Ecological Questions 25/2017: 51–66 http://dx.doi.org/10.12775/EQ.2017.005
Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land
Mi osz eptu a n rze ienarto icz Marta icka nna ilbran t-Cza a
1LFRODXV&RSHUQLFXV8QLYHUVLW\)DFXOW\RI%LRORJ\DQG(QYLURQPHQW3URWHFWLRQ&KDLURI*HRERWDQ\ DQG/DQGVFDSH3ODQQLQJ/ZRZVND7RUXĔ3RODQG *[email protected] Received: 25 April 2017/Accepted: 05 June 2017
bstract.,QWKHVWUXFWXUHRIDIRUHVWVWDQGJURZLQJRQIRUPHUDJULFXOWXUDOODQGLQVXEXQLWQRIWKH3U]\PXV]HZR)RUHVW 'LYLVLRQ 5HJLRQDO'LUHFWRUDWHRI6WDWH)RUHVWV5'6)LQ7RUXĔ ZDVGHVFULEHG7KHVWXG\DUHDZDVDIIRUHVWHGLQ ±PRVWO\ZLWK Scots pine (Pinus sylvestris L.), silver birch (Betula pendula Roth) and several seedlings of the European beech – after many years of agricultural cultivation of grain and potato crops. Characteristics of the forest stand comprised the following parameters: species composition and species diversity, density of individual components, the average tree diameter at breast height (DBH), and the height and aboveground biomass of trees divided into individual species. The species structure, dendrometric characteristics and spatial dis- tribution of trees studied in 2015 were compared with the situation assessed in 2000. Based on DBH and height values, as well as the use of dendrometric tables and basic wood density for tree species, the above- ground biomass and total biomass of trees with a minimum diameter of 7 cm were calculated. The aboveground and total biomass for trees with DBH less than 7 cm was calculated on the basis of density and weight of trees according to the classification into species and height classes. The aboveground and total biomass of the whole tree stand, including spontaneous non-native and invasive Padus serotina (KUKZDVFRPSDUHGZLWKWKHVWDQGLQJELRPDVVRIDJULFXOWXUDODUHDVLQWKH7XFKROD)RUHVWUHJLRQDVZHOODVZLWKRWKHU6FRWV pine-silver birch plantations on former agricultural lands described in ecological literature. The paper presents also the differences in assessments of aboveground pine biomass at the study site obtained when using conver- VLRQIDFWRUVHVWDEOLVKHGGXULQJGLUHFWPHDVXUHPHQWVFDUULHGRXWLQIRUHVWVRIWKH3U]\PXV]HZR)RUHVW'LYLVLRQDQGFRQYHUVLRQIDFWRUV applied during inventories conducted in Poland according to the IPCC recommendations for international reporting submitted to the 8QLWHG1DWLRQV(FRQRPLF&RPPLVVLRQIRU(XURSH)$2DQG81)&&&
Ke or s afforestation, dendrometry, plantation, Padus serotinaSRVWDJULFXOWXUDOODQGVVSHFLHVGLYHUVLW\7XFKROD)RUHVW
1. ntro uction ment. One of the initiatives undertaken at the global level is the approach promoted in the documents adopted at the The process of forest reconstruction on former agricultural United Nations Conference on Environment and Develop- lands is one of the most important economic objectives of ment (“Earth Summit”) in Rio de Janeiro in 1992 calling forestry, as well as one of the most important and inter- for efforts towards “greening the world” and programmes esting problems of modern ecology. The growing interest developed under the IPPC (IPPC, 1996). The European in increasing the afforestation rate at the country, regional Union has implemented some legal regulations that cre- and global level results from the significant role of for- ate the basis for granting the financial assistance to those est ecosystems in mitigating the “greenhouse effect” and Member States who engage in efforts aimed at increas- increasing the ecological balance in the natural environ- ing the forest area through afforestation of uncultivated 52 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
and agriculturally inefficient lands. In Poland, a significant ricultural soils. In the circumstances when urgent measures increase in the afforestation rate after World War II was aimed at mitigating the greenhouse effect are required, the implemented within the scope of three projects: 1 – a res- most important issue is to assess the environmental effect toration plan for forest ecosystems after World War II dev- of afforestation of fallow lands, which is manifested in the astation, 2 – a national programme to increase the affores- increased plant biomass and amounts of organic carbon ac- tation rate, developed in the 1990s by the Minister of the cumulated in that biomass, observed at different levels of (QYLURQPHQWDO3URWHFWLRQ1DWXUDO5HVRXUFHVDQG)RUHVWU\ biosphere organization – from an individual and a popula- ±³WKH6WDWH)RUHVW3ROLF\´DGRSWHGE\WKH&RXQFLORI tion of trees to a landscape and region. Ministers in 1997. The objective of the latter initiative is Pomerania is one of the regions in Poland with the to increase the afforestation rate in the country up to 30% highest contribution of forests restored on former farm- in 2020 and 33% in the mid-21st century (Gorzelak, 1999). lands. In this region, there are many secondary forest ar- These objectives are to be implemented mainly through af- eas resulting from intensive afforestation carried out after forestation of unproductive agricultural lands. previous extensive deforestation. Deforestation of the re- In 1946–1994, ca. 1.200 thousand ha of previously cul- gion was carried out from the 16th century to the late 19th tivated or uncultivated lands were afforested across Po- century, since when intensive afforestation was undertaken ODQG :yMFLN 7KH SUHIHUUHG VSHFLHV DW WKDW WLPH by the Prussian authorities and continued by the Polish was Scots pine, less frequently silver birch, Norway spruce foresters after 1920. There are many forests on former and other species. The consequence of such actions are farmlands in the region of Pomerania, including mainly monospecies forest stands of the same age developed over WKH3U]\PXV]HZR)RUHVW'LYLVLRQZKLFKEHORQJVWR5'6) relatively large areas, which are exposed to many destruc- LQ 7RUXĔ 8QWLO WKH DUHD RI WRGD\¶V 3U]\PXV]HZR tive factors, i.e. annosum root rot (Heterobasion annosum )RUHVW'LYLVLRQEHORQJHGWRWKH3UXVVLDQ=ZDQVKRII)RU- )U %UHI JUDGDWLRQV RI LQVHFWV RU ILUH 7KHVH IDFWRUV est District, which was established in 1890 on the grounds cause major losses in the growth of trees and forest area of former Prussian estates as well as the Polish ones pur- and, consequently, losses in the production of wood. On chased or taken over by the Prussian government as part the other hand, despite the impact of unfavourable factors RIWKH³.XOWXUNDPSI´SROLF\DQGLQWHQGHGIRUDIIRUHVWDWLRQ and compared to the previously existing agroecosystems, )XUWKHULQWHQVLYHDIIRUHVWDWLRQWRRNSODFHDIWHU:RUOG:DU forests reconstructed on former arable lands are systems , ZKHQ WKH DUHD RI *GDĔVN 3RPHUDQLD ZDV LQFOXGHG LQ with a high carbon content accumulated in the living plant the Polish forest district established in this area. The next biomass and in the soil of the forest ecosystem. major afforestation of former agricultural lands was carried The importance of ecological effects of afforestation out after World War II when large estates were national- and problems encountered during the implementation of L]HGDQGLQFRUSRUDWHGLQWRWKH6WDWH)RUHVWVPDQDJHPHQW