 37.9 67.4 7.5
Sapling stage 16.4 4.2 20.9 51.1 92.6 10.3
Middle-aged 81.7 42.8 135.5 154.1 414.1 46.2
Ripening 20.8 9.5 25.1 70.8 126.2 14.1
Mature 18.0 8.8 36.2 70.2 133.2 14.9
Overmature 12.7 2.7 21.3 25.3 62.0 6.9
Total 160.3 73.5 252.3 409.4 895.5 100
* Note: jim - juvenile and mature; v1 - virginal; g1 - young generative; g2 - middle-aged generative; g3 - mature generative; g3 s - old generative.
Results and discussion
Typological structure. Formation of spruce forests in the Carpathian forest cover occupies 36.2 %, fir - 18.6%, beech - 42.6 %. Accordingly, in their structure, there are 5, 4 and 5 subformations, respectively (Table 2).
Table 2. The subformation structure of mountain forest ecosystems of the Ukrainian Carpathians (within the boundaries of the State forestry agency land, thousand
№ P/P Basic forest subformations Macroslope Carpathians
northeast, thousand /ha southwest thousand /ha thousand /ha %
Spruce formation 268.3 101.3 396.6 36.2
1. Unmixed Spruce 80.5 16.2 96.7 9.5
2. Cedar/Fir 4.9 0.3 5.2 0.5
3. Beech/Fir 10.1 14.3 24.5 2.4
4. Spruce/Fir 10.5 1.2 11.7 1.1
5. Beech/Spruce/F ir 162.3 69.3 231.6 22.7
Fir formation 168.9 21.8 189.7 18.6
6. Oak/Fir 3.2 0.0 3.2 0.3
7. Hornbeam/Beech/Fir 6.7 0.3 7.0 0.7
8. Beech/Fir 18.9 3.3 22.2 2.2
9. Beech/Spruce/F ir and Spruce/Beech/Fir 139.2 18.2 157.4 15.4
Beech formation 114.5 316.4 430.9 42.2
10. Unmixed Beech 10.4 188.3 198.7 19.4
11. Hornbeam/Beech and Rocky Oak/Beech 4.3 79.1 83.3 8.2
12. Fir/Beech and Hornbeam/Fir/Beech 30.0 12.3 42.3 4.1
* Note: 1) nature conservation; 2) recreation; 3) protecting; 4) exploitable.
Most massive deforestation in the Carpathians took place in the 1950s and 1960s, when logging was carried out in all functional categories of forests. As a consequence, middle-aged forest stands are prevailing age groups in the current forest cover - 46.2 %. Young stands account for 17.8 % of the area, ripening and mature age groups - 14.1% and 14.9 %, respectively. Overmature stands occupy only 6.9 %, or 62 thousand hectares. In terms of the age structure, the modern Carpathian forest stands are generally divided, by convention, into even-aged and uneven-aged (Krynytsky, 2012; Tsurik, 1974). In the context of structure, uneven-aged forest stands are less studied than other ones.
According to origin, level of preservation, the rate of natural cycle development and anthropogenic influence in a modern forest cover of the Carpathians, it is proposed to distinguish:virgin forests; natural (old-growth) forests on the site of virgin forests; natural-artificial forests with permanent human intervention and artificial (culture) with constant economic activity (Parpan, 2014). According to the identification of old-growth forests and virgin forests in the Ukrainian Carpathians as of the late 2016, the forests subordinated to the State forestry agency contain about 85 thousand hectares of old-growth forests, including about 54 thousand hectares of virgin forests (Parpan, Parpan & Hudyma, 2016; Parpan & Chystiakova, 1990).
The impact of forest management and the forest stands formed by the age groups are critically evaluated in the manual "Close-to-nature forestry and multifunctional forest management in the Carpathian region of Ukraine and Slovakia", which is the result of joint intellectual work by Uk-
rainian and Slovak forest scientists and practitioners (Parpan, 2014). The manual and modern publications offer general positive principles for the introduction of a close-to-nature forestry and the possibility of its use in the Ukrainian Carpathians (Chernevyj, 2006; Cherniavskyi, Krynytskyj & Parpan, 2011).
