Leaf characteristics and morphophysiological features of selected vascular plants in goltsy deserts of the Khibiny Mountains (Kola Peninsula)

Zonal tundra vegetation occupies slightly sloped watershed surfaces, weakly convex tops and gentle slopes of moraine hills and ridges with moderate snow cover and loamy soils (plakors). Environmental conditions of such sites are most relevant to macroclimate (Aleksandrova, 1971; Matveyeva, 1998). For the East European sector of the Arctic this vegetation was described in 30–70 years of last century by the Soviet geobotanists V. D. Aleksandrova (1956), ­ V. N. Andreyev (1932), I. D. Bogdanovskaya-Giye­nef (1938), A. A. Dedov (2006), A. E. Katenin (1972), Z. N. Smirnova (1938), who, following the dominant approach, attributed the described associations mainly to the moss vegetation type. In the Asian sector of the Arctic (Yamal and Taymyr peninsulas, Arctic Yakutia, Wrangel Isl.) and in Alaska some associations of zonal communities with Carex bigelowii s. str., C. bigelowii subsp. arctisibi­rica and C. lugens have been described according to Braun-Blanquet approach: Carici arctisibiricae–Hylocomietum alaskani Matveyeva 1994, Dryado integrifoliae–Caricetum bigelowii Walker et al. 1994, Salici polaris–Hylocomietum alaskani Matveyeva 1998, Carici lugentis–Hylocomietum alaskani Sekretareva 1998 ex Kholod 2007, Salici polaris–Sanionietum uncinatae Kholod 2007, Tephrosero atropurpureae–Vaccinietum vitis-idaeae Telyatnikov et Pristyazhnyuk 2012, Festuco brachyphyllae–Hylocomietum alaskani Lashchinskiy ex Telyatnikov et al. 2014. Our study area in the East European tundras (730 km of south–north and 550 km of west–east directions) covers 3 tundra subzones (arctic, typical and southern) and two floristic subprovinces (Kanin-Pechora and Ural-Novaya Zemlya) (Fig. 1). 7 associations (one with 5 subassociations) based upon 101 authors’ relevés as well 95 ones by geobotanists-predecessors were described or validated on plakors and habitats close to these. Zonal communities are comprised by thick multi-species moss layer formed by mesophylous bryophytes (Aulacomnium turgidum, Hylocomium splendens, Ptilidium ciliare, Racomitrium lanuginosum and Tomentypnum nitens), the presence of Carex bigelowii subsp. arctisibirica, Deschampsia borealis or D. glauca in the herb layer, the high abundance of dwarf-shrubs, the presence, but not always, of Dryas octopetala and shrubby willows. Their plant cover is closed or discontinuous with frost-boils (3-component module of patch of bare ground – rim – trough or 2-component one of flat surface – patches of bare ground — see Fig. 2, 3). Zonal syntaxa are the richest in species number, compare to all others because the placor habitats are moderate in such important environmental characters as moisture and nutrition of soil and snow depth. That’s why they contain, with the same constancy and sometimes abundance, some character species of alliances and classes of intrazonal vegetation: Kobresio-Dryadion Nordh. 1943 (dryad fell-fields on well drained snowless or poor snowy habitats with slightly carbonated loamy-gravelly soils at outcrops of bedrock) and Carici rupestris–Kobresietea bellardii Ohba 1974, Loiseleurio-Arctostaphylion Kalliola ex Nordhagen 1943 (dwarf-shrub and dwarf-shrub-lichen (often with Betula nana) communities on sandy soils) and Loiseleurio procumbentis–Vaccinietea Eggler ex Schubert 1960, Rubo chamaemori–Dicranion elongati Lavrinenko et Lavrinenko 2015 (dwarf-shrub-cloudberry-moss (Dicranum elongatum, Polytrichum strictum)-lichen communities of oligotrophic palsa and polygon peatlands) and Oxycocco-Sphagnetea Br.-Bl. et Tx. ex Westhoff et al. 1946. The basic syntaxon, whose communities occupy the placor habitats in the arctic tundra subzone (southern­ variant) is Salici polaris–Polytrichetum juniperini Aleksandrova 1956, described on the Southern Island of Novaya Zemlya (Table 1). Similar syntaxa in the typical tundra subzone are Carici arctisibiricae–Hylocomietum splendentis Andreyev 1932 nom. mut. propos. hoc loco (Table 5, Fig. 14–17) and Dryado octopetalae–Hylocomietum splendentis Andreyev 1932 nom. mut. propos. hoc loco salicetosum nummulariae (Bogdanov­skaya-Giyenef 1938) subass. nov. (stat. nov.), nom. corr. hoc loco, described by us and earlier by I. D. Bogdanov­skaya-Giyenef (1938) and Z. N. Smirnova (1938) on the Kolguyev Isl. (Table 2, Fig. 3, 5, 6); D. o.–H. s. caricetosum redowskianae subass. nov. hoc loco and D. o.–H. s. caricetosum arctisibiricae (Koroleva et Kulyugina in Chytrý et al. 2015) subass. nov. (stat. nov.) hoc loco (Table 4, Fig. 4, 9–13) — in the most eastern part of the studied area (Vaygach Isl., Yugorskiy Peninsula and Pay-Khoy Range); D. o.–H. s. typicum subass. nov. hoc loco (Tab­le 3), described by us with use the V. N. Andreyev (1932) relevés on Vangureymusyur Upland (Bolshezemelskaya tundra). In the southern tundra subzone the basic zonal association is Calamagrostio lapponicae–Hylocomietum splenden­tis ass. nov. hoc loco (Table 6, Fig. 20–22). Even small deviations from placor habitat conditions are reflected in the community species composition. In such habitats the following syntaxa are described: Deschampsio borealis–Limprichtietum revolventis Aleksandrova 1956 nom. mut. propos. hoc loco and Flavocet­rario nivalis–Dryadetum octopetalae Aleksandrova 1956 nom. mut. propos. hoc loco on gentle slopes and loamy soils, not in moderate soil moisture, but in wet or, on the contrary, well-drained ecotopes on the Novaya Zemlya (Table 1); Dryado octopetalae–Hylocomietum splendentis caricetosum capillaris subass. nov. hoc loco — on the deluvial tails, in the mid and lower parts of the gentle slopes in Bolshezemelskaya and Malozemelskaya tundras (Table 4, Fig. 2, 7, 8); Oxytropido sordidae–Hylocomietum splendentis ass. nov. hoc loco — in the Pakhancheskaya Bay area (the northern part of the Bolshezemelskaya tundra) on strongly sloping well drained slopes (Table 6, Fig. 18, 19). We attributed these syntaxa to zonal vegetation due to the presence of such taxa of its differential combination as shrub Salix glauca, dwarf-shrub Salix polaris, herbs Bistorta major, Carex bigelowii subsp. arctisibirica, Deschampsia borealis, D. glauca, Eriophorum brachyantherum, Juncus biglumis, Luzula arcuata, Pedicularis lapponica, Petasites frigidus, Poa arctica, Saxifraga hieracifolia, S. hirculus, Stellaria peduncularis, Valeriana capitata, mosses Aulacomnium turgidum, Hylocomium splendens, Ptilidium ciliare, Racomitrium lanuginosum, Tomentypnum nitens and lichens Lobaria linita, Nephroma expallidum, Protopannaria pezizoides, Psoroma hypnorum. This combination of taxa differentiates (by the presence, constancy, abundance) the zonal communities in studied area from vegetation of other classes (Carici rupestris–Kobresietea bellardii, Loiseleurio procumbentis–Vaccinietea, Oxycocco-Sphagnetea) (Table 7). The borders of many species area distribution are held in the East European tundras, so the variation of the community species composition along the latitudinal and longitude gradients is quite natural. Thus, in zonal communities Ledum palustre subsp. decumbens and Salix phylicifolia occur and Betula nana as well as hypoarctic dwarf-shrubs Arctous alpina, Empetrum hermaphroditum, Vaccinium uliginosum subsp. microphyllum­ and V. vitis-idaea subsp. minus are most active only in the southern tundra subzone; Salix polaris (its activity increases to the north) and, in some syntaxa, Dryas octopetala are common in the subzones of typical and arctic tundras. In zonal conditions shrubs Salix glauca, Betula nana (prostrate) and all hypoarctic dwarf-shrubs occur in the typical tundra subzone on the mainland and on Kolguyev Isl., while in the northern part of this subzone on Vaygach Isl. they are already absent, ­except the Vaccinium spp. (with low constancy). In the arctic tundra subzone there are no shrubs and hypoarctic dwarf-shrubs on plakors, while Salix polaris is abundant. We believe that these floristic differences of zonal communities can be considered as markers of their subzonal affiliation. A similar shift in species distribution on the latitudinal gradient is established (Matveyeva, 1998) for the zonal biotopes on Taymyr Peninsula. Some species (Arctagrostis latifolia, Cerastium regelii subsp. caespitosum, Saxifraga oppositifolia, Silene acaulis) have high constancy in zonal communities within the Ural-Novaya Zemlya subprovince, as opposed to the Kanin-Pechora one. Herbs Oxyria digyna, Papaver polare, Parrya nudicaulis, Pedicularis sudetica subsp. arctoeuropaea, Saxifraga cernua and S. cespitosa occur with high constancy only in zonal communities on Novaya Zemlya that brings them closer to syntaxa described in the arctic and typical tundra subzones on Taymyr Peninsula (Matveyeva, 1994, 1998). Already in 1994, N. V. Matveyeva stated the need to describe a new class for zonal vegetation. The name Carici arctisibiricae–Hylocomietea alaskani cl. prov. has been reserved for this class in Yalta’s conference on the classification of Russian vegetation (Lavrinenko et al., 2016), Prague’s “Circumpolar Arctic Vegetation Archive and Classification Workshop” (presentation by N. V. Matveyeva) and “Arctic Science Summit Week 2017” (Lavrinenko et al., 2017). We do not attribute the newly described syntaxa to alliance Dryado octopetalae–Caricion arctisibiricae Koroleva et Kulyugina in Chytrý et al. 2015, which was described at the base of 15 relevés by geobotanists-predecessors (V. N. Andreyev, A. A. Dedov) and as well the 11 ones by E. E. Kulyugina for zonal habitats in the East European tundras (Koroleva, Kulyugina, 2015). At least, it is necessary to revise this alliance, since the name of ass. Pediculari oederi–Dryadetum octopetalae (Andreev 1932) Koroleva et Kulyugina 2015 are not legitimate (nomen superfluum), ass. Salici reticulatae–Dryadetum octopetalae Koroleva et Kulyugina 2015 need to be revised and the rank of the third one (Dryado octopetalae–Caricetum arctisibiricae Koroleva et Kulyugina in Chytrý et al. 2015 was lowered by us (in this paper) to subass. Dry