this process resulted in the increasing number of studies as part of the so-called agrarian reform. The last major af- that relate to the forest development on former agricul- IRUHVWDWLRQLQWKH3U]\PXV]HZR)RUHVW'LYLVLRQWRRNSODFH tural lands. The establishment success of plantations, their in the 1990s due to the lack of profitability of agriculture resistance to pests and adverse abiotic factors, the biomass in small areas within forests on poor sandy soils, and as growth of trees as well as the species diversity of forest a result of demographic processes, such as aging of farm- stands and the whole forest ecosystem are particularly thor- ers, migration of successors and users of agricultural lands oughly researched. to the cities. They relinquished their lease and the owner The issue of afforestation on former agricultural lands of the land, i.e. the state forest administration afforested LQ3RODQGKDVEHHQVWXGLHGLHE\%HUQDG]NLDQG.RZDO- the fallow lands. VNL %HUQDG]NL 1L]LĔVNL 5\NRZVNL Changes in the range of forests occurring in the Przy- 6REF]DN 6]XMHFNL 7XV]\ĔVNL PXV]HZR)RUHVW'LYLVLRQLQWKHSHULRGRI±ZHUH Gorzelak (1999). Research on former agricultural lands GHVFULEHGE\:LONRĔ0LFKDOVNDHWDO 1LHQDUWRZLF] was also carried out in other EU and non-EU countries, the HWDO DQG'HSWXáD 7KHODWWHURIWKH results of which have been published in a number of papers above authors assessed the changes in the plant biomass DQGERRNV HJ6DOELWDQR :DWNLQV -RKQVWRQ and the amount of carbon contained in that biomass on HWDO -RKDQVVRQ )OLQQ 9HOOHQG a map sheet at a scale of 1: 25000, covering an area of +HLOHWDO .LUNE\ :DWNLQV 0X\VHWDO about 120 km2. Nienartowicz et al. (1998, 2015) assessed 8ULHODO %DUGXOLVHWDO 6RPHRI the biomass changes resulting from afforestation of a land- the interesting aspects, studied include changes in the spe- ed estate covering 612 ha, where heaths used for sheep and cies composition, changes in the productivity of trees, and cattle grazing dominated before the afforestation. development of forests on former agricultural lands, which The objective of this work was to present the effects prompts to further, more profound analysis of the impact of afforestation on the accumulation of plant biomass and exerted on these processes by substances contained in ag- carbon contained in this biomass based on the example of Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 53
RQHIRUPHUO\DUDEOHSORWLQWKH3U]\PXV]HZR)RUHVW'LYL- established mainly by direct measurements carried out in sion afforested in 1991. In 2000, all trees still growing on WKH3U]\PXV]HZR)RUHVW'LYLVLRQDQGFRQYHUVLRQIDFWRUV this permanent study plot after nine years of cultivation used in inventories carried out in Poland according to the were measured (Dykiert, 2000). In 2015, measurements IPCC recommendations for international reporting submit- of all trees in the forest stand growing at the same plot ted to the United Nations Economic Commission for Eu- were repeated. This paper presents a comparison of den- URSH)$2DQG81)&&&" drometric features of the forest stand between 2000 and 2015 and described a range of changes occurring in its structure over the period of 15 years. Dendrometric pa- 2. The stu area rameters and the determined differences were examined in terms of ecological problems described in the literature as The research was conducted in subunit 277n of the Przy- those encountered in the process of forest reconstruction PXV]HZR)RUHVW'LYLVLRQZKLFKLVWKHQRUWKHUQPRVWHFR- on former arable lands. The main objective of our study QRPLFXQLWRI5'6)LQ7RUXĔ )LJ ,QWHUPVRIWKHWHU- was to answer the following three questions: 1) What is ritorial administration, the study area and the whole forest the standing crop after 25 years since afforestation of the district are located in the Pomerania province, the Choj- fallow land? 2) To what extent this biomass is greater nice county (poZiat) and the Brusy commune. Accord- compared to the field and the fallow land before affores- ing to the nature and forest regionalization proposed by tation? 3) What are the differences between assessments 7UDPSOHUHWDO WKH3U]\PXV]HZR)RUHVW'LYLVLRQ of the pine biomass with the use of conversion factors EHORQJV WR WKH :LHONRSROVND.XMDZ\ ,,, QDWXUDOIRUHVW
)LJXUH/RFDWLRQRI3U]\PXV]HZR)RUHVW'LYLVLRQLQ1RUWKHUQ3RODQG 54 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
UHJLRQ DQG WKH7XFKROD )RUHVW GLVWULFW ,Q WKH V\VWHP RI slope. A small number of Picea abies / +.DUVWBetula SURWHFWHGDUHDVWKHVWXG\DUHDLVORFDWHGLQWKH=DERUVNL pendula L., Quercus robur L. and Fagus sylvatica L. oc- Landscape Park, and in the Special Area of Conservation curred locally in the main tree stand. According to the State PLH 220026 Wielki Sandr Brdy and the Special Protection )RUHVW0DQDJHPHQW3ODQIRUWKH3U]\PXV]HZR)RUHVW'L- Area of Birds PLB 220009 Bory Tucholskie. The area is YLVLRQ 3ODQ8U]ąG]DQLD*RVSRGDUVWZD/HĞQHJR« DOVRDGMDFHQWWRWKHQDWXUHUHVHUYH.XODZD9DOOH\ Dolina the site index class of this tree stand was Ia,5 in 1999 and KulaZy) DQGWKH7XFKROD)RUHVW%LRVSKHUH5HVHUYH7KH the trees were then 46 years old, thus in 2015 (i.e. 16 years study area is located in the vicinity of the Laska forest later, at the time of our studies) they were 62 years old. The village on the slope with N-E exposure descending gently EDQNV RI WKH .XODZD ULYHU QHDU WKH VWXG\ DUHD DUH RYHU- WRZDUGV /DNH 6LHF]RQHN )LJ 7KH ODNH ZDV IRUPHG grown with alluvial alder forest, while the banks of the LQWKHSODFHZKHUHWKHVPDOO.XODZDULYHUIORZVLQWRWKH =EU]\FDULYHU±ZLWKEODFNDOGHUDQG1RUZD\PDSOHWUHHV =EU]\FD ULYHU ZKLFK LV D WULEXWDU\ RI WKH %UGD 5LYHU The study area is an elongated rectangle of 117 x 27 m LH RQH RI WKH PDLQ ULYHUV LQ WKH7XFKROD )RUHVW UHJLRQ (0.3159 ha). Until the mid-1980s, the area was used as &KRLĔVNL 7KHDUHDLVERUGHUHGE\DQDUURZVWULS SDUWRIWKHOHDVHE\DZRUNHURIWKH3U]\PXV]HZR)RUHVW of slightly younger pine woods adjacent to the lower-lying Division. Rye and potatoes were cultivated alternately in reed beds at the lake. WKHDUHD)RU\HDUVEHIRUHWKHDIIRUHVWDWLRQWKHDUHDZDV In the south-west and in the north, the study area is a fallow land. In 1990, the study area was afforested main- surrounded by pine forest growing in the upper part of the ly by Scots pine and silver birch. The pine was planted in
)LJXUH/RFDWLRQRIWKHVWXG\DUHD VXEXQLWQ LQ3U]\PXV]HZR)RUHVW'LYLVLRQ Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 55