We analyzed the age structure of the all-aged confero-us-beech and beech stands, which can serve as identification criteria and models for selection felling and re-forming of stands into complex mixed cenoses within forest types in spatially different watersheds.
Population structure of uneven-aged coniferous/beech and beech stands. The population structure of edificator species has common variants of ontogenesis, self-maintenance, distribution by age groups (generations) and specific for fir and spruce. An analysis was carried out for four forest types on 50 hectare trial plots, of which 34 belong to old-growth forests, and 16 to climax virgin forests. In connection with a significant amount of research material, averaged data, for better perception, on the number and productivity of cenopopulations according to forest types, age periods and age groups are given (Table 4, Table 5).
In all types of forest, the coenopopulation of beech is complete, the cenopopulation of fir and especially spruce in coniferous-beech types is fragmentary (gappy). In the generative period, the number of reproductive individuals of the beech varies from 174 to 235 specimens /ha and is sufficient to reproduce the juvenile generation. For sub-edificator species - fir and spruce- they are are not enough. The reserve stock of cenopopulations is the number of pregenerative individuals (virgin and juvenile-immature age group). It is also characteristic that the number of individuals of the virgin age for all species cenopopulations in coniferous-beech and beech stands is not enough, while the number for the juvenile-immature group, especially beech, is sufficient. For fir and spruce, special regulatory measures should be introduced to promote natural regeneration. From the standpoint of the population ecology, the juvenile-immature generation will be analyzed in more detail in later publications.
Table 4. The number of cenopopulations (specimen / ha) of tree species in the age groups of uneven-aged
Table 5. The productivity of cenopopulations (m / ha) of tree species in the age groups of the uneven-aged
Code, amount of samples Coeno-population * Generative period Pre-generati-ve period
Age groups (generation) *
6 5 4 3 Z 2 1
P3-SF-BE 9 samples (old-aged) Fg.s 60 50 42 83 235 96 6,377
Ab.al 1 6 12 17 42 15 733
P.ab - 2 5 11 18 18 144
A.ps - 2 6 8 16 3 4,557
P3-S-BE 13 samples (old-aged) Fg.s 39 37 53 85 214 153 1,090
Ab.al 8 11 15 18 52 116 1,992
P.ab - - 1 4 5 7 185
OK3-BE 12 samples (old-aged) Fg.s 35 37 36 121 224 326 12,206
Ab.al 1 1 2 2 8 1 2,467
P.ab - 1 1 1 6 1 2,858
OK2-3-BE 16 samples (virgin forests) Fg.s 56 48 29 41 174 105 21,240
Ab.al 3 3 2 2 10 - 6,458
Fr.ex 4 1 - - 5 - 3,725
Code, amount of samples Coeno-population * Age groups (generation) *
6 5 4 3 2 Z %
P3-SF-BR 9 samples (old-aged) Fg.s 218.6 78.4 24.4 10.4 3.1 335.0 71.0
Ab.al 3.6 33.3 17.7 7.0 0.5 61.9 13.1
P.ab - 10.2 32.5 9.7 3.1 54.3 11.5
A.ps - 1.8 13.1 5.6 0.4 20.9 4.4
P3-S-BE13 samples (old-aged) Fg.s 148.6 73.7 46.4 19.9 6.3 294.9 81.0
Ab.al 29.9 15.8 11,2 4.5 4.0 65.4 18.0
P.ab - 0.6 1.0 1.5 0.4 3.5 1,0
OK3-BE 12 samples (old-aged) Fg.s 253.2 106.0 33.1 28.0 12.9 433.3 96.4
Ab.al 1.1 3.2 3.6 1.4 - 9.3 2.0
P.ab - 0.6 5.6 0.2 0.4 6.8 1.6
OK2-3-BE 16 samples virgin forests Fg.s 330.8 142.1 38.5 20.5 7.8 539.7 98.5
Ab.al 1.2 3.9 0.7 0.2 - 6.0 1.1
Fr.ex 2.0 0.2 - - - 2.2 0.5
* Note: 1- juvenile and mature (young growth); 2- virginal; 3- young generative; 4 - middle-aged generative; 5 - mature generative; 6 -
old generative; Fg.s - beech, Ab.al -more, Fr.ex - ash tree.