The tiny fishing settlement of Tareya (73.253389° N, 90.596806° W) on the right bank of the river Pyasina (Fig. 1, this and others see in text) in its middle reaches (Western Taymyr) is well known in the circumpolar scientific community due to the long-term Biogeocenological field station of the Komarov Botanical Institute of the Academy of Sciences of the USSR, which operated in 1965-1977. A huge amount of complex researches has been done by numerous scientists, and the results were published in a lot of proceedings, reports at the Arctic conferences, and papers published in various journals, which formed the basis of several monographs as well as the large article in the multi-volume international edition «Ecosystems of the world» [Chernov, Matveyeva, 1997]. It was the reason why just this site was considered as point number one for doing work within the project “Back to the Future” (hereinafter BTF). The idea of visiting the sites of long-term work carried out in circumpolar Arctic within UNESCO “International Biological Program” arose in connection with the popular concept of global warming. The BTF task suggested to assess the current state of arctic ecosystems in details studied half a century ago. In several sites in the North American Arctic this was achieved on the eve of the International Polar Year (2008) [Callaghan et al., 2011a]. The Taymyr trip, took place in July–August 2010. Only the first author worked at the station from its beginning in 1965 and last time was there 40 years ago (1970). The period of field works in 2010 (July 21 – August 8), was not promising for detailed researches due both to the extremely short (18 days) stay and unfavorable weather. Botanists managed to re-inventory the flora of vascular plants and assess their activity in landscape, to make relevés at two permanent experimental stands and selectively some communities as well walk around the territory with vegetation map [Matveyeva, 1978]. The results on the flora were published [Matveyeva et al., 2014]. This paper presents the results of assessing the state of plant cover. We were well aware that opportunities for such a short time of repeated study in assessing the state of ecosystems and making not just expert conclusions about any changes, but to evaluate these quantitatively and to explain their reasons, were minor. In our case, different not only at moments far apart in time, but also at the same time in the past were the methodology doing relevés, including the size of sample plots, the totality of species records and quantitative assessment of their presence in communities, as well as professionalism by researchers, including their field work experience. We kept all this in mind when assessing the results, trying to distinct objectivity, subjectivity and expertise when interpreting these. In the past, detailed studies were carried out at six permanent sites [see: Matveyeva, 1968, 1969; Matveyeva et al., 1973], the most important of which were zonal communities on watersheds – frost-boils and hummock stands. DRyad–sedge–moss frost-boils stand (Matveyeva, 1968: Fig. 1, 3-5, Table 1; Matveyeva et al., 1973: Fig. 4, Table 1. Site N 2) is located on terrace above the floodplain close to high river bank of approx. 10 m high. In the checklist of Taymyr communities, according to the dominant classification [Matveyeva, 1985] it is classified as the ass. Hylocomium splendens var. alaskanum+Aulacomnium turgidum+Tomentypnum nitens–Carex ensifolia+Dryas punctata; according to the Zürich-Montpellier (hereinafter Z-M) school floristic one (Braun-Blanquet (hereinafter B-B) approach) – to the ass. Carici arctisibiricae–Hylocomietum alaskani Matveyeva 1994. In the past, the relevés were carried out on two sample plots located in close proximity to each other, 10 × 10 m (in 1966) and 15 × 15 m (in 1969), with lists of species (vascular plants, mosses, liverworts, lichens) according to 3 nanorelief elements (soil patch, rim, trough), with measurements of their size, and the horizontal structure schemes on both. We did not find those sample plots in 2010, so the relevé was performed on a new plot of 10 × 10 m. Only vascular plants were guaranteed to be identified totally with assessment of their abundance/cover on the B-B scale while that of bryophytes and lichens was estimated only for the most common and large-sized species, relatively easy identified in the field. Due to nanorelief of cryogenic genesis community horizontal structure is of 3-item regularly cyclic type [see. Matveyeva, 1988, 1998], with module repeating in space: soil patch (up to ~0.8 m diam.) at different stages of overgrowth on the medallion (up to 1.3 m diam.) +rim along its periphery (up to ~0.5 m wide)+trough (~0.3 m wide) between medallions (Fig. 2). This type of horizontal structure was preserved in 2010, although some values of element sizes were close, but not identical (Appendix 1, Table П1). However, the fact that after more than 40 years the number of modules per 100 m2 (32 and 31) and the ratio of their elements (patches 30%, rims 50%, troughs 20%) are the same, is rather evidence in favor of the horizontal structure stability, with variances due to measurement error of items widely varying in shape. Visually, the share of bare soil decreased slightly (no more than 2-3%), that caused a minor increase of total community plant cover, ~90% in 1966, 1969. up to ~92% in 2010. The dominating species in the ground layer were Hylocomium splendens var. alaskanum, Aulacomnium turgidum, Tomentypnum nitens, in the sparse upper one – Carex bigelowii ssp. arctisibirica and Dryas punctata. There were 197 species (60 vascular plants, 49 mosses, 27 liverworts, 61 lichens) on two sample plots, being different (135 and 180) on each one, due to some nuances of methodology (the lower number in 1966 is the work of a beginning graduate student, while later is the professional job by specialists in bryophytes; the lichen number was close because lichenologists were working on both plots). This community is the richest in species known in circumpolar Arctic [Matveyeva, 2009]. In 2010, on 10 × 10 m plot the composition of vascular plant species was identified with assessment of their abundance/coverage in points on the B-B scale for the entire area; that of bryophytes and lichens was estimated only for the most common and large-sized species. The most abundant ( 1%) species in the sparse low dwarf shrub-herbaceous layer were the same as before – sedge Carex bigelowii subsp. arctisibirica and dryad Dryas punctata. 11 species (all previously with low abundance/occurrence or single specimen) were not found, including two (underlined) in the past were recorded only on one of the two plots – Androsace chamaejasme, Cardamine bellidifolia, Koeleria asiatica, Orthilia obtusata, Papaver pulvinatum, Pedicularis capitata, P. hirsuta, Petasites frigidus, Nardosmia gmelinii, Ranunculus nivalis, Saxifraga oppositifolia, Vaccinium vitis-idaea subsp. minus) and found, also single specimen, 6 (Carex misandra, Eriophorum brachyantherum, Hedysarum arcticum, Polygonum bistorta, Ranunculus affinis, Saxifraga foliolosa). Such small variances gave practically the same species richness of vascular plants – 55/57 and 56. The abundance of species and their pattern at nanorelief elements remained unchanged except the cover increase of the most active species in the landscape – sedge Carex bigelowii subsp. arctisibirica. For entire community with rims occupying half of its area, this gives an increase of ~10% in layer density, i. e. the sedge abundance over the whole area remained the same (2 points). As cryptogams composition was not completely assessed, we cannot comment their richness, however all co-dominants in ground layer (mosses Hylocomium splendens var. alaskanum, Aulacomnium turgidum, Tomentypnum nitens and liverwort Ptilidium ciliare), as well species with previously significant ( +) cover kept their abundance. The obtained results provide the basis for a partly objective, partly expert conclusion that there are no significant changes in the composition of species and in their distribution within this stand. DRyad–sedge–moss hummock stand [Matveyeva, 1968: Fig. 6, 8, Table 1; Matveyeva et al., 1973: Fig. 3, Table 