Regeneration dynamics, causal factors, and characteristics of pacific ocean-type beech (Fagus crenata) forests in Japan: A review

This paper reviews the differences in the distribution and regeneration ofFagus crenata between two types of Japanese beech forests, the Japan Sea (JS)-type and the Pacific Ocean (PO)-type, and discusses the causal factors and characteristics of these forests, particularly the PO-type.F. crenata in PO-type forests regenerates sporadically rather than constantly, whereas regeneration in the JS-type forests is relatively constant with gap dynamics.F. crenata has dominated in snowy areas both in the past, after the last glacial age, when there was less human disturbance, and in the present. Snow accumulation facilitates beech regeneration in snowy JS-type forests, but not in the less snowy PO-type. Snow protects beechnuts from damage caused by rodents, desiccation, and freezing. In addition, snow suppresses dwarf bamboo in the spring, thus increasing the amount of sunlight available for beech seedlings on the forest floor. Snow also supplies melt water during the growing season and limits the distribution of herbivores. Moreover, snow reduces the number of forest fires during the dry winter and early spring seasons. The low densities ofF. crenata impede its regeneration, because disturbed wind pollination lowers seed fertility and predators are less effectively satiated. In snowy JS-type beech forests,F. crenata dominates both at the adult and the juvenile stages because it regenerates well, while other species are eliminated by heavy snow pressure. On the less snowy PO-side, deciduous broad-leaved forests with various species are a primary feature, althoughF. crenata dominates because of its large size and long lifespan.     

Comparison of ecophysiological responses to heavy snow in two varieties ofAucuba japonica with different areas of distribution

Aucuba japonica varieties are common evergreen understory shrubs in Japan.Aucuba japonica var.borealis is distributed on the Sea of Japan side of Honshu and Hokkaido where heavy snow cover lasts for more than 3 months in winter.Aucuba japonica var.japonica is distributed in areas with shallow or no snow on the Pacific Ocean side of Honshu and Shikoku. The ecophysiological characteristics of var.borealis were compared with those of var.japonica to examine the effects of heavy and long-term snow cover on the life cycle of var.borealis. Shoots of both varieties were shaded in crushed ice for 110 days, but their photosynthetic activities, chlorophyll contents and the chlorophylla/b ratio was not affected. The leaves of var.borealis were no less frost tolerant than those of var.japonica. In spite of the difference in environmental factors, both varieties had similar characteristics in seasonal changes of photosynthesis, respiration and chlorophylla/b ratio. These results suggest that var.japonica could survive in areas with heavy snow where it does not normally occur. Leaf net production (LNP) was estimated based on the microclimatic data and seasonal photosynthetic and respiration rates. The difference in the annual LNP between the two varieties was equivalent to the difference in the LNP during the snow season. One of the major effects of snow cover is to interrupt and reduce the production period of var.borealis.     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.     

Snowcover as a factor controlling the distribution and speciation of forest plants

Effects of climatic factors on plant distribution in the Chubu District, central Japan, were examined with direct gradient analysis. The distribution of many species was controlled not only by temperature but also by snowfall.Sasa kurilensis, Heloniopsis orientalis andTripterospermum japonicum preferred snow regardless of the thermal conditions. The snow seemed to exert more influence on endemics than on the more widespread plants, on evergreen plants more than on deciduous plants, and on trees or shrubs more than on herbs or climbing plants. Thus, the snow factor contributes to the high endemic ratio of central Japan. Among the snow-tolerant evergreen shrubs there were many neo-endemics derived from snow-intolerant parent populations of the species which have restricted distributions in the laurel forests. They could occupy habitats in the summergreen forests, where their life form can withstand heavy snowfall. Because of the heavy snowfall on the Sea of Japan side of central Japan in the postglacial period, evergreen coniferous trees have lost many of their previous habitats since most of them are intolerant of heavy snowfall.     

Differences in beech (Fagus crenata) regeneration between two types of Japanese beech forest and along a snow gradient