the central part of the study area, and the birch – along DIWHU-DEáRĔVNL %XGQLDN 7KHYDOXHZDVIRU longer sides of the area, in five rows on each side, as well Scots pine and 0.52 grams dry matter/cm3 for silver birch. as at its northern edge. In one of the extreme rows of birch, )RUEODFNFKHUU\WKHYDOXHRIJUDPVGU\PDWWHUFP3 a few dozen beech trees were planted along a small section. ZDVDGRSWHG .R]DNLHZLF] 7KHELRPDVVRIWUHHV Based on the distribution maps of trees, it has been found with a breast height diameter of at least 7 cm was summed that Scots pine occurs over an area of 0.1994 ha, whereas up by species. silver birch over an area of 0.1165 ha. Apart from Scots Trees with breast height diameter less than 7.0 cm and pine and silver birch, the black cherry was noted in the up- juniper bushes were classified into established classes ac- per tree layer of the study area. The age of the main forest cording to their height. The frequency in each height class stand was 25 years. Scots pine, silver birch, grey alder, was calculated for each species. The frequency was mul- Norway spruce, common juniper, common rowan, black tiplied by the average aboveground dry biomass per tree cherry, European beech, Norway maple and pedunculate or shrub in a height class. The average biomass of one in- oak occur in the groundcover and in the undergrowth. dividual was obtained after cutting trees in the study area In the herb layer, Deschampsia flexuosa L., Festuca ovina or adjacent area. Trees were dried at 80°C for 48 hours. L., Corynephorus canescens L. and mosses – Pleurozium In the analysis, we also used the results of our previous schreberi (Willd.) Mitten., Dicranum undulatum Schrad., studies of the primary production and the biomass of forest Pohlia nutans (Hedw.) Lidb. are the dominant species. VWDQGVRIGLIIHUHQWDJHFODVVHVLQWKH3U]\PXV]HZR)RUHVW 'LYLVLRQ 'HSWXáD -DU]ĊEVNLHWDO 7KHXQ- derground biomass and the total dry biomass of all species 3. Materials an metho s of trees in this category were calculated using the conver- sion factors R/S defining the relationship between under- The location of each tree and shrub, including both in- ground and aboveground biomass of trees for the younger dividuals in the main stand and in the understorey as well age classes, obtained in the previous studies (Barcikowski as seedlings growing in the herb layer was determined over /RUR 'HSWXáD 7KHUHOHYDQWIUDFWLRQVRI the whole area using two measuring tapes. While mov- biomass of trees with a breast height diameter below 7.0 ing along the rows of trees, x and y coordinates of each of cm were added to the relevant values for trees with a larger them (identified to the species level), height and diameter breast height diameter. The results are presented for the at breast height were determined. The height of trees be- entire area and for 1 ha. The groundcover biomass value longing to the main stand (overstorey) was measured us- of 0.2384 kg/m2 (i.e. 2.384 t/ha, including 2.004 t/ha of ing the device Suunto PM-5/1520. The height of younger above-ground and 0.38 t/ha of under-ground biomass, re- specimens belonging to the lower forest layers was deter- spectively) in age class II (21–40 years) forests on former mined using a pole. These specimens were classified into arable lands was added to the values calculated for trees, height classes every 0.5 m. DBH was determined using a 40 determined in our previous studies conducted in the vicin- cm calliper. Tree diameter at breast height (including the ity of Laska village at the study sites representing the same thickness of a bark) was determined to the nearest 0.5 cm. category of habitat. After conducting field observations and measurements, the The biomass of the forest phase at the study site was composition of tree species occurring in the study area in also compared with the biomass of grain crop fields located 2015 was compared with that of 2000, described by Dykiert in the forest and on fallow lands near the village of Laska (2000). The species composition and the number of trees by assessed in our previous studies (Nienartowicz et al., 1998, species were compared and indicators of species diversity 'HSWXáD DQGHYHQQHVVZHUHFDOFXODWHGXVLQJ0963 .RYDFK Based on dendrometric measurements for each Scots pine, silver birch and black cherry with a breast height 4. Results diameter of at least 7 cm, the volume of the aerial parts ZDVGHWHUPLQHGXVLQJWDEOHVRI&]XUDM )RUWKHIHZ 4.1. Species composition an en rometric specimens of black cherry, for which no tables of thick- characteristics o the orest stan ness were available, abundance tables for birch were used DFFRUGLQJWRWKH)RUHVW0DQDJHPHQW,QVWUXFWLRQ SDUW A total of 905 trees belonging to 9 species and one spe- Instruction on how to prepare a forest management plan for cies of shrubs – Juniperus communis L., represented by 9 a forest division, Annex to Ordinance no. 43 of the Direc- specimens, were recorded in the study area. Compared to WRU*HQHUDORIWKH6WDWH)RUHVWVRI$SULO:DUVDZ 2000 when the total number of trees amounted to 2140, 2003). Next, the value of volume for each tree was multi- the number of trees decreased more than twice, while the plied by the specific gravity of wood (wood density) used number of species doubled from 5 to 10. The species di- LQWKH3RODQG¶V1DWLRQDO,QYHQWRU\5HSRUW .2%L=( versity index calculated by the Shannon formula increased 56 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
from 0.894 in 2000 to 1.235 in 2015, whereas the index of birch (Betula pendula) and black cherry (Padus serotina). species evenness slightly decreased (Table 1). The decline The number of the most abundant species, i.e. Scots pine, in the number of saplings was mainly due to the early decreased more than twice, from 1.178 in 2000 to 499 in- cutting and intraspecific and interspecific competition of dividuals in 2015. The maximum height of this species plants, game browsing and death of seedlings. Whereas increased in that period from 4.0 m to 21.0 m (Table 1). the increase in the number of tree and shrub species re- The maximum height of birch changed in a similar sulted from the import of their diaspores from the sur- range (from 4.0 to 22.0 m), but the number of specimens roundings. decreased almost five times (from 847 to 175). The con- The comparison of species composition indicates that tribution of the two additional species, i.e. common rowan three species dominated in both phases of the phytocoeno- (Sorbus aucuparia L.) and grey alder (Alnus incana (L.) sis development, i.e. Scots pine (Pinus sylvestris), silver Moench) significantly decreased in both compared phas-
Table 1. Comparison of the number of trees and their average height according to the state of the study area in 2000 and in 2015 Year 2000 2015 Maksimal umber Maksimal umber Species height o trees height o trees (m Scots pine - Pinus sylvestris L. 1178 4.0 499 21.0
Silver birch - Betula pendula Roth 847 4.0 175 22.0
American black cherry - Prunus serotina 82 2.0 161 11.0 (Ehrh.) Borkh.
European beech - Fagus sylvatica L. 62 1.5
Common juniper - Juniperus communis L. 9 1.5
Mountain ash - Sorbus aucuparia L. emend 10 0.5 3 6.0 Hedl.
Norway maple - Acer platanoides L. 2 0.5
Grey alder - Alnus incana (L.) Moench 23 0.5 1 3.0
Pedunculate oak - Quercus robur L. 1 0.5
Norway spruce - Picea abies / +.DUVW 1 0.5
Total number o trees 2140 914
Number of species 5 10
Species diversity H’ 0.894 1.235
Evenness e 0.556 0.537 Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 57