1. Site N 1] is located on the first terrace above the floodplain in the upper part of stream valley gentle slope at 1.5 km from the riverbank. In the checklist of Taymyr communities, according to the dominant classification [Matveyeva, 1985] it is classified as the ass. Hylocomium splendens var. alaskanum+Aulacomnium turgidum+Tomentypnum nitens–Carex ensifolia+Dryas punctata. This community with closed cover is the same in dominants as the above frost-boils one: Hylocomium splendens var. alaskanum, Aulacomnium turgidum, Tomentypnum nitens, and Ptilidium ciliare in the ground layer, and Carex bigelowii ssp. arctisibirica and Dryas punctata in the sparse upper one. Despite the common dominants and significant number of species with similar abundance, communities with closed cover are poorer in species due to the lack of species obligate to bare or partly overgrown soil. The positioning of such communities in the classification of the Z-M school (B-B approach) was not proposed. In the future, it is possible either to describe new association or to identify a subassociation. There is nanorelief of cryogenic genesis, caused by frost ground cracking and its consequences – hummocks 0.10-0.12 m high and 0.15-0.30 m diam. which sometimes, merging together, form chains or almost locked rims, and troughs 0.15-0.20 m wide, with no patches of bare soil (Fig. 3). The type of horizontal structure is irregular mosaic (Matveyeva, 1988). In 2010, what awaited us in this community was not just a surprise, but rather a shock. A transformation took place that we [Matveyeva et al., 2011; Matveyeva, Zanokha, 2013] formulated as “polygonization” of loamy watersheds – the previously leveled surface (with described nanorelief) turned into a system of mounds (7-10 m diam.) and trenches (2-5 m wide) with significant (0.5-1.0 m) excess in height (Fig. 4). In terms of the area size and the pattern of heterogeneity with rows of mounds and trenches, these are most similar to the massifs of bajdzharakhs (the Yakutian name for mounds that appears a result of the fossil ice wedge melting). Such serious changes occurred without disturbances in the plant cover, as well as in the absence of erosion, with the previous nanorelief and the same irregular mosaic type of horizontal structure both on the surface of mounds and their almost vertical slopes, and in trenches. Since there were no signs of this until 1994 (evidence from colleagues who worked here after 1970), and the system already existed in 2003 (Google Earth Quick Birds, 8.11.2003), the transformation has occurred in less than 9 years. We were not able to find the old sample plots in 2010, and only a wooden stick and small (10 × 20 and 50 × 50 cm) metal frames (used for horizontal structure study) near it convinced us that this was the same permanent stand. More than 40 years later, the horizontal structure on both new microrelief elements looked the same: the familiar combination of hummocks and troughs, but visually the surface became smoother due to the decrease in the height of the elements relative to each other. The link of species with nanorelief elements did not change, with the same dominants on hummocks (mosses Hylocomium splendens var. alaskanum and Aulacomnium turgidum, sedge Carex bigelowii ssp. arctisibirica and dwarf shrub Dryas punctata) and in troughs (Тоmentypnum nitens and Ptilidium ciliare and the same vascular plants but with lower abundance). In general, the variances in species composition between the sample plots in 1966 and 1969 were similar to those recorded in the frost boils stand, but noticeably more dissimilar (69 and 141), and not only in cryptogams but in vascular plants (Appendix 1, Table П3). In 2010, full information was obtained only about vascular plants: 43 species (32 and 33 on 2 mounds) with the same dominants both on mounds and in trenches that were previously on the flat stand surface. The abundance of sedge Carex bigelowii ssp. arctisibirica has increased up to 3 points versus 2 and that of cotton grass Eriophorum angustifolium to 2 versus 1 and +, with the same abundance of dwarf shrubs Dryas punctata and Cassiope tetragona. We found no changes in species composition or abundance in dry trenches compare to the formerly flat surface of the community and the current mound one. The second object is 3-element rim-polygonal mire. RIM-POLYGONAL MIRE [Matveyeva et al., 1973: Fig. 4, Table 3. Site N 4] in 1969 was located in: flat-concave lake depression on a river terrace above the floodplain in about 1 km from the riverbank. There are from hundreds to thousands of modules polygon center+rim+trench – wet polygon 15-20 m diam. with 1) concave center and 2) rim along it periphery 1.0-1.5 m wide, rising (0.15-0.20 m) above central part and 3) water trenches between polygons in a polygonal system (Fig. 6). Quite arbitrarily, without assigning their vegetation to any units of any classification, lists of species were made for three microrelief elements. Altogether there are 110 species (vascular plants 24, mosses 47, liverworts 24, lichens 15) were identified, with respectively 34 (10, 24, 0, 0) on polygon centers, 80 (16, 28, 21, 15) on rims, and 34 (8, 23, 3, 0) in trenches. Co-dominants in continuous moss layer are Cinclidium latifolium, Sarmentypnum sarmentosum, Scorpidium revolvens, Meesia triquetra on polygon centers and in trenches, and Aulacomnium turgidum, Hylocomium splendens var. alaskanum, Tomentypnum nitens on rims; these in the sparse above moss layer are Carex aquatilis subsp. stans and C. chordorrhiza on polygon centers and Carex aquatilis subsp. stans in trenches, and Betula nana, Dryas punctata and Salix pulchra on rims. The classification of such complex object is debatable in all respects, beginning from the relevé methodology (choice of sample plots, their size, number) as well as defining the object status. It is most logical to consider the plant cover of each of the 3 elements as communities, trying to classify them independently, however this is not too obvious: there are 18 numbers in the scheme legend, that demonstrates both the obvious cover complexity (3 types of communities) and the mosaic nature of each type – 7 units on polygon centers, 8 on rims, 3 in trenches. In the Z-M school system (B-B approach), vegetation on polygon centers and in trenches is classified as mires of the class Scheuczerio–Caricetea nigrae (Nordh. 1936) R. Tx. 1937; while that on rims as communities close to zonal ones of the class Carici arcrtisibiricae–Hylocomietea alaskani Matveyeva Lavrinenko 2023 (ass. Carici arcrtisibiricae–Hylocomietum alaskani Matveyeva 1994). In 2010, we not only failed to make relevé on previous sample plot, but could not determine its exact location in wet depression. This was because the general picture of microrelief in the area, where site in question was situated, was so different from described above, that an attempt to obtain a photo of a “classical” rim-polygonal mire for a lecture course for students (which was so easy to do before) turned in vain: there were only isolated hummocks due to partial going down (subsidence) of most rims (Fig. 7), In another massif (south of Lake Bolshoye), which vegetation on map [Matveyeva, 1978] is shown as a 3-item rim-polygonal mire, all rims went downwards, and the polygon surface became flat (Fig. 8, а). As a result, the previously clearly heterogeneous plant cover visually (from a human height) became looked homogeneous. Although heterogeneity remained (Fig. 8, б): in 2010, obviously hygrophilic grasses (Carex aquatilis subsp. stans, Eriophorum medium, Hierochloë pauciflora) and mosses (Sarmentypnum sarmentosum, Cinclidium latifolium, Scorpidium revolvens, Meesia triquetra, etc.) and just as obviously mesophilic shrub/dwarf shrub (Betula nana and Dryas punctata) and mosses (Aulacomnium turgidum, Hylocomium splendens var. alaskanum, Tomentypnum nitens, etc.) cohabit at the same surface level with high soil moisture. Anyone who has seen this would not be able to find an adequate explanation for this phenomenon without knowing the past of such areas. Our expert conclusion is that, despite significant transformations in microrelief, the heterogeneity of plant cover as well as species composition are the same as