Differences in beech (Fagus crenata) regeneration were quantitatively investigated using power function analysis for the size–class (diameter at breast height, DBH) distribution and juvenile-to-canopy tree (J/C) ratio along a snow gradient throughout Japan. In snowy areas, all species combined, as well as F. crenata alone, showed constant regeneration, with parameter b≈−1.6 for the power function y=ax ^b (x=DBH, y=density), which is related to the DBH–class distribution. The good fit of the data to the function suggests that beech regenerates constantly with self-thinning patch dynamics. Parameter a, which indicates the abundance of small trunks, was high. Furthermore, the mean J/C ratio was ≈8, i.e., each parent beech tree produced eight juveniles. These results suggest that beech regenerates constantly with gap dynamics in snowy beech forests on the Japan Sea side of Japan (snowy). However, the fit of F. crenata was lower and nonsignificant in some forests in less snowy areas, despite the high fit of all species combined. In these areas, the mean of a was low, and b was often near zero for F. crenata regressions. These results suggest that the abundance of beech was low, and self-thinning was not evident because of the initial low abundance. Moreover, the mean J/C ratio was <1.0, suggesting that juvenile density was lower than that of canopy trees. Thus, the regeneration of F. crenata on the Pacific Ocean side of Japan (less snowy) is rather sporadic. Less snowy conditions may promote seed desiccation, predation of beechnuts and seedlings, and water stress. Lower F. crenata density may also reduce predator satiation and wind pollination.     

Ecological studies on the plants of fagaceae V. Growth in the sapling stage and minimal participation of reserve materials in the formation of annual new shoots of the evergreenQuercus glauca thunb

The growth in the sapling stage and participation of reserve materials in the formation of annual new shoots were studied in the evergreen treeQuercus glauca. The growth and some allometric relationship were analyzed for 1-to 4-year-oldQ. glauca plants. Each individual was felled at the foot, or all the leaves were removed at the end of the growth season (December). Sprouts were formed on the remaining stump and new shoots were formed from winter buds under dark or light conditions. The dry weight of each plant part was measured before and afte,, the formation of these new shoots. The amount of reserve materials in each plant part was estimated from the difference in allometric relatationships before and after the formation of new shoots. Although the results showed that a small amount of reserve material existed in roots, the participation of reserve materials in the formation of annual new shoots was negligible inQ. glauca growing under usual conditions, and substrates for the formation of new shoots were derived from the products of photosynthesis in old leaves and developing new leaves. Some of the growth characteristics ofQ. glauca were compared with those of the deciduous speciesQuercus variabilis to explain ecological behavior of the two species in warmtemperate secondary forests.     

Warming effects on shoot developmental growth and biomass production in sympatric evergreen alpine dwarf shrubs Empetrum nigrum and Loiseleuria procumbens

Effects of experimental warming on shoot developmental growth and biomass production were preliminarily investigated in two evergreen dwarf shrubs Empetrum nigrum and Loiseleuria procumbens, using the International Tundra Experiments open-top chamber (OTC) method, in the Tateyama Range, central Japan. An OTC was installed over shrub (E. nigrum and L. procumbens) -dominated vegetation and over shrub-forb (such as Anemone narcissiflora var. nipponica and Solidago virga-aurea ssp. leiocarpa) mixed vegetation, and stem samples of the evergreen shrubs were obtained at 26 months after installing the OTC. The OTC increased the daily mean temperature by 0.1°C to 1.8°C, on average, during the growing season. Shoot developmental growth and biomass production were considerably different between species of different vegetation types. The boreal species E. nigrum generally showed better growth inside the OTC than the arctic and subarctic species L. procumbens. Both species showed significantly larger shoot elongation and biomass production inside the OTC over shrub-dominated vegetation, whereas smaller or reduced growth was detected inside the OTC over shrub-forb mixed vegetation. The variations of growth responses to warming between species of different vegetation types are discussed, especially in relation to interspecific competition under a simulated environmental change.     

常绿阔叶林植物叶片N、P化学计量特征对毛竹扩张的响应

为从生态化学计量内平衡角度解释常绿阔叶林不同层次植物对毛竹(Phyllostachys edulis)扩张的生存响应差异性,该研究采用空间代替时间的方法,在江西井冈山国家级自然保护区沿毛竹扩张方向选取典型毛竹-常绿阔叶林界面,依次设置毛竹林、竹阔混交林和常绿阔叶林样地,比较分析了毛竹扩张方向上样地内不同乔木层、灌木层、草本层植物叶片及土壤N、P含量及比例。结果表明:(1)从毛竹林到阔叶林,土壤N含量上升,P含量下降,N:P上升(P<0.05); 乔木层树种 [红楠(Machilus thunbergii)、赤杨叶(Alniphyllum fortunei)及交让木(Daphniphyllum macropodum)]叶片P含量下降,N:P上升(P<0.05); 除灌木层的红果山胡椒(Lindera erythrocarpa)外,各林分中的灌木层和草本层植物N、P含量及比例变化较小。(2)土壤N:P与乔木层、草本层和灌木层植物叶片N:P分布呈显著正相关、负相关与不相关。(3)在各林分中,毛竹叶片N、P含量及比例较稳定。综上认为,毛竹通过改变土壤N、P化学计量特征进行扩张,引起植物体N、P元素化学计量特征发生变化。灌木及草本植物受土壤异质性影响较小,但是乔木层植物N、P元素化学计量特征却因此失衡,这可能是阔叶林乔木层树种存亡受威胁的重要原因。     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