es (Table 1). New species in 2015 (as compared to 2000) of trees of this species were in the class of 7.0–10.4 cm were beech (Fagus sylvatica), Norway maple (Acer plata- (Table 2). noides L.), common oak (Quercus robur), Norway spruce The number of three species of trees with DBH above (Picea abies) and common juniper (Juniperus communis). 7.0 cm and their dendrometric characteristics are presented Fagus sylvatica – represented by a relatively large number in Table 3. The volume of aerial parts of 604 trees repre- of specimens (62) – was introduced in the process of af- senting the three species with a DBH of at least 7.0 cm forestation. The other four species (three tree species and was 91.75 m3, calculated with the use of Czuraj’s (1991) one shrub species) probably came from self-seeding. Three tables. The volume of 499 pine trees, 118 birch trees and species of trees – maple, oak and spruce were represented 3 black cherry trees was 80.57 m3, 11.03 m3 and 0.15 m3, by only one specimen. respectively. The volume of pine accounted for almost 88% Table 2 presents the abundance of 10 species deter- of the total volume (Table 3). mined in 2015 – 914 specimens per DBH classes. It ap- Only trees of the three most abundant tree species with pears that only three species of trees, i.e. Scots pine, silver DBH values below 7.0 cm reached great heights (Table 4). birch and black cherry reached DBH of 7.0 cm or more – The height of Scots pine and black cherry trees was 10.0 m. 604 trees in total. The remaining 310 trees had DBH below Most of the trees representing the latter species with small 7.0 cm. They belonged to all ten species. DBH values were included in the lowest height class. This In the group of the three most abundant species, 16 relation also applies to relatively common beech trees and pine trees, 57 silver birch trees and only three black cherry almost all species represented by a small number of indi- trees had DBH below 7.0 cm. The vast majority of indi- viduals. Rowan is an exception in this case, as two speci- viduals of the latter species occurred in the form of sev- mens of this species reach the height of 5.0 m and one – eral-year-old seedlings and their diameter at breast height 6.0 m, while their DBH does not exceed 4.0 cm (Table 4). was below 3.5 cm. All specimens of the common juniper and almost all trees of the six other species, except for 4.2. bo egroun plant biomass in 2015 one specimen of the common rowan, occurred in the low- est DBH class. The aboveground biomass of 483 Scots pine trees with Scots pine was characterised by the highest variability DBH above 7.0 cm was 37.868 t, the biomass of under- in DBH. Its specimens occurred in as many as ten de- ground parts of pine trees with the same DBH was 8.422 t, fined DBH classes. DBH of the thickest tree in the entire and the total aboveground and underground biomass was analysed population was in the class of 31.5–34.9 cm. 46.290 t (Table 5A). The aboveground biomass of silver Birch occurred in six DBH classes. The largest number birch with DBH of at least 7.0 cm was 5.736 t. The bio-
Table 2. The number of trees of 10 species by DBH classes in 2015
BH class Pinus Betula Padus Fagus Juniperus Sorbus Acer Alnus Quercus (cm sylvestris pendula serotina sylvatica communis aucuparia platanoides incana robur
0.1–3.4 6 24 152 62 9 2 2 1 1 3.5–6.9 10 33 6 1 7.0–10.4 112 52 2 10.5–13.9 160 29 14.0–17.4 126 30 1 17.5–20.9 51 7 21.0–24.4 19 24.5–27.9 4 28.0–31.4 10 31.5–34.9 1 Total 499 175 161 62 9 3 2 1 1 58 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
7DEOH'HQGURPHWULFFKDUDFWHULVWLFVDQGWKHVWDQGYROXPHRIWKHDERYHJURXQGSDUWVRIWUHHVZLWK'%+FPE\VSHFLHV
erage erage umber Stan olume Species BH height o trees (cm (m (m3 ( Pinus sylvestris 483 13.75 13.18 80.57 87.82 Betula pendula 118 11.54 12.27 11.03 12.02 Prunus serotina 3 10.67 7.33 0.15 0.16 Total 604 91.75 100.00
Table 4. The number of trees of 10 species with DBH < 7 cm in subsequent classes of height and the maximum DBH in individual species
Total Height class Pinus Betula Padus Fagus Juniperus Sorbus Acer Quercus Picea Alnus number (m sylvestris pendula serotina sylvatica communis aucuparia platanoides robur abies incana o trees 0.01 – 0.50 4 96 51 7 2 1 1 162 0.51 – 1.00 1 1 11 10 1 24 1.01 – 1.50 3 39 1 1 44 1.51 – 2.00 6 4 10 2.01 – 2.50 1 1 2 2.51 – 3.00 1 8 1 10 3.51 – 4.00 7 1 8 4.51 – 5.00 2 9 3 2 16 5.51 – 6.00 11 1 1 13 6.51 – 7.00 3 6 1 10 7.51 – 8.00 4 4 8 8.51 – 9.00 1 1 9.51 – 10.00 1 1 2 Total 16 57 158 62 9 3 2 1 1 1 310 Max. DBH 6.5 6.0 6.5 1.0 1.0 4.0 0.5 0.5 1.0 1.5 (cm)
mass of underground parts was assessed at 1.451 t and the 7.0 cm was 53.566 t, while the contribution of individual total aboveground and underground biomass of birch with components to the total biomass was as follows: 86.417% the same DBH – at 7.187 t. In the case of three black cherry for pine, 13.417% for birch and 0.166% for black cherry. specimens, the biomass of aboveground and underground parts as well as of the whole trees amounted to 0.071 t, The total aboveground biomass of all 310 trees with 0.018 t and 0.089 t, respectively (Table 5A). DBH below 7.0 cm was 0.316 t (Table 5B). The biomass of The total aboveground and underground biomass of the underground parts was 0.074 t, which amounted to 0.390 three species represented by trees with DBH of at least t for the entire study area. The biomass of these trees con- [59] Total .965 53.956 n er- groun Biomass (t 0.633 0.120 0.753 44.624 10.085 54.709 bo e- groun trees um- ber ber o he two DBH categories and the herb-layer biomass Biomass (t trees um- ber ber o and the total dry biomass of trees by species and the size of t Total bo egroun n ergroun Total n er- groun Biomass (t bo e- groun . Trees ith . Trees BH 7 cm ith B. Trees BH 7 cm ll C. trees o trees umber umber 9 0.0024 0.00046 0.0029 9 0.0024 0.00046 0.0029 3 0.012 0.00402 0.016 3 0.012 0.00402 0.016 2 0.0000022 0.0000012 0.000003 2 0.000002 0.0000012 0.0000034 3 0.071 0.018 0.089 158 0.05006 0.014 0.064 161 0.121 0.032 0.153 62 0.00035 0.00021 0.00056 62 0.00035 0.00021 0.00056 483 37.868 8.422 46.290 16 0.047 0.012 0.059 499 37.915 8.434 46.349 118 5.736 1.451 7.187 57 0.203 0.043 0.246 175 5.939 1.494 7.433 1 0.0000053 0.0000021 0.0000074 1 0.0000053 0.0000021 0.0000074 1 0.00103 0.00034 0.0014 1 0.00103 0.00034 0.0014 1 0.000018 0.0000092 0.000027 1 0.000018 0.0000092 0.000027 as well as the total biomass of vegetation in the study area Species Herbs, mosses and lichens biomass biomass Total (trees, shrubs, herbs, mosses and lichens) Total treestan Total 604 43.675 9.891 53.566 310 0.316 0.074 0.390 914 43.991 9 Picea abies Quercus robur Quercus Acer platanoides Alnus incana Sorbus aucuparia Fagus sylvatica Juniperus communis Prunus serotina Betula pendula Pinus sylvestris Table 5. Table The number of trees and the aboveground, underground, 60 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