before, with slight change in the abundance of some dominants. Another type of polygonal complexes is developed in the upper reaches of numerous brook valleys. BOG-TUNDRA POLYGONAL COMPLEX [Matveyeva et al., 1973: Fig. 5, Table 3. Site N 3] in 1969 was located on a river terrace above the floodplain in 1 km from the riverbank in a depression in the upper reaches of a short valley directly close to settlement. The structure of sample plot (50 × 60 m) is a complex of drained polygons of diverse shape and size (15-30 m diam.) and trenches (0.5-6.0 m wide and 0.2-0.3 m deep), filled with water (Fig. 9). The area ratio polygons/trenches is 80/20%. The name of the complex reflects the heterogeneity of its vegetation. Plant cover on polygons is close to that of low watersheds with dominance of willows Salix reptans, S. pulchra and dwarf birch Betula nana in the shrub layer, sedge Carex bigelowii subsp. arctisibirica and cotton grass Eriophorum angustifolium and dwarf shrubs Dryas punctata, Cassiope tetragona, Vaccinium vitis-idaea subsp. minus in dwarf shrub–herbaceous, and Aulacomnium turgidum, Hylocomium splendens var. alaskanum, Tomentypnum nitens in moss one; and mire in trenches with the same shrubs as on the polygons, sedge Carex aquatilis subsp. stans and cotton grass Eriophorum angustifolium and hygrophilic mosses Sarmentypnum sarmentosum, Cinclidium latifolium, Scorpidium revolvens, Meesia triquetra, Polyrichum jensenii. There were 85 species (35 vascular plants, 41mosses, liverworts were not detected, 9 lichens), respectively – 59 (30, 20, 9) on polygons and 35 (12, 23, 0) in trenches. The classification of this object is no less problematic in all respects, as of rim-polygonal mire vegetation. Most logical is to consider the vegetation on each of two elements as communities and try to classify them separately, which is quite difficult. There are 19 numbers in the map legend – two community types with 13 inside units on polygons and 6 ones in trenches. Such complexes so far have not been described in literature. In 2010, visually everything looked as before, however this conclusion is subjective being based upon only on two routes through a vast complex system, including a stationary site with wooden sticks. TUNDRA AND NIVAL-MEADOW COMMUNITIES ON THE SOUTHERN SLOPE OF THE RIVER BANK [Matveyeva et al., 1973: Fig. 7, Table 4, Site N 5]. The steep slope, is cut by hollows (with 3-5 m snow beds) formed due to the ice wedge melting (Fig. 10). Ridges, in winter with little snow, melting completely in June, are in summer the warmest biotopes with the maximum (up to 1.5 m) depth of frozen ground seasonal thawing. The great biotope diversity determines the heterogeneity of the plant cover, with elements small (2-3 m2) in size that form ecological series, contrasting in soil moisture and heating. There are 13 community types on sample plot (70 × 70 m). The most contrasting in comparison with stands in zonal habitats were in 1969 and remained (visually) in 2010 are herb communities on ridges (Fig. 11) with grasses (Festuca brachyphylla, Koeleria asiatica, Trisetum sibiricum subsp. litorale) and forbs (Astragalus alpinus, Cerastium maximum, Myosotis alpestris subsp. asiatica, Oxytropis adamsiana, Pachypleurum alpinum, Pedicularis verticillata, Polemonium boreale) from 0.10-0.15 to 0.30-0.35 m high and thin (up to 0.01 m), and sparse moss layer of Hypnum revolutum, Sanionia uncinata, Thuidium abietinum. Later such community types became the object of close attention [Zanokha, 1993] in different areas of Taymyr (but not in Tareya), and was classified as the ass. Pediculari verticicillatae–Astragaletum arctici Zanokha 1993, but with no placing in any higher unit. The plant cover of such herb communities, in terms of life form set and horizontal and vertical cover structure is closest to boreal meadows of the class Molinio-Arrhenatheretea Tüxen 1937, however composed of not boreal, but of arctic and arctic-alpine species, that stops these from being placed in this class. As well, conditional is the positioning [Matveyeva, Lavrinenko, 2021] of such communities in the class. Mulgedio-Aconitetea Hadač et Klika in Klika et Hadač 1944. In 2010, the lists of vascular plant species were compiled for such herb communities along the whole riverbank of the field station area, and no differences were recorded in their activity [Matveyeva et al., 2014]. It is worth to notice that the methodology for getting data in the past is not described, and it differs from that adopted in the Z-M school. This will not allow objectively assessing possible changes in the future that should be kept in mind by those who will manage to visit this area. VEGETATION UNDER MAN IMPACT [Matveyeva et al., 1973; Fig. 8. Site n 6]. In 1965, when six BIN researchers arrived to Tareya, life in tiny fishing settlement was in full swing. The basis of this was a vast man-made cave in the permanently frozen ground of the high river bank. It was used to store fish that was caught by teams of fishermen from Norilsk State Industrial Enterprise, scattered across the vast Western Taymyr territory. Fishermen were flown to “points” on AN-2 planes, from where the catch was regularly taken, brought to Tareya, frozen and stored until the autumn fishing season, when ships with barges arrived along the river from the Norilsk city. There were three small houses (at the edge of the floodplain) for living and a house where the radio operator lived and worked. In addition, there was a large plank house owned by the Arctic and Antarctic Research Institute (AARI), permission for its use became the basis for organization of a long-term BIN field station (Appendix 2, Fig. П1). In the first summer (1965), the pioneer group lived in a plank house (future laboratory). The following summer, scientific field station began to function, which gathered from 18 (1966) to 30-40 (1967-1969) people from various scientific institutes, who lived in numerous tents located on a gentle slope between the laboratory and the radio operator' house. After 1977, the living buildings continued to be used by fishermen, as well as geologists. Fishing intensity gradually decreased becoming private. In a spring (the year is unknown) high flood, three small houses were carried away by water; the laboratory house was burned down in 1998. Before 1965, the plant cover was quite changed, since for a long time the base of the geological expedition of the AARI was located here. Its initial state is dryad-sedge-moss hummock tundra, common on gentle slopes with the dominance of mosses Aulacomnium turgidum, Hylocomium splendens var. alaskanum, Tomentypnum nitens, sedge Carex bigelowii subsp. arctisibirica, dwarf shrub Dryas punctata. During the field station functioning, the load (trampling) on plant cover in summer (late June–early September) was quite strong. In 1968, the vegetation of the territory was verbally described, and a map-scheme was made, with 12 items in legend [Matveyeva et al., 1973]. In 1968, the most of area between houses, where the original vegetation was damaged almost completely, was occupied by suppressed and sparse grass cover. In 2010, it looked like the original dryad-sedge-moss tundra, with no obvious signs of disturbance and with no high abundance of apophytic grasses (Alopecurus alpinus and Poa alpigena). However walking along, it at the end of July–beginning of August was possible only in rubber boots, i. e., the soil moisture was significantly higher than before, when we lived in tents and walked in light sports shoes. Vegetation map. The conclusion that in 2010 communities have kept their belonging to the same earlier classified community types is made according to their look when walking around the territory with vegetation map that would not have to be changed (Fig. 13). Some of the objectivity of this opinion is supported by the fact that it was done by the researcher who made this map, as well as by few relevés, where the community structure and species composition remained the same. Flora of vascular plants. There were 212 species on the territory that was studied in 2010 [Polozova, Tikhomirov, 1971]. After 40 years, we did not find 29 species (all rare in the landscape) and discovered 10 new ones (all in the