Differences in Shoot Size and Allometry between Two Evergreen Broad-Leaved Shrubs, Aucuba japonica Varieties in Two Contrasting Snowfall Habitats

Aucuba japonica , an evergreen broad-leaved shrub. Aucuba Japonica var. borealis is widely distributed in heavy snowfall areas in Japan and is covered, shaded and physically pressured by snow for more than four months of the year. On the other hand, var. japonica is widely distributed in light snowfall areas. The sizes of new shoots and leaves were significantly different between the two varieties with different critical shoot sizes for flowering. The average new shoot dry mass of var. borealis was about one third of that of var. japonica. Despite the differences in growing conditions and shoot size, no significant differences were observed in the allometry of their shoot organs between the two varieties. Large new shoots had thicker and longer stems per biomass than small shoots because of their larger pith volume. The large shoots showed higher efficiency of stem growth per invested biomass and had a higher rate of annual height increase than small shoots. When the size of new shoot rapidly increased from year to year, i.e. the plants are growing well, initiation of flowering was postponed and vegetative growth continued. Small new shoots were tolerant of low productivity conditions but traded vertical growth for an increase in matter allocation to leaves. Received 8 July 1999/ Accepted in revised form 1 September 2000     

Two new species and one new variety of Oudemansiella (Agaricales) from Japan

Two new species and one new variety of Oudemansiella (Agaricales) from Japan are described and illustrated: (1) Oudemansiella latilamellata sp. nov. (subgenus Xerula, section Radicatae) has relatively broad lamellae and broadly ellipsoid spores with a subacute apex; (2) Oudemansiella rhodophylla sp. nov. (subgenus Xerula, section Radicatae), growing in deciduous forests of Fagus crenata, Quercus crispula, etc., is characteristic in having lamellae tinted reddish; and (3) Oudemansiella amygdaliformis var. bispora var. nov. (subgenus Xerula, section Albotomentosae) is distinguished from the type variety by having two-spored basidia and grows in evergreen oak forests (Castanopsis sieboldii, Quercus glauca, etc.) or bamboo groves.     

Effects of snow pressure on growth form and life history of tree species in Japanese beech forest

Abstract. In the cool temperate zone in Japan Fagus crenata (beech) is a highly dominating climax species, especially in the snow-rich regions at higher altitudes. The explanation for this dominance was studied with special reference to the tolerance of tree trunks to snow pressure. Traits of six tree species: trunk bend, trunk height decrease, trunk damage and basal sprouting rate were measured in Fagus forest in the Echigo Mountains, central Japan along a snow pressure gradient. The following general trends were recognized: (1) trunk bend and trunk height decrease; (2) snow pressure caused trunk injury; (3) when the trunk bend exceeded a critical angle, the number of sprouts increased remarkably. Although the tolerances were different among the six species, most of them were seriously damaged by snow pressure, and many trunks were reduced in size. On the other hand, many sprouts emerged from deformed trunks and contributed to the maintenance of the population under heavy snow pressure. F. crenata was the only species whose trunk form and maximum size were hardly influenced by snow pressure.     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

Snow cover and extreme winter warming events control flower abundance of some,but not all species in high arctic Svalbard

The High Arctic winter is expected to be altered through ongoing and future climate change. Winter precipitation and snow depth are projected to increase and melt out dates change accordingly. Also, snow cover and depth will play an important role in protecting plant canopy from increasingly more frequent extreme winter warming events. Flower production of many Arctic plants is dependent on melt out timing, since season length determines resource availability for flower preformation. We erected snow fences to increase snow depth and shorten growing season, and counted flowers of six species over 5 years, during which we experienced two extreme winter warming events. Most species were resistant to snow cover increase, but two species reduced flower abundance due to shortened growing seasons. Cassiope tetragona responded strongly with fewer flowers in deep snow regimes during years without extreme events, while Stellaria crassipes responded partly. Snow pack thickness determined whether winter warming events had an effect on flower abundance of some species. Warming events clearly reduced flower abundance in shallow but not in deep snow regimes of Cassiope tetragona, but only marginally for Dryas octopetala. However, the affected species were resilient and individuals did not experience any long term effects. In the case of short or cold summers, a subset of species suffered reduced reproductive success, which may affect future plant composition through possible cascading competition effects. Extreme winter warming events were shown to expose the canopy to cold winter air. The following summer most of the overwintering flower buds could not produce flowers. Thus reproductive success is reduced if this occurs in subsequent years. We conclude that snow depth influences flower abundance by altering season length and by protecting or exposing flower buds to cold winter air, but most species studied are resistant to changes.     