sisted mainly of silver birch – 63.120%, wild black cher- cherry was the second most abundant species after silver ry – 16.376% and Scots pine – 15.142%. Common rowan birch in this group (category) of trees. Our study deter- contributed in only 4.189%, while each of the remaining mined that the contribution of trees with DBH below 7 cm species – no more than 1%, in total – 1.173%. Contribu- in the total aboveground biomass of the whole tree stand tion of all trees with DBH below 7 cm in aboveground was 0.72%. A slightly higher percentage contribution of biomass of the tree stand was only 0.72%. The total the understory and undergrowth in aboveground biomass aboveground biomass of all 914 trees growing at the study ZDVUHSRUWHGE\2U]HáHWDO E LQWKHRDNWUHHVWDQGV site – i.e. those with a diameter at breast height below 7.0 LQWKH1LHSRáRPLFH)RUHVW,QWKHDOGHUIRUHVWVWDQGRIWKH cm and those with a larger DBH – was 43.991 t (Table 5C). )RUHVW WKLV FRQWULEXWLRQ ZDV VOLJKWO\ DERYH 2U]Há The biomass of underground parts was assessed at 9.965 t. et al., 2005). The lower contribution (estimated by us at The total biomass of trees, i.e. aerial and underground parts only 0.72%) could result from the fact that the pine-birch was 53.956 t (Table 5C). Pine contributed to the total bio- IRUHVWVWDQGLQWKH7XFKROD)RUHVWZDV\RXQJHUWKDQWKHRQH mass of all trees in 85.901%, birch in only 13.776%, and LQWKH1LHSRáRPLFH)RUHVWDQGWKH\RXQJHUGHYHORSPHQW black cherry – in 0.283%, seven other species – 0.038% in phase is characterised by stronger competition of the main total. tree stand in relation to the understory and undergrowth. After conversion to 1 ha unit area, the aboveground Czuraj’s (1991) tables were used in the assessment of biomass of the forest stand was 139.256 t/ha, the under- aboveground biomass of trees with DBH above 7.0 cm. ground biomass – 31.545 t/ha and aboveground and under- The tables provide information on the overall volume ground biomass in total – 170.801 t/ha. of standing trees (in m3) as a total value for large tim- When taking into account the aboveground and under- ber, i.e. the fraction subject to inventories and parts of the ground biomass of the groundcover in the total biomass of WUXQNZLWKDGLDPHWHUEHORZFP9DOXHVUHDGIURPWKH the forest stand, i.e. 2.004 t/ha and 0.38 t/ha respectively tables of Czuraj (1991) were converted to dry matter using (0.633 t and 0.12 t for the whole study area of 0.3159 ha), specific wood density of a given tree species. However, in we arrived at the value of 54.709 t for the whole study area LQYHQWRULHVFRQGXFWHGE\WKH6WDWH)RUHVWVDQGUHSRUWLQJWR (Table 5C). Trees with DBH of at least 7.0 cm, trees with WKH81(FRQRPLF&RPPLVVLRQIRU(XURSH)$2IRU DBH below 7.0 cm and groundcover contributed to this bio- 7HPSHUDWHDQG%RUHDO)RUHVW5HVRXUFHV$VVHVVPHQW±7%- mass in 97.911%, 0.713% and 1.376%, respectively. After )5$ WKH%LRPDVV([SDQVLRQ)DFWRU %() LVXVHG conversion to 1 ha unit area, the total plant biomass (com- i.e. the ratio of the total aboveground biomass of trees to prising trees, herbs, mosses and lichens) was 173.184 t/ha. the biomass of large (merchantable) timber according to Szymkiewicz’s tables (2001). $FFRUGLQJ WR -DEáRĔVNL DQG %XGQLDN WKH UD- 5. iscussion an conclusions tio for coniferous forest stands in age class II is 1.67 and for over 140-year-old forest stands – 1.10, and the gener- Directions of changes in the phytocoenosis structure ob- alized index for coniferous species in total is 1.265 and served in the study area are similar to those observed in for deciduous species – 1.170. In ecology, this index was other forest communities developed on former agricultural also described as a Total/Merchantable timber (T/M) ra- lands. They include an increase in the number of tree spe- WLR -RKQVRQ 6KDUSH 0XULOOR5RGULTXHV cies due to natural spread of diaspores from neighbouring 1997). It is calculated as a ratio of the total aboveground phytocoenoses. A typical phenomenon is also a decrease biomass (including the trunk, branches and leaves) to the in the density of planted trees, both due to the competition inventoried biomass of merchantable timber. Based on the and the effect of parasites, especially annosum root rot. direct measurements conducted in our previous studies, we Such interactions are manifested in fallen trees, consider- calculated that the T/M ratio for Scots pine in age class II able gaps in the forest canopy and relatively rapid change ZDVDQGIRU6FRWVSLQHLQFODVV9± in the type of spatial distribution of trees from regular to 'HSWXáD :KHQXVLQJWKH70LQGH[HTXDOWR rather random distribution. and wood density of 0.430 t/m3, we found that the above- The observations also proved the invasive nature of Pa- ground biomass of pine trees with DBH of at least 7.0 cm dus serotina, which is reflected in the presence of many amounts at the study site to 30.7202 t. When using the seedlings. Perhaps they germinated from the seeds of the %() LQGH[ DQG ZRRG GHQVLW\ RI WP3, the three older living trees. These trees spread at the study site aboveground biomass of large pine timber at the study site most likely before afforestation, i.e. at the time when the amounts to 36.630 t. When using Czuraj’s tables (1991), land was left fallow. They were left in the process of affor- the value was 37.868 t. estation to increase the species diversity of the forest stand. The difference between these two last values is 1.238 t, Padus serotina was the most common species among which accounts for 3.27% of the aboveground biomass de- trees with DBH below 7 cm. In terms of biomass, black termined at the study site using Czuraj’s (1991) tables. This Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 61
value is similar to the percentage contribution of needles in HIIHFW ZDV DOVR FRQILUPHG E\ +HOLĔVND5DF]NRZVND DQG the aboveground biomass of pine tree stands quoted in the )DELVLDN ZKRUHSRUWHGWKDWWUHHVFKDUDFWHULVHGE\ OLWHUDWXUHZKLFKLVXVXDOO\QRWLQFOXGHGLQWKH%()LQGH[ rapid growth (the so-called “wolf trees”) are characterised )RUH[DPSOH2U]HáHWDO D UHSRUWWKDWWKHQHHGOHV E\ORZDYHUDJHGHQVLW\)RU\HDUROGSLQHWUHHV LHRI account for 3.2% of the total aboveground biomass of pine a similar age as pines growing in our study area), the au- WUHHVWDQGVLQWKH1LHSRáRPLFH)RUHVW2XUDQDO\VLVVKRZV thors determined that the default wood density is in the that the ratio of the total biomass 37.868 t/m3 and the bio- range of 0.321–0.360 t/m3. mass of large (merchantable) timber 21.934 t/m3 of the On the other hand, other authors found (though based main tree stand, calculated on the basis of Czuraj’s (1991) on less extensive material) that the default value for wood tables, was 1.726. The higher value, above 1.67, may result density of pine on former agricultural lands is higher com- from the contribution of leaves in the total aboveground SDUHG WR IRUHVWHG ODQGV )RU H[DPSOH EDVHG RQ WKH UH- biomass of trees. VHDUFK FRQGXFWHG LQ WKH 0LDVWNR )RUHVW 'LYLVLRQ 5'6) When using Szymkiewicz’s (2001) tables for pine Szczecinek), thus near the areas managed by the Przy- with the site index class Ia based on average DBH and PXV]HZR)RUHVW'LYLVLRQ-HORQHNHWDO IRXQGWKDW the height of trees, as well as using the biomass extension wood of trees growing on the habitat of fresh coniferous factor of 1.67 and the wood density index of 0.43 