floodplain of the Pyasina River, rare, many in a single specimen). We refer a reader to the publication [Matveyeva et al., 2014], the main conclusions of which are as follows: 1) the main reason for the incomplete identification of the flora is the short duration of the research in 2010; 2) there is no firm conviction that newly found species were absent 40 years ago; 3) assuming that the last are still present, the systematic and geographical structure flora remains unchanged. It is possible to assess changes in species activity within landscape only for a total of 184 species – in 162 (88.5%) it remained unchanged, in 5, with the same activity, abundance slightly increased or decreased; activity decreased by 1 point in 22 (mean and low active) species. Small changes in the landscape pattern of species with low activity may be considered both objective and subjective (short duration of observations in 2010 and uncertainty in estimations in the 1971 annotated list). No information was obtained on the cryptogam flora (mosses, liverworts, lichens), earlier detailed studied. Our partly expert opinion is that their composition and presence in communities have not undergone noticeable changes. However, for an objective assessment it is necessary to conduct studies similar to those that were done at high professional level [Pijn, Trass, 1971; Blagodatskikh, 1973; Zhukova, 1973]. THE DISCUSSION OF THE RESULTS. The most general conclusion based on the results of various observations in the course of repeated (after 40 years) visit to the area of long-term field station functioning can be formulated as follows: stability in the plant cover with significant transformations in the landscape, micro- and nanorelief, and as a consequence in changes in surface/inside soil water flow. From the diverse cryometamorphic processes, we focus the most significant and noticeable one, that might considerably change the plant cover on the above-floodplain terrace, where previously there were 2 systems, both in depressed landscape sites: 1) rim-polygonal mires (in lake depressions, bottoms of drained lakes of thermokarst origin) and 2) bog-tundra complexes (concave surfaces of watersheds, dissected by trenches as a result of backward erosion). The third one, with flat mounds of different height and size and trenches of various width and depth, appeared in zonal sites. (Fig. 15). This happened on a large area, lot of watersheds is transformed completely with some (most wide and flat) being so far rather uniform. The beginning or early stages of this process in the form of future polygonal system were recorded already in 1968 by geocryologist [Danilov et al., 1971]. In 2010, already in the field on many interfluves between brook valleys, especially on the widest ones, with a horizontal surface in their middle part, we observed the beginning of polygonization so far with no upcoming mound exceeding the trenches in height ( 1-2 cm), which is clearly visible on satellite images (Fig. 16). Potentially, the presence of trench system on watersheds may strengthen the hydrological cycle through higher inside soil flow (that will eliminate trench wetting), however as drainage system it will reduce the moisture amount on watersheds, that may lead to larger frozen soil seasonal thawing, and greater thermokarst in zonal sites. What will be a result of such large transformation is a subject of professional interest for geocryologists. We can only state the landscape instability, which was not recorded 40 years ago in Tareya. The second phenomenon of significant change is the coming down of rims in rim-polygonal mires, in the place of which only isolated hummocks remained, or the surface of the polygons has become flat, the most important consequence of which was a radical change in hydrological regime. In classic rim-polygonal mire systems, the water on the isolated concave polygon centers surrounded by rims is standing water, while in trenches between polygons it is running, and there is a general waterway, which gathers water from connected trenches. This is the source of brooks through which the general (surface and inside) water flow is running away the wetland (Fig. 17). Without rims, the previously standing water on polygons, being no longer isolated, has become running, that increased the total flow (a kind of drainage). On the downed rims, the plant cover is so far (visually) the same. Although the fact that the mire, heavily watered throughout the growing season in 1967-1969 (and according to satellite image in 2003), in 2010 has lost part of water, affected the activity of the most important grasses – the abundance of Hierochloë pauciflora and Carex chordorrhiza previously dominated on the most watered polygons became less, while that of Carex aquatilis subsp. stans (previously also rather abundant) increased. This expert conclusion is based on difficulty in finding the first two species, which previously were common in these biotopes. At first sight, our judgment about stability in plant cover along with great landscape transformation, looks at least contradictory. In our defense, we propose thesis that stability does not mean the absence of any changes. The latter includes changes in the activity (abundance, occurrence) of some vascular plant species, dominants in communities in zonal sites (Carex bigelowii subsp. arctisibirica) and in mires (Carex chordorrhiza, C. aquatilis subsp. stans, Hierochloë pauciflora). However, for the majority (88.5%) of species it remained unchanged; for few ones the abundance slightly increased or decreased, which did not cause noticeable changes in the structure of communities and their diversity. To explain the slight increase in the cover density on ground patches in frost-boils stands and that of main dominant, the long-rhizome sedge Carex bigelowii subsp. arctisibirica, is hardly makes sense to attach the argument, most common in the last decade, about global warming. A series of questions arises – what do we know about vegetation before we worked in this area 40 years ago? how much do we know about the life cycles of Arctic species populations, about the species individual growth? as well, without single-vector climate trend, changes in vegetation do not occur? or we ignore natural succession? Our conclusion about the stability of syntaxonomic diversity, with small changes in the communitiy structure and with minor variation in vascular plant species set in local flora and their activity in landscape, in general coincides with the opinion of colleagues, who worked within the BTF project in Canada and Greenland, and repeated studies over shorter periods in Alaska and the European North, differing in minor details. This is inspiring and at the same time amazing, because only on Taymyr (besides Tareya, in the Dickson area) this stability takes place against the background of spectacular landscape transformation – polygonization of watersheds and modification of rim-polygonal mires. The formation of the third polygonal system on watersheds, in addition to the widespread polygonal mires and bog-tundra polygonal complexes in depressions, may continue, which gradually lead to radical transformation of the Arctic landscape on the plains. However, to predict exactly, what consequences will follow, is difficult. The existence of new formed trenches proposes their greater moisture, in comparison with mounds and the former flat surface, but the fact that these are not isolated, but form system, suggests a drainage effect. We are not ready to predict to what extent the intra-soil moisture runoff increasing will balance or exceed the current greater moisture in trenches, this is a matter for soil scientists. However, there is no doubt that the dynamics of vegetation in zonal sites depends on this, and significant changes in the plant cover may be expected over vast areas. The data obtained by us and other researchers in different Arctic regions indicate the stability of the plant cover in the course of the period that coincides with the ascending wave of climate warming in high latitudes, which is the second in the 20th century [Vize. 1937; Rosenbaum, Shpolyanskaya, 2000; Malinin, Vainovsky, 2018], even in situations of mobile landscape.