Spatial distribution of canopy and subcanopy species along a sloping topography in a cool-temperate conifer-hardwood forest in the snowy region of Japan

The spatial distributions of canopy and subcanopy species (50cm stem length) were investigated within a plot extending from the top of a ridge to the valley bottom in a cool-temperate old-growth mixed forest, dominated by Cryptomeria japonica and Fagus crenata, in the snowy region of Japan. Based on the longitudinal profile of the slope, the study slope was divided into the relatively gentle upper slope position (US), the steep lower slope position (LS), the flat valley bottom (VB) and the boundary zone between the upper and lower slopes (BS). Spatial dispersal and the association patterns of species were analyzed in upperstory (10cm d.b.h) and understory (<10cm d.b.h) layers. Dominant species in the upperstory layer abruptly changed from Cryptomeria to Fagus at the BS site. In contrast, the understory trees of many species, including shade-intolerant and evergreen species, were independent of the location of conspecific upperstory trees or canopy gaps and extended their distributions on and around the BS site. Significant, diverse canopy and subcanopy species occurred at this site in both upperstory and understory layers. On the BS site, which is the lower margin of Cryptomeria-dominated vegetation, there were many medium-sized C.japonica that were killed by uprooting or breaking of the stems as a result of heavy snow pressure. It is suggested that the snow pressure gradient along a slope has a strong influence on community structure and the maintenance of diverse canopy and subcanopy species in this snowy mixed forest.     

不同微生境下齿肋赤藓(Syntrichia caninervis)生理生化特性对不同季节的响应

齿肋赤藓(Syntrichia caninervis)作为典型的耐旱藓类,在古尔班通古特沙漠的藓类结皮中占优势地位。该沙漠的季节气候差异较大,冬季低温湿润,春季干旱,夏季高温且干旱。荒漠藓类植物叶片仅具单层细胞,对外界环境的变化十分敏感。而有关荒漠藓类植物在生理上如何适应这种剧烈环境变化还不得而知。研究测定了生长于两种不同微生境下的齿肋赤藓,经由低温湿润的冬季到干旱的春季再到高温干旱的夏季过程中生理生化变化特征,以探究不同微生境下齿肋赤藓在水热变化剧烈的不同季节的适应机制。研究发现:季节、微生境及二者的交互作用能够显著影响齿肋赤藓的游离脯氨酸、可溶性糖、可溶性蛋白及丙二醛(MDA)含量、过氧化氢酶(CAT)、过氧化物酶(POD)、超氧化物歧化酶(SOD)活性。夏季的高温干旱使齿肋赤藓的脯氨酸、可溶性糖、MDA含量及3种抗氧化酶活性均显著高于冬季及春季,而可溶性蛋白含量却呈相反趋势。干旱的春季齿肋赤藓脯氨酸及可溶性糖显著高于冬季。而在低温湿润的冬季,齿肋赤藓丙二醛含量及过氧化氢酶活性均显著高于春季。表明夏季齿肋赤藓所受胁迫最大,脯氨酸、可溶性糖含量及抗氧化酶活性大幅度提高。同时,在具有积雪覆盖的冬季,两种微生境下的齿肋赤藓生理生化特性无显著差异;而在无积雪覆盖的春夏季节,灌丛的遮阴作用为齿肋赤藓提供了水含量相对较高的良好生活环境,其下齿肋赤藓的渗透调节物质含量和抗氧化酶活性均显著低于裸露地。表明生长在裸露地的齿肋赤藓较活灌丛拥有更强的胁迫耐受性。     

Differential ecophysiological response of deciduous shrubs and a graminoid to long-term experimental snow reductions and additions in moist acidic tundra, Northern Alaska