t/m3, the forest (on ancient forest soils) was characterised by aver- aboveground biomass of the pine forest stand growing over age default density of 0.4257 t/m3, while trees growing an area of 0.1994 ha was only 29.961 t, which is 7.907 t on former agricultural lands – 0.47046 t/m3. On a slightly less than the value obtained based on Czuraj’s (1991) ta- more fertile habitat of fresh mixed coniferous forest, the EOHV:KHQXVLQJWKH%()LQGH[RIWKHGLIIHUHQFH wood density was at the level of 0.43413 t/m3 on ancient was 6.902 t. forest soils and 0.47407 t/m3 on a former agricultural land. The biomass assessment accuracy was also affected by )XUWKHUPRUH WKH DERYH DXWKRUV IRXQG WKDW ZRRG GHQVLW\ adopted values of weight of one individual in the height depends on the biosocial class of trees. In the case of classes defined every 0.5 m within the range from 0 to trees growing on former arable lands, wood of predomi- 10.0 m. Direct measurements were used only for the low- QDQW WUHHV .UDIW FODVV , LV FKDUDFWHULVHG E\ WKH KLJKHVW est classes of the most abundant species, i.e. pine, birch density – 0.50561 t/m3, while the wood of codominant DQGEODFNFKHUU\)RUKLJKHUDQGWKLFNHUVSHFLPHQVRISLQH WUHHV .UDIWFODVV,,, ±E\WKHORZHVWGHQVLW\RI and birch, but with DBH below 7.0 cm, values determined t/m37KHZRRGRIGRPLQDQWWUHHV .UDIWFODVV,, KDVDQ for individuals growing at the adjacent sites in 2015 were average density of 0.45135 t/m3. In the case of trees grow- used. This principle was also applied in the case of rare ing on typical forested lands (no agricultural use in the species (beech, common juniper), usually occurring in low- past), codominant trees (class III) are characterised by the er height classes, to avoid their elimination or reduction of highest wood density – 0.44994 t/m3 and the dominant WKHLUFRQWULEXWLRQDWWKHVWXG\VLWH)RUVRPHVSHFLHVVXFK trees (class II) – by the lowest density of 0.42073 t/m3. as beech, spruce, grey alder, literature data were also used. The wood of predominant trees (class I) has a medium )DFWRUV FRQYHUWLQJ WKH ELRPDVV WR ZRRG GHQVLW\ DOVR density of 0.42671 t/m3. DIIHFWHGWKHTXDOLW\RIELRPDVVDVVHVVPHQW)RUWKHGRPL- According to Jelonek et al. (2010), basic density of nant species, i.e. pine and birch, conversion factors of wood on the habitat of fresh coniferous forest on typi- 0.430 and 0.520 t dry matter /m3 were used (respectively) cal forest soils is 0.43026 t/m3, whereas on former arable LQWKH3RODQG¶V1DWLRQDO,QYHQWRU\5HSRUW .2%L=(DIWHU soils – 0.47231 t/m3, with the average general value of -DEáRĔVNL %XGQLDN 7KHVHYDOXHVDUHPXFKORZHU 0.45070 t/m3. WKDQWKRVHSURYLGHGE\.U]\VLN WGU\PDWWHU Witkowska and Lachowicz (2012, 2013) determined m3 for pine and 0.610 t dry matter/m3 for birch. The value that pine in age class II, growing on the habitat of fresh adopted for pine, i.e. 0.430 t/m3, is also lower than the PL[HGFRQLIHURXVIRUHVWVLQWKH7XFKROD)RUHVWLVFKDUDF- value of 0.450 t/m3 proposed for pine by Trendelenburg terised by the default wood density of 0.419 t/m3 at DBH, and Mayer-Wegelin (1955). It is, however, similar to the 0.382 t/m3 halfway up the trunk, and 0.375 t/m3 in the up- value of 0.435 kg/m3 determined by Tomczak and Jelonek per end of merchantable timber. Based on the analysis of (2013) based on the analysis of a large material collected 400 samples of trees from four regions of Poland, grow- in e.g. forest divisions of Trzebielino, Drawsko Pomorskie LQJRQIRXUW\SHVRIKDELWDWV '&)±GU\FRQLIHURXVIRUHVW DQG:DUFLQR 5'6)6]F]HFLQHN LQWKH3RPHUDQLDUHJLRQ )&)±IUHVKFRQLIHURXVIRUHVW)0&)±IUHVKPL[HGFRQLI- DQGLQWKH2OHĞQR)RUHVW'LYLVLRQ 5'6).DWRZLFH )RU HURXVIRUHVW)0)±IUHVKPL[HGIRUHVW DQGUHSUHVHQWLQJ pines growing in forested areas, the above authors reported IRXUDJHFODVVHV IURP,,WR,9 DQGWKUHHGLIIHUHQWKHLJKWV a higher value of 0.478 t/m3. The lower value for pines of the trunk, the above authors arrived at an average value growing on former agricultural lands is usually explained of 0.417 t/m3. The default value of wood density for the by their rapid increments (Tomczak et al., 2009). This 7XFKROD)RUHVWUHJLRQLQWKHVDPHUDQJHRIKDELWDWW\SHV 62 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
age classes and trunk heights, was 0.420 t/m3, which is value of 0.24. The value of 0.2224 is thus an intermediate similar to the value of 0.430 t/m3 used in our calculations. value in the range of 0.1442–0.2831. Biomass density of Scots pine wood also depends on Based on the data provided by Cannell (1982), the R/S its maturity and varies along the position of the cross-sec- value for a 42-year-old Betula verrucosa stand in the Mos- tion on the longitudinal stem profile. Based on the analy- cow Province is 0.207 and for a 35-year-old forest stand sis carried out in six pine forest stands on the habitat of of B. verrucosa and Populus sp. in the region of Novosi- fresh coniferous forest and fresh mixed coniferous forests, ELUVN± )RUDVLOYHUELUFKIRUHVWVWDQGLQWKH8. Tomczak and Jelonek (2012) determined that the default the above values of underground and aboveground biomass density of young and mature wood decreases with increas- yielded the R/S value of 0.272.)RUELUFKWUHHVZLWK'%+ ing distance from the base of the trunk. At the breast height of 7 cm and more, the value of 0.253 was used, and for de- and in the middle of the section between DBH and the ciduous trees with DBH below 7 cm – 0.25. Bardulis et al. base of the live crown, the density of juvenile wood was (2015) reported that for young grey alder stands under 10 lower compared to mature wood. At the base of the live years of age growing on abandoned agricultural lands in crown, juvenile wood was characterised by a higher den- Central Latvian lowlands, the average ratio of the above- sity, but the differences were not statistically significant. ground biomass to roots is 3:1. Similar data were provided )RUDOOVDPSOHVFROOHFWHGDWWKUHHKHLJKWVRIWUHHVJURZLQJ by Uri et al. (2002, 2007) for grey alder and silver birch at six sites in two types of forest habitat, the above au- in Estonia. thors arrived at a density of 0.396 t/m3 for juvenile wood The values obtained for the aboveground biomass and 0.423 t/m3 for mature wood. Taking into account the (139.256 t/ha) and for the underground biomass (31.545 t/ effect of spacing and arrangement of seedlings in planta- ha) of the studied forest stand are similar to those quoted tions of Scots pine harvested for construction timber as in the literature for similar ecological systems. According GHILQHGE\6SáDZD1H\PDQHWDO RQHFDQDVVXPH WR%LMDNDQG=DVDGD WKHDERYHJURXQGELRPDVVIRU that artificially regenerated forest stands (i.e. developed a 23-year-old pine tree stand on the habitat of fresh mixed from planting), usually having a greater density and more FRQLIHURXV IRUHVW LQ WKH /XEVNR )RUHVW 'LYLVLRQ 5'6) regular distribution of trees, are characterised by a higher =LHORQD *yUD LV WKD ZKLOH IRU D \HDUROG proportion of mature wood, which results in higher wood pine wood in the habitat of fresh coniferous forest in the GHQVLW\ 7RPF]DN -HORQHN -DEáRĔVNL %XGQLDN