fir, P.ab - spruce, A.ps - syca-
* Note: see Table 4 for designations.
A brief analysis of the productivity of the cenopopulati-ons indicates its features in coniferous/beech and beech uneven-aged stands. In coniferous-beech cenoses, a beech population of type P3-SF-BE accounts for 71 % of the stock; fir - 13,1%; spruce - 11 %, and in type P3-S-BE -81 %, 18% and 1 %, respectively. In monodominant beech forests, the cenopopulation of beech is predominant and to amounts to 96.4-98.5 % of the stock. The distribution of the stock over the age groups is such that almost half of it is concentrated in the overmature (old generative post vegetative) generation (P3-SF-BE - 47 %; P3-S-BE - 49 %; OK2-BE - 56-61 %). In the mature generative stand, the volume of timber amounts to 24-26 %; in middle-aged - 17-18 %; in young generative - 4-7 %, and in virginal - 1.5-3.0 %. For selective regulation from the standpoint of rejuvenating the cenopopulation, it is necessary to use predominantly overmature and mature age groups of beech, and, more rarely, fir and spruce.
Therefore, coniferous/beech and beech uneven-aged stands of the Carpathians include all age generations with the dominance in the number of the pre-generative generation, and in terms of productivity - the generative generation. They need to be identified as old-growth forests and virgin forest ecosystems. Only old-growth stands underwent insignificant anthropogenic impact, and the virgin forests emerged and develop naturally through the action of only natural elements and events without significant human intervention (Parpan, 1993). Both groups of forest stands are need of preservation and protection.
Conclusions
According to the age classes, middle-aged forest stands predominate in the mountain forests of the Carpathians (46.2 %), overmature stands occupy 6.9 % (62 thousand ha). According to the inventory, about 85 thousand hectares of old-growth forests are identified, including virgin forests that occupy about 54 thousand hectares (Volosyanchuk, Protsia & Kahala, 2015). According to the age structure, as shown in this publication, they are uneven-aged. For their identification, a population matrix with a division into age
groups is proposed taking into account the age condition of the edificator. There are six age groups: pre-generative - juvenile-immature, virginal; generative - young generative, middle-aged, mature and old generative. In terms of the number of uneven aged coniferous-beech and beech stands, the complete cenopopulation of beech is dominated. The age spectra of fir and spruce, sycamore maple, ash-tree, and elm are generally non-stabilized, or fragmentary. On average, on one hectare in all-aged coniferous-beech and beech stands on the generative generation there are 174-235 beech individuals, 42-52 - fir, 5-16 spruce, 5-10 - sycamore and ash-tree. Half of the timber stock (50-60 %) is over-mature, and 25 % is in mature age condition. In uneven-aged stands, a uniform or non-uniform selection felling system can be recommended. The introduction of population paradigm in forest synecology opens up a new opportunity to divide stands into age generations and treat them as a set of cenopopulations of key species for old-growth forests and virgin forests.