Синтаксономия луговин тундрового пояса гор Мурманской области

Studies on terrestrial ecosystems in the high Arctic region have focused on the response of these ecosystems to global environmental change and their carbon sequestration capacity in relation to ecosystem function. We report here our study of the photosynthetic characteristics and biomass distribution of the dominant vascular plant species, Salix polaris, Dryas octopetala and Saxifraga oppositifolia, in the high Arctic tundra ecosystem at Ny-Alesund, Svalbard (78.5 degrees N, 11.5 degrees E). We also estimated net primary production (NPP) along both the successional gradient created by the proglacial chronosequence and the topographical gradient. The light-saturated photosynthesis rate (A (max)) differed among the species, with approximately 124.1 nmol CO(2) g(-1)leaf s(-1) for Sal. polaris, 57.8 for D. octopetala and 24.4 for Sax. oppositifolia, and was highly correlated with the leaf nitrogen (N) content for all three species. The photosynthetic N use efficiency was the highest in Sal. polaris and lowest in Sax. oppositifolia. Distributions of Sal. polaris and D. octopetala were restricted to the area where soil nutrient availability was high, while Sax. oppositifolia was able to establish at the front of a glacier, where nutrient availability is low, but tended to be dominated by other vascular plants in high nutrient areas. The NPP reflected the photosynthetic capacity and biomass distribution in that it increased with the successional status; the contribution of Sal. polaris reached as high as 12-fold that of Sax. oppositifolia.

The influence of abiotic factors on the growth of two vascular plant species (Saxifraga oppositifolia and Salix polaris) in the High Arctic

Ranunculus glacialis (L.) A. Löve & D. Löve is a rare species that is included in the Red Data Book of the Murmansk region. It belongs to a group of northern species that, under climate change conditions, will be exposed to a reduction of range and loss of genetic diversity. The objective of this study was to estimate the adaptive potential of this species in the Khibiny Mountains, which is the edge of the eastern limit of its range. Plants growing in natural conditions of the Khibiny Mountains and in the nurseries of the Polar-Alpine Botanical Garden-Institute (PABGI) were compared in terms of leaf mesostructure and pigment content. Under nursery conditions, at higher temperature than in the field, R. glacialis plants showed quantitative rearrangement of leaf mesostructure. Changes associated with increases in internal leaf volume and disturbance of ontogeny, changes in morphometric indicators of assimilating organs (mass and leaf area), reduced productivity and, consequently, reduced resistance to growing conditions were also found in the PABGI-cultivated plants. In this study, we show that this species has a low level of genetic diversity and a limited adaptive potential in the extreme eastern edge of its range in Russia (Kola Peninsula), as evidenced by numerous experiments on acclimatization of R. glacialis under nursery conditions in the Khibiny Mountains.