same forest division – 169.770 t/ha. The average biomass )RUWKLVUHDVRQWKHYDOXHRIWP3, similar to RI URRWV LQ /XEXV] 3LQH )RUHVWV FDOFXODWHG EDVHG RQ WKH the conversion factor of 0.430 t/m3 used in our studies, ap- 56UDWLRZDVWKD %LMDN =DVDGD 9DOXHV pears to be more relevant to our study area. for the biomass of roots in young and middle-aged forest The adopted value of the R/S index, i.e. 0.2224, also stands were similar: 26.577 and 27.201 t/ha, respective- affected the biomass assessment. This value indicates that ly. They were much lower compared to the oldest forest underground parts account for 18.195% and aboveground stands – 36.408 t/ha. In terms of habitat, values of the un- parts – for 81.805% of the total biomass of the forest stand. derground biomass of trees increased with increasing mois- The value of 18.195% for pine stands is therefore higher ture content and habitat fertility: 33.626 t/ha on the habitat than 14% quoted by Laurow (1966). of fresh coniferous forest and 34.111 t/ha on the habitat of The adopted value is lower than 0.26 obtained by Xiao fresh mixed coniferous forests. and Ceulemans (2004) for 10-year-old pine trees, with un- The biomass value stated for our study site is also simi- derground parts accounting for 20.557% and aerial parts – lar to the value obtained by Cannell (1982) for a 33-year- for 79.443%. Xiao et al. (2003) suggested R/S = 0.1442 ROGSLQHIRUHVWVWDQGLQWKH8.$FFRUGLQJWRWKLVVRXUFH for 73-year-old pine, which means that the aboveground the value of aboveground biomass was 140.1 t/ha and of part accounted for 87.4% and the underground part only the underground biomass – 36.1 t/ha, which gives a total 12.6% of the total biomass. Oleksyn et al. (1999) reported value of 176.2 t/ha. Whereas Johansson (1999) reported R/S = 0.2831 for 12-year-old pine (underground 22.07% that aboveground biomass of a 26-year-old silver birch and aboveground 77.93% of the total biomass of a tree). stand growing on fallow lands in Sweden at a latitude of &DQQHOO LQ KLV ERRN ³:RUOG )RUHVW %LRPDVV DQG 60°09’ was 175.3 t/ha. Primary Production Data” provided R/S = 0.1985 for Assuming that carbon accounts on average for 50% of a 22-year-old Pinus nigra Arn. stand in the Netherlands GU\SODQWPDWWHU :KLWWDNHU /LNHQV $MWD\HWDO and 0.2577 for a 33-year-old Pinus sylvestris forest stand. 'HZDU &DQQHOO +ROOLQJHU HW DO The former value is similar to R/S equal to 0.2, proposed +RXJKWRQ .DUMDODLQHQ *RZHUHWDO by the committee on climate change (IPCC, 1996) for co- -DJRG]LĔVNLHWDO :\VRFND)LMRUHN =DMąF niferous forests in the temperate zone. Cairns et al. (1997) plant biomass at the study site contained ca. 86.592 t suggest the value of 0.26, thus the same as proposed by C /ha. When assessing the environmental effects of affor- ;LDRDQG&HXOHPDQV .U|QHU VXJJHVWVWKH estation on former arable lands implemented in terms of Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 63
greenhouse effect mitigation, the current biomass of a for- [Assessment of the belowground biomass in Scots pine est community can be compared to the plant biomass of stands of Bory Lubuskie]. Sylwan 151(12): 21–29. a field existing and cultivated before the afforestation. If &DQQHOO0*5 :RUOG)RUHVW%LRPDVVDQG3ULPDU\ the maximum plant biomass before the afforestation of fal- Production Data. Academic Press, A subsidiary of Har- low lands was 4.566 t/ha (as noted in our previous studies FRXUW%UDFH-RYDQRYLFK3XEOLVKHUV/RQGRQ1HZ
URSH )URP ILHOG REVHUYDWLRQ WR 'HFLVLRQ 6XSSRUW -RKQVRQ :& 6KDUSH '0 7KH UDWLR RI WRWDO Springer, Dordrecht, The Netherlands. merchantable forest biomass and its applicationto the +HOLĔVND5DF]NRZVND/ )DELVLDN( =PLHQQRĞü JOREDOFDUERQEXGJHW&DQ-)RU5HV ± JĊVWRĞFLGUHZQDVRVQ\]GU]HZQDOHĪąF\FKGRJUXS\ -RKQVWRQ0++RPDQQ36(QJVWURP-.*ULJDO') W]Z UR]SLHUDF]\ >9DULDWLRQRI WKH 6FRWV SLQH ZRRG 1996. Changes in ecosystem carbon storageover 40 density index among so-called wolf trees]. Prace years on an old-field / forest landscape in east-central .RPLVML7HFKQRORJLL'UHZQD>6WXGLHVRI:RRG7HFK- 0LQQHVRWD)RUHVW(FRORJ\DQG0DQDJHPHQW ± nology Committee] 13: 41–49. .DUMDODLQHQ7 0RGHO FRPSXWDWLRQV RQ VHTXHVWUD- +ROOLQJHU'<0DFODUHQ-3%HHWV31 7XUODQG- tion of carbon in managed forests and wood products &DUERQVHTXHVWUDWLRQLQ1HZ=HDODQG¶VSODQ- XQGHUFKDQJLQJFOLPDWLFFRQGLWLRQVLQ)LQODQG-RXUQDO WDWLRQIRUHVWV1HZ=HDODQG-RXUQDORI)RUHVWU\ of Environmental Management 47: 311–328. 194–208. .LUNE\.- :DWNLQV&K7KH(FRORJLFDO+LVWRU\ Houghton R.A., 1996, Converting terrestrial ecosystems RI(XURSHDQ)RUHVWV&$%,3XEOLVKLQJ&3,$QWKRQ\ from sources to sinks of carbon. Ambio 25: 267–272. Rowe, Eastbourne. IPPC, 1996, IPCC Guidelines for National Greenhouse .RYDFK:/ 0963±$0XOWL9DULDWH6WDWLVWLFDO Gas Inventories: Reference Manual. 3DFNDJHIRU,%03&¶VYHUVLRQ.RYDFK&RPSXWLQJ -DEáRĔVNL 0 %XGQLDN 3 6]DFRZDQLH QD- 6HUYLFHV3HQWUDHWK:DOHV8. G]LHPQHM ELRPDV\ GU]HZQHM ODVyZ Z 3ROVFH QD .R]DNLHZLF]3&]HUHPFKDDPHU\NDĔVND Prunus SRWU]HE\ VSUDZR]GDZF]RĞFL (.*)$2 L 81)&&& serotina(KUK ±GU]HZR]$PHU\NL3yáQRFQHM>%ODFN [Estimating above-ground woody biomass of forests cherry (Prunus serotina Ehrh.) – a tree from North LQ 3RODQG IRU 81(&()$2 DQG 81)&&& UHSRUW- $PHULFD@ 3U]HP\Vá 'U]HZQ\ >:RRG ,QGXVWU\@ /;, LQJ@/HĞQH3UDFH%DGDZF]H )RUHVW5HVHDUFK3DSHUV ±:\GDZQLFWZRĝZLDW 75(3): 277–289. .U|QHU& %LRPDVVIUDFWLRQDWLRQLQSODQWV DUHFRQ- -DJRG]LĔVNL $ 0 -DURVLHZLF] * .DUROHZVNL 3 sideration of definitions based on plant functions, [in:] 2OHNV\Q-=DZDUWRĞüZĊJODZELRPDVLHSR- J. Roy, E. Garnier (eds), A whole plant perspective on VSROLW\FKJDWXQNyZNU]HZyZSRGV]\FLDOHĞQHJR>&DU- carbon-nitrogen interactions. SPB Academic, Haga: bon concentration in the biomass of common species 173–185. of understory shrubs]. Sylwan 156(9): 650–662. .U]\VLN) 1DXNDRGUHZQLH>:RRGVFLHQFH@3:1 -DU]ĊEVNL01LHQDUWRZLF]$'HSWXáD0%XEQLFNL- Warszawa. 'RPLQ'-3DVWFXUUHQWDQGSRWHQWLDOUH- /DXURZ = .RPSOHNVRZH XĪ\WNRZDQLH DUERPDV\ sources of carbon and above-ground plant biomass in OHĞQHM>([WHQVLYHXVHRIIRUHVWWUHHELRPDVV@:\GDZ- WKHODQGVFDSHZLWKKHDWVLQ7XFKROD)RUHVW(FRORJL- QLFWZRĝZLDW:DUV]DZD cal Questions 13: 9–27. Murillo-Rodriquez J.C., 1994, The carbon budget of the -HORQHN 7 3D]GURZVNL : 7RPF]DN $ Spanish forests. Biogeochemistry 24: 1–21. :áDĞFLZRĞFL GUHZQD VRVQ\ ]Z\F]DMQHM Pinus syl- Murillo-Rodriquez J. C., 1997, Temporal variations in the vestris/ QDJUXQWDFKSRUROQ\FKZSyáQRFQHM3ROVFH carbon budget of forest ecosystems in Spain. Ecologi- [Selected properties of wood in Scots pine (Pinus syl- cal Applications 7 (2): 461–469. vestris L.) growing on post-agricultural land in north- 0X\V%/XVW1 *UDQYDO3 (IIHFWVRIJUDVVODQG HUQ3RODQG@/HĞQH3UDFH%DGDZF]H )RUHVW5HVHDUFK afforestation with different tree species on earthworm Papers) 70(3): 277–286. communities, litter decomposition and nutrient status. Jelonek T., Pazdrowski W., Arasimowicz-Jelonek M. Soil Biology and Biochemistry 24: 1459–1466. 