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Т. В. Парпан, В. I. Парпан
Прикарпатський нацюнальний утверситет iм. Василя Стефаника, м. 1вано-Франювськ, Украта
ПОПУЛЯЦ1ЙНО-ЕКОЛОГ1ЧНА СТРУКТУРА БУКОВИХ I ХВОЙНО-БУКОВИХ КЛ1МАКСОВИХ
ДЕРЕВОСТАН1В УКРАШСЬКИХ КАРПАТ У Л1СОВОМУ ПОКРИВ1
За класами вжу прських лгав Карпат переважають середньовшж деревостани (46,2%), перестшш займають 6,9% (62 тис. га). За сучасною швентаризащею видшено близько 85 тис. га юлмаксових старовжових лгав, зокрема, близько 54 тис. га ктмаксових рiзновiкових пралгав. Для 1х вдентифжацп запропоновано еколопчну матрицю з подшом на вшж групи з урахуванням вжового стану едифжатора. Видшено 6 вжових груп: догенеративна - ювшшьно^матурна i вiргiнiльна; генеративна - молода, середньовжова, стигла i стара (перестшна). За чисельшстю в рiзновiкових хвойно-букових i букових де-ревосташв домшуе повночленна ценопопулящя бука. Вшж спектри ялищ та смереки, клена-явора, ясена, шьма, зазвичай, неповночленш або фрагментарш. У середньому на одному гектарi елементарно! демографiчноl ктмаксово! ценопопуляци букових i хвойно-букових деревосташв на генеративне поколшня припадае 174-235 особин бука, 42-52 - ялищ, 5-16 - яли-ни, 5-10 - клена-явора та ясена звичайного, вдаовдао. Запас деревини на 50-60% приурочений до перестшних i на 25% -до спшого вжового стану. У ктмаксових ценозах доцшьно проводити рiвномiрно або нерiвномiрно вибiркову систему ру-
бань. Впровадження популяцшно! парадигми в люову синекологiю вiдкривае нову можливiсть членувати деревостани на вь ковi поколшня та трактувати 1х для старовжових лiсiв i пралгав як сукупшсть ценопопуляцiй ключових видiв, встановити стратепчну поведiнку.
Ключовi слова: типолопя; популящя; структура; буковий i хвойно-буковий деревостани.
Т. В. Парпан, В. И. Парпан
Прикарпатский национальный университет им. Василия Стефаника, г. Ивано-Франковск, Украина
ПОПУЛЯЦИОННО-ЭКОЛОГИЧЕСКАЯ СТРУКТУРА БУКОВЫХ И ХВОЙНО-БУКОВЫХ КЛИМАКСОВЫХ ДРЕВОСТОЕВ УКРАИНСКИХ КАРПАТ В ЛЕСНОМ ПОКРОВЕ
По классам возраста горных лесов Карпат преобладают средневозрастные древостои (46,2%), перестойные занимают 6,9% (62 тыс. га). По современной инвентаризации выделено около 85 тыс. га климаксовых старовозрастных лесов, в т.ч. около 54 тыс. га климаксовых девственных лесов разновозрастных. Для их идентификации предложена популяционно-эко-логическая матрица с разделением на возрастные группы с учетом возрастного состояния эдификатора. Выделено 6 возрастных групп: догенеративная - ювинильно-иматурная и виргинильная; генеративная - молодая, средневозрастная, спелая и старая (перестойная). По численности в разновозрастных хвойно-буковых и буковых древостоях доминирует полночленная ценопопуляция бука. Возрастные спектры пихты и ели, клена-явора, ясеня, ильма, как правило, неполночленны или фраг-ментары. В среднем на одном гектаре элементарной демографической климаксовой ценопопуляции буковых и хвойно-буко-вых древостоев на генеративное поколение приходится 174-235 особей бука, 42-52 - пихты, 5-16 - ели, 5-10 - клена-явора и ясеня обыкновенного, соответственно. Запас древесины на 50-60% приурочен к перестойному и на 25% - к спелому возрастному состоянию. В климаксовых ценозах целесообразно проводить равномерно или неравномерно выборочную систему рубок. Внедрение популяционной парадигмы в лесную синекологию открывает новую возможность членить древостой на возрастные поколения и трактовать их для старовозрастных и девственных лесов как совокупность ценопопуляций ключевых видов, наметить стратегическое поведение.
Ключевые слова: типология; популяция; структура; буковый и хвойно-буковый древостой.