Plants respond to herbivory either by maximizing resource acquisition and compensatory growth or by minimizing loss of resources, e.g., by investing in chemical or structural defence. We studied the response to simulated browsing by the deciduous dwarf shrub, Salix polaris, on high Arctic Spitsbergen. Salix polaris is browsed by Svalbard reindeer, and its response to browsing may influence subsequent utilization. We compared leaf characteristics and flowering of S. polaris from areas with relatively short, intermediate, and long growing season, and their responses 1 yr after simulated browsing in early, mid, and late summer. Leaf numbers, total and individual biomass of leaves, and the number of inflorescences were greatly reduced the year after treatment. There was no increase in phenolics but a tendency to an increase in N content in the leaves of S. polaris 1 yr after treatment. Salix polaris showed little variation in the response to simulated browsing with local variation in resource availability (length of growing season) or with time of browsing. The results suggest that S. polaris responds to summer browsing the previous year by allocating resources to compensatory growth rather than to defence. For reindeer, browsing of S. polaris leads to a sizeable decrease in food quantity and, possibly, to a limited increase in food quality.

One of the factors of abiotic stress is ultraviolet radiation. Acting on the photosynthetic apparatus of the cell, radiation causes a change in the intensity of photosynthesis - the main physiological process that determines the growth, development and productivity of plants. Quantitative changes in the content of photosynthetic pigments and the intensity of growth processes are indirect indicators of plant resistance to the effects of various environmental factors. With the right treatment regimes, UV radiation can be used to improve the sowing quality of seeds and increase the productivity of grain crops. The article presents the results of studying the effects of small doses of UV-C radiation on laboratory germination, the content of chlorophylls a and b, carotenoids in soft spring wheat seedlings, as well as indicators of the amount of chlorophylls, the ratio of chlorophylls a / b, the ratio of the sum of chlorophylls and carotenoids. According to the results obtained, the effect of UV radiation in the studied time range (30-180 seconds) led to an increase in laboratory germination and an increase in the content of chlorophylls and carotenoids, no dependence on the change in pigment ratios on exposure time was revealed.

OPTIMIZATION OF LIGHTING SPECTRUM FOR PHOTOSYNTHETIC SYSTEM AND PRODUCTIVITY OF LETTUCE BY USING LIGHT-EMITTING DIODES

Changes in some leaf characteristics: leaf mass area (LMA), content of photosynthetic pigments and nitrogen in the leaves, leaf mass ratio (LMR) and leaf area ratio (LAR) were investigated in steppe plants of the Volga land along the gradient of aridity. When drought stress became stronger, the content of chlorophylls in the leaves, LMR and LAR decreased, whereas LMA and the proportion of carotenoids in the leaves rose. In the North to South direction, the content of pigments and nitrogen per unit whole plant weight considerably decreased (4 and 2 times, respectively). The relationship between leaf indices (chlorophyll and nitrogen contents and LMA) differed along this gradient. It was concluded that adaptation of steppe plants to drought stress generally depended on predominant development of heterotrophic tissues in the leaf and the whole plant. During aridization, the stress-tolerant species became more numerous.

PDF HTML阅读 XML下载 导出引用 引用提醒 4种高大树木的叶片性状及WUE随树高的变化 DOI: 10.5846/stxb201304020586 作者: 作者单位: 中国林业科学研究院林业研究所,华南农业大学林学院,中国林业科学研究院林业研究所 作者简介: 通讯作者: 中图分类号: Q945.79 基金项目: 教育部博士点基金资助项目(20124404110007);中国林科院林业所所长基金资助项目(RIF2013-08) Changes of leaf traits and WUE with crown height of four tall tree species Author: Affiliation: State Key Laboratory of Tree Genetics and Breeding,Research Institute of Forestry,Chinese Academy of Forest,Beijing,College of Forestry,South China Agricultural University,Guangzhou,State Key Laboratory of Tree Genetics and Breeding,Research Institute of Forestry,Chinese Academy of Forest,Beijing Fund Project: 摘要 | 图/表 | 访问统计 | 参考文献 | 相似文献 | 引证文献 | 资源附件 | 文章评论 摘要:为了解西双版纳北热带雨林高大树木树顶叶片性状对通道阻力增长引起的水力限制增强及高光和季节性干旱等气候条件的响应,对该区乔木浆果乌桕(Sapium baccatum Roxb)、思茅木姜子(Litsea pierrei var. szemois liou)、小叶藤黄(Garcinia cowa Roxb)及共生木质藤本黑风藤(Fissisfigma polyanthum (Hook. f. et Thoms.)Merr.)的叶片形态解剖结构、光合色素、水分利用效率(WUE)等随冠层高度的变化及种间差异进行了研究。结果表明:小叶藤黄和思茅木姜子的叶片(310.14、319.73 μm)和角质层(6.06、5.13 μm)都较厚、细胞较大(21.48、27.09 μm),光合色素含量则较低;黑风藤栅栏组织所占的比例最大、光合色素含量也最高,但叶片薄、WUE最低;浆果乌桕的WUE最高。随着冠层高度的增加,4种树木的叶厚、栅栏组织及角质层厚度、LMA、P/S和TPM/LT均增加、细胞变小,其中黑风藤的变幅最大。4树种的叶绿素和类胡萝卜素含量均随冠层的增高而减少,δ13C和WUE则随树冠增高而增大(黑风藤的变幅小于3种乔木);Δ则相反。上述结果表明4种树木冠层上部叶片偏向旱生结构和水分利用效率增加,暗示树顶叶片可能受到了水分胁迫,从而在结构上偏向于减少水分散失、功能上提高对水分的利用效率以适应水分亏缺;同时,随冠层增加光合色素含量减少,暗示其光合碳同化能力也降低。上述结果支持了水力限制假说中由于通道阻力增大引起树顶水力限制增强,大树可能会通过减少光和碳的获得而减慢树高生长的假设。 Abstract:Leaves as the main photosynthetic organs are sensitive to exterior environments. Generally the structure and physiological characteristics of leaves are influenced by water deficiency significantly. In the northern rainforest of Xishuangbanna in China, strong radiation and seasonal drought occur frequently. With the increasing of tree height, the xylem pathways increase, resulting in the increasing of water gravity and pathway resistance. In the rainforest, tree is higher and is liable to be influenced by hydraulic limitation. The response of treetop leaves to hydraulic limitation and seasonal drought in higher trees is particularly important to the study of seasonal rainforest. In order to understand the adaptation strategies of rainforest trees, the changes of leaf anatomical structure, pigment content, and water use efficiency (WUE) under varied tree heights of a liana plant Fissisfigma polyanthum (Hook. f. et Thoms.)Merr. and three co-existing arbors of Sapium baccatum Roxb, Litsea pierrei var. szemois liou and Garcinia cowa Roxb were studied in the rainforest of Xishuangbanna, China. Differences in leaf structure, pigment content and WUE were compared among the four tree species. The results showed that the G. cowa and L. pierrei have thicker leaf (310.14, 319.73 μm), thicker cuticle (6.06, 5.13 μm) and bigger cells (21.48,27.09 μm). In addition, their photosynthetic pigments content were lower compared to the other two species. The palisade tissue occupied larger proportion of the leaf transverse section, and the pigments content were higher in the F. polyanthum than three arbor species. However, the F. polyanthum had thinner leaf (147.67 μm) and lowest WUE, and the S. baccatum had the highest WUE. The leaf thickness, palisade tissue and cuticle thickness, LMA, P/S, and TPM/LT of measured four tree species increased with the increasing of crown height, and the F. polyanthum had the largest increasing. Meanwhile, the leaf chlorophyll and carotenoid content of four tree species reduced with the increasing of crown height. The leaf δ13C value and WUE of four tree species increased with the tree height, where the △ value has an opposite trend. The WUE of F. polyanthum had a small change along tree height than the arbor species. The above results showed that the upper crown leaves of four tree species exhibited xeromorphic structure and high WUE. These xeromorphic structure and physiological characters suggests that the trees may suffer from water stress. In order to reduce the influences of relative water shortage, the water loss of treetop leaves was minimized through reducing stomatal conductance, and the WUE was increased resultantly. Simultaneously, the reduced stomatal conductance led to the decreasing of photosynthesis. The fact of low photosynthetic pigment contents in the upper crown leaves indicates the reduced ability of photosynthetic carbon assimilation in treetop leaves. Xeromorphic structure may limit cell division and expansion, then the gas exchange and carbon assimilation capability are restricted resultantly. However, respiration consumption increases with the spread of tree crown. The nutrient shortage limits carbon investment on new leaf growth, and the tree growth is limited ultimately. These findings support the hypothesis of hydraulic limitation that the height growth of big trees slowed down through reduced absorption of irradiance and carbon, which is caused by the hydraulic limitation at treetop arising from increased pathway resistance. 参考文献 相似文献 引证文献