Changes in winter precipitation that include both decreases and increases in winter snow are underway across the Arctic. In this study, we used a 14-year experiment that has increased and decreased winter snow in the moist acidic tussock tundra of northern Alaska to understand impacts of variation in winter snow depth on summer leaf-level ecophysiology of two deciduous shrubs and a graminoid species, including: instantaneous rates of leaf gas exchange, and δ¹³C, δ¹⁵N, and nitrogen (N) concentrations of Betula nana, Salix pulchra, and Eriophorum vaginatum. Leaf-level measurements were complemented by measurements of canopy leaf area index (LAI) and depth of thaw. Reductions in snow lowered summer leaf photosynthesis, conductance, and transpiration rates by up to 40 % compared to ambient and deep snow conditions for Eriophorum vaginatum, and reduced Salix pulchra conductance and transpiration by up to 49 %. In contrast, Betula nana exhibited no changes in leaf gas exchange in response to lower or deeper snow. Canopy LAI increased with added snow, while reduced winter snow resulted in lower growing season soil temperatures and reduced thaw depths. Our findings indicate that the spatial and temporal variability of future snow depth will have individualistic consequences for leaf-level C fixation and water flux by tundra species, and that these responses will be manifested over the longer term by changes in canopy traits, depth of thaw, soil C and N processes, and trace gas (CO₂ and H₂O) exchanges between the tundra and the atmosphere.     

Shoot structure and dynamics of saplings and canopies of three deciduous broad-leaved trees of a coppice forest in central Japan

Three deciduous broad-leaved trees, Quercus serrata, Castanea crenata and Carpinus laxiflora, were the main constituents of a coppice forest in central Japan. The shoot elongation and leaf emergence modes of both saplings and the canopy of the three species were investigated. The shoot elongation modes of Q. serrata and C. crenata were the same in saplings and the upper layer of the canopy. The second shoots of these two species were formed after the first shoots were elongated. C. laxiflora was different between saplings and the upper layer of the canopy. In saplings, only the first shoots took a long time to elongate. In the upper canopy layer, higher order shoots were formed in the same way as in the other two species. In the lower layer of the canopy, all three species showed the same shoot elongation mode, in which only the first shoot and its duration of elongation was short. Leaf longevity, individual leaf area, leaf mass per unit leaf area and the stem mass per unit stem length of C. laxiflora were significantly shorter or significantly smaller than those of Q. serrata and C. crenata. The length of the stem per unit leaf area of C. laxiflora was three times that of Q. serrata and five times that of C. crenata. The elongation growth of C. laxiflora was highly efficient as it occurred with a small leaf area. The shoot dynamics and the shoot structure of C. laxiflora are more suitable for elongation growth than in Q. serrata and C. crenata. Furthermore, the shoot structures of the three species were compared and ecological characteristics of the three species are discussed. Received: 29 September 1998 / Accepted: 17 September 1999     

不同厚度雪被对高山森林6种凋落物分解过程中酸溶性和酸不溶性组分的影响

高山森林冬季不同厚度雪被格局可能通过影响凋落物的分解过程中酸溶性和酸不溶性组分特征,改变凋落物分解过程,但缺乏必要关注。采用凋落物分解袋法,研究了高山森林林窗中央至林下形成的天然雪被厚度梯度(厚型雪被、中型雪被、薄型雪被和无雪被)覆盖下,6种典型物种岷江冷杉(Abies faxoniana)、红桦(Betula albo-sinensis)、四川红杉(Larix mastersiana)、方枝柏(Sabina saltuaria)、康定柳(Salix paraplesia)和高山杜鹃(Rhododendron lapponicum)凋落物在不同关键时期(雪被形成期、雪被覆盖期和雪被融化期)的酸溶性组分和酸不溶性组分变化特征。经历一个冬季的分解后,6种凋落物酸溶性组分绝对含量呈降低趋势,除红桦外5种凋落物酸不溶性组分绝对含量呈增加趋势。不同厚度雪被显著影响雪被覆盖期和融化期凋落物酸不溶性和酸溶性组分绝对变化量;其中方枝柏、红桦和康定柳凋落物酸不溶性组分增加量在厚型雪被下显著高于其它雪被覆盖;而相对于阔叶凋落物酸溶性组分变化量在薄型雪被和无雪被梯度达到最大值,针叶凋落物酸溶性组分在厚型雪被下具有最大的变化量。一个冬季分解结束后,表征6种凋落物酸溶性和酸不溶性组分含量相对比例的LCI指数(Lignocellulose index)总体升高,雪被对LCI指数的影响主要表现在雪被覆盖期和融化期,且方枝柏、岷江冷杉和康定柳凋落物LCI在冬季分解后均在厚型雪被达到最高值。同时统计分析结果表明,物种极显著影响冬季不同阶段凋落物酸溶性和酸不溶性组分的变化。这些结果意味着气候变暖情景下,高山森林冬季雪被和冻融格局的改变将显著影响凋落物分解过程中酸溶性、酸不溶性组分以及LCI指数代表的抵抗性组分结构的变化,且影响趋势受到凋落物质量的调控。