7RPF]DN$:áDĞFLZRĞFLGUHZQDVRVQ\]Z\ 1LHQDUWRZLF]$)LOEUDQGW&]DMD$'HSWXáD0 %RLĔVNL czajnej (Pinus sylvestris/ SRFKRG]ąFHM]JUXQWyZ M., 1998, Changes in land use and their effect on spe- porolnych [Properties of Wood of Scots pine (Pinus cies diversity and above-ground standing crop in the sylvestris L.) growing on former farmlands]. Sylwan surroundings of lake Milachowo, [in:] S. Borsuk (ed.), 154(5): 299–311. Research of the lakes in national parks and other pro- Johansson T., 1999, Biomass equations for determining tected areas, II Int. Scientific Conference. Inst. Wyd. fractions of pendula and pubescent birches growing Habitat, Bydgoszcz: 20–31. on abandoned farmland and some practical implica- 1LHQDUWRZLF]$.XQ]0'HSWXáD0 'RPLQ' tions. Biomass and Bioenergy 16: 223–238. Ecological consequences of changes in landscape struc- Johansson T., 2007, Biomass production and allometric ture in the neighbourhood of Brusy in 19th and 20th cen- above- and below-ground relations for young birch tury. Ecological Questions 1: 117–135. stands planted at four spacings on abandoned farm- Nienartowicz A., Lewandowska-Czarnecka A., Ortega ODQG)RUHVWU\ ± ('HSWXOD0)LOEUDQGW&]DMD$ .RZQDFND0 Biomass of Scots pine-silver birch tree stand 25 years after afforestation of former agricultural land 65
2015, Afforestation of heathlands and its influence on 6]\PNLHZLF] % 7DEOLFH ]DVREQRĞFL L SU]\URVWX the land cover, accumulation of plant biomass and en- GU]HZRVWDQyZ>:RRG\LHOGDQGYROXPHLQFUHPHQWWD- HUJ\IORZLQWKHODQGVFDSH $QH[DPSOHIURP=DERUVNL bles]. PWRiL, Warszawa. Landscape Park. Ecological Questions 21: 91–99. 7RPF]DN $ -HORQHN 7 3DUDPHWU\ WHFKQLF]QH 1L]LĔVNL = =DJRVSRGDURZDQLH L SURGXNF\MQRĞü PáRGRFLDQHJR L GRMU]DáHJR GUHZQD VRVQ\ ]Z\F]DMQHM GU]HZRVWDQyZ QD JUXQWDFK SRUROQ\FK QD SU]\NáDG]LH (Pinus sylvestris L.) [Technical parameters of juvenile 2=/3Z6]F]HFLQNX>0DQDJHPHQWDQGSURGXFWLYLW\RI and mature wood in Scots pine (Pinus sylvestris L.). stands on post-agricultural grounds exemplified by Re- Sylwan 156(9): 695–702. JLRQDO$GPLQLVWUDWLRQRI6WDWH)RUHVWLQ6]F]HFLQHN@ 7RPF]DN $ 3D]GURZVNL : -HORQHN 7 6\OZDQ&;;;,9 ± ± Wybrane elementy budowy makrostrukturalnej drew- 2OHNV\Q - 5HLFK 3%&KDOXSND : 7MRHONHQ 0* QDDGRMU]DáRĞüVRVQ\]Z\F]DMQHM Pinus sylvestris L.) 1999, Differential above- and below-ground biomass Z\URVáHMZZDUXQNDFKJUXQWyZSRUROQ\FK>&RUUHODWLRQ accumulation of European Pinus sylvestris populations between selected elements of wood macrostructure and in 12-yr-old provenance experiments. Scandinavian maturity of Scots pine (Pinus sylvestris L.) growing on -RXUQDORI)RUHVW5HVHDUFK ± SRVWDJULFXOWXUDOODQG@/HĞQH3UDFH%DGDZF]H )RUHVW 2U]Há6)RUJLHO06RFKD- 2FKDá:%LRPDVV Research Papers) 70(4): 373–381. and Annual Production of Common Alder Stands of the 7RPF]DN$ -HORQHN73URPLHQLRZD]PLHQQRĞü 1LHSRáRPLFH)RUHVW(-3$8 ZZZHMSDXPHGLDSO ZáDĞFLZRĞFLGUHZQDVRVQ\]Z\F]DMQHM Pinus sylvestris volume8/issue1/forestry/art-25.html. / Z\URVáHMQDJUXQWDFKSRUROQ\FK>5DGLDOYDULDWLRQLQ 2U]Há 6 )RUJLHO 0 2FKDá : 6RFKD - D the wood properties of Scots pine (Pinus sylvestris L.) 1DG]LHPQDELRPDVDLURF]QDSURGXNFMDGU]HZRVWDQyZ JURZQRQIRUPHUDJULFXOWXUDOVRLO@/HĞQH3UDFH%DGDZ- VRVQRZ\FK 3XV]F]\ 1LHSRáRPLFNLHM >$ERYHJURXQG F]H )RUHVW5HVHDUFK3DSHUV ± biomass and annual production in stands of the 7UDPSOHU 7 .OLF]NRZVND $ 'P\WHUNR ( 6LHUSLĔVND 1LHSRáRPLFND)RUHVW@6\OZDQ ± $ 0DWXV]NLHZLF] : 5HJLRQDOL]DFMD 2U]Há62FKDá:)RUJLHO0 6RFKD-E%LRPDVD SU]\URGQLF]ROHĞQD QD SRGVWDZDFK HNRORJLF]QRIL]MR- LURF]QDSURGXNFMDGU]HZRVWDQyZGĊERZ\FK3XV]F]\ graficznych [Nature and forest regionalisation on the 1LHSRáRPLFNLHM>%LRPDVVDQGDQQXDOSURGXFWLRQRIRDN ecological and physiographic basis]. PWRiL, Warsza- VWDQGV LQ WKH 1LHSRáRPLFND )RUHVW@ 6\OZDQ wa. 30–43. 7UHQGHOHQEXUJ5 0D\HU:HJHOLQ+ 'DV+RO] 3ODQ 8U]ąG]DQLD *RVSRGDUVWZD /HĞQHJR 1DGOHĞQLFWZD als Rohstoff, München. Przymuszewo na okres 01.01.1999–31.12.2008. RDLP 7XV]\ĔVNL0 :áDĞFLZRĞFLJOHESRUROQ\FKDJRV- 7RUXĔ >6WDWH )RUHVW 0DQDJHPHQW 3ODQ IRU WKH 3U]\- SRGDUNDOHĞQD>3URSHUWLHVRISRVWDJULFXOWXUDOVRLOVDQG PXV]HZR )RUHVW 'LYLVLRQ IRU WKH SHULRG RI -DQ IRUHVWHFRQRP\@6\OZDQ&;;;,9 ± ± 'HF 5'6)7RUXĔ@ 8UL9/RKPXV.2VWRQHQ,7XOOXV+/DVWLN5 9LO- 5\NRZVNL. 3UREOHP\RFKURQ\ODVXQDJUXQWDFK do M., 2007, Biomass production, foliar and root char- porolnych [Problems of forest protection on afforested acteristics and nutrient accumulation in young silver DJULFXOWXUDOJURXQGV@6\OZDQ&;;;,9 ± ± birch (Betula pendula Roth.) stand growing on aban- 6DOELWDQR) HG +XPDQLQIOXHQFHRQ)RUHVW(FR- GRQHGDJULFXOWXUDOODQG(XURSHDQ-RXUQDORI)RUHVW5H- system Development in Europe. Pitagora Editrice, Bo- search 126: 495–506. logna. 8UL97XOOXV+ /RKPXV.%LRPDVVSURGXFWLRQ 6REF]DN 5 2 SU]\ZUDFDQLX ODVyZ QD JUXQW\ SR- and nutrient accumulation in short-rotation grey alder UROQHZ3ROVFH>2Q5HVWRUDWLRQRIWKH)RUHVWRQ)RU- (Alnus incana (L.) Moench) plantation on abandoned mer Agricultural Waste Land in Poland]. Sylwan CXL, DJULFXOWXUDOODQG)RUHVW(FRORJ\DQG0DQDJHPHQW 1996(5): 35–41. 161–179. 6SáDZD1H\PDQ63D]GURZVNL: 2ZF]DU]DN= Watkins Ch., 1993 Ecological Effects of Afforestation. Biometryczne parametry budowy drewna sosny zwyc- Studies in the history and ecology of afforestation in zajnej (Pinus sylvestris/ ZDVSHNFLHZLĊĨE\VDG]HQLD Western Europe. CABI International Redwood Press, upraw [Biometric parameters of the Scots pine (Pinus Melksham. sylvestris L.) wood in the aspect of initial plantation :KLWWDNHU5+ /LNHQV*( &DUERQLQWKHELRWD VSDFLQJV@)ROLD)RUHVWDOLD3RORQLFD ± >LQ @*0:RRGZHOO(93HFDQ HGV &DUERQDQGWKH Szujecki A., 1990, Ekologiczne aspekty odtwarzania eko- Biosphere. U.S. Atomic Energy Comission, Washing- V\VWHPyZOHĞQ\FKQDJUXQWDFKSRUROQ\FK>(FRORJLFDO ton DC: 281–302. aspects of re development of forest ecosystems on old :LONRĔ0LFKDOVND-1LHQDUWRZLF]$.XQ]0 'HSWXáD IDUPODQGV@6\OZDQ&;;;,9 ± ± M., 1999, Old land-use maps as a basis for interpreting 66 Miłosz Deptuła, Andrzej Nienartowicz, Marta Iwicka, Anna Filbrandt-Czaja
of the contemporary structure of forest communities in lues of carbon accumulated in Scots pine stands [in:] =DERU\/DQGVFDSH3DUN3K\WRFRHQRVLV ± Economic evaluation of forests and their functional :LWNRZVND- /DFKRZLF]+$QDOL]D]PLHQQRĞFL SDUWV.RáREU]HJ ±=DNáDG=DU]ąG]DQLD JĊVWRĞFL XPRZQHM GUHZQD VRVQ\ ]Z\F]DMQHM Pinus =DVREDPL /HĞQ\PL ,QVW\WXW %DGDZF]\ /HĞQLFWZD sylvestris/ Z]GáXĪZ\VRNRĞFLSQLDZ]DOHĪQRĞFLRG >'HSDUWPHQW RI )RUHVW 5HVRXUFHV 0DQDJHPHQW ,Q- Z\EUDQ\FKF]\QQLNyZ>$QDO\VLVRIYDULDWLRQLQSXUH VWLWXWH RI )RUHVW 5HVHDUFK@ http://www1.up.poznan. density of Scots pine wood (Pinus sylvestris L.) along pl/kel/sites/default/files/dok_konferencje/6.%20 DWUXQNKHLJKWGHSHQGLQJRQVHOHFWHGIDFWRUV@3U]HJOąG :\VRFND)LMRUHN(=DM&F6 Papierniczy 68(9): 573–578. Warto%C5%9B%C4%87%20wi%C4%85zanego%20 :LWNRZVND- /DFKRZLF]+=PLHQQRĞüJĊVWRĞFL w%C4%99gla%20w%20drzewostanach%20sos- umownej drewna sosny zwyczajnej (Pinus sylvestris nowych.pdf [Assessed 24.03.2017]. / Z]DOHĪQRĞFLRGZ\EUDQ\FKF]\QQLNyZ>9DULDELOLW\ ;LDR &: &HXOHPDQV 5 $JURPHWULF UHODWLRQ- of conventional wood density of Scots pine (Pinus syl- ships for below and above-ground biomass of young vestris L.) depending on the selected factors], Sylwan 6FRWV SLQHV )RUHVW (FRORJ\ DQG 0DQDJHPHQW 157(5): 336–347. 177–186. :yMFLN5 6XNFHVMDZWyUQDQDJUXQWDFKSRUROQ\FK ;LDR &: &XULHO