<p>The glacial history of the Kola Peninsula, northwest Arctic Russia, during the Last Glacial-Interglacial Transition (LGIT; c. 18-10 ka) is poorly understood, with some researchers suggesting that the region was glaciated by the Fennoscandian Ice Sheet (FIS; e.g. Hughes et al., 2016), and others suggesting that it was glaciated by an independent Ponoy Ice Cap (e.g. Astakhov et al., 2016). Furthermore, it is unclear if and where there was a periodic ice standstill during the Younger Dryas (c. 12.9-11.7 ka) cold stadial. This is the largest sector of Fennoscandia where glaciation is poorly constrained, which stems from low resolution geomorphological mapping, a lack of sedimentary analyses, and limited dating of glacial landforms and deposits on the Kola Peninsula.</p><p>Initial interpretations of geomorphological mapping and sedimentological analyses are presented. High resolution geomorphological mapping has, so far, demonstrated that the Kola Peninsula was glaciated by the FIS, which flowed from the Scandinavian mountains in the west and across the shield terrain of the Kola Peninsula, and not an independent Ponoy Ice Cap, as indicated by the west-east orientation of glacial lineations (e.g. drumlins, crag and tails, mega-scale glacial lineations), moraines, and meltwater channels. Up to four ice streams located in the western Kola Peninsula and the White Sea demonstrated in the glacial lineation record have also been identified. Furthermore, the Younger Dryas margin is proposed to be aligned north-south across the Kola Peninsula, flowing around the Khibiny Mountains, and forming an ice lobe in the White Sea, which is demonstrated by the moraine and meltwater landform assemblage. Moraines and lateral meltwater channels also suggest the Monche-tundra Mountains were exposed as nunataks, and that there were independent cirque and valley glaciers in the Lovozero and Khibiny Mountains at the periphery of the FIS during the Younger Dryas. In addition, glaciotectonised sediments identified in sedimentary analyses indicates the FIS underwent sustained readvances during retreat. This research will provide crucial empirical data for validating numerical model simulations of the FIS, which in turn will further our understanding of (de)glacial dynamics in other Arctic, Antarctic, and Alpine regions.</p><p> </p><p>Astakhov, V., Shkatova, V., Zastrozhnov, A. and Chuyko, M. (2016). Glaciomorphological map of the Russian Federation. <em>Quaternary International, 420</em>, pp.4-14.</p><p>Hughes, A.L., Gyllencreutz, R., Lohne, Ø.S., Mangerud, J. and Svendsen, J.I. (2016). The last Eurasian ice sheets - a chronological database and time-slice reconstruction, DATED-1. <em>Boreas, 45</em>(1), pp.1-45.</p>

As a part of the study on soil carbon flow in a deglaciated area in Ny-Alesund, Svalbard (79°N), we estimated the contribution of the belowground respiration of vascular plants to total soil respiration in August 1996. Four study sites were set up along a primary successional series, ranging from newly deglaciated moraine to older moraine with well-developed vegetation cover. Respiratory activity of the belowground parts (roots + belowground stems) of three dominant species, Salix polaris, Saxifraga oppositifolia and Luzula confusa, was determined under laboratory conditions. The respiratory activity and the Q10 value of the respiration were higher in S. polaris than in the other two species. Total soil respiration rates measured in the field varied widely. The areas with dense vegetation cover tended to show high respiration rates. Belowground respiration of vascular plants was estimated based on the respiratory activity and biomass of the belowground parts at each study site. The contribution to the belowground respiration to total soil respiration was negligible in the early stages of succession. On the other hand, the respiration of the belowground parts contributed to a significant proportion (?29%) of the total soil respiration in the latter stages of succession.

A comprehension of the effects planting density and nitrogen (N) fertilization have on the physiological and morphological characteristics of trees is critical for optimizing the require size and characteristics of wood products. We evaluated the growth traits and the leaf and wood characteristics of three clone poplars including Populus simonii × P. nigra ‘Xiaohei’, ‘Xiaohei-14’ and ‘Bailin-3’ under five planting densities (1666, 1111, 833, 666, and 555 tree ha−1) and four N fertilization rates (0, 100, 160, and 220 g tree−1 year−1). The results show that the clone type significantly affected all observed indicators, while planting density and N fertilization treatments had a significant effect on growth traits and leaf characteristics, but not on wood characteristics. Specifically, the clone ‘Bailin-3’ exhibited the largest annual increments in tree height and diameter at breast height (DBH), leaf width, N content, and soluble protein content. A decrease in initial planting density (from 1666 to 555 tree ha−1) led to an increased annual incremental tree height and DBH, regardless of clone type and N fertilization treatment. N fertilization treatment significantly impacted the annual increment in DBH, but not that of tree height. Further, the annual increments in tree height and DBH were positively correlated with leaf width, N content, chlorophyll content, and soluble protein content, and negatively correlated with hemicellulose content. In addition, the chlorophyll and soluble protein contents were identified as the most reliable predictors of the annual increments in tree height and DBH. Our results demonstrate the clone ‘Bailin-3’ with 555 tree ha−1 under 160 g N tree−1 yr−1 showed superior growth traits and leaf characteristics. Thus, it is recommended for future poplar silviculture of larger diameter timber production at similar sites. The results contribute to understanding of the effects of planting density and fertilization on the growth traits and the leaf and wood characteristics of three poplar clones, offering valuable guidance for the sustainable development and long-term productivity of poplar plantations.

Medieval climate warming and aridity as indicated by multiproxy evidence from the Kola Peninsula, Russia