西南喀斯特山地森林群落土壤种子库研究综述

土壤种子库作为喀斯特山地研究的重要组成部分,其研究成果在深入探讨喀斯特地区植被结构、功能与动态方面具有十分重要而独特的作用.本文系统介绍了该地区种子库的研究方法及主要内容.在对已有成果进行综合分析的基础上,就目前喀斯特土壤种子库研究中亟待解决的一些问题进行了探讨,认为今后应加强森林群落中优势种或建群种土壤种子库动态变化,外来入侵物种土壤种子库动态及其对群落物种多样性的影响机理和土壤种子库在退化植被恢复中的作用与潜力等领域的研究工作,以期为从事西南喀斯特地区土壤种子库研究和该地区植被群落恢复实践提供理论指导.     

哀牢山亚热带常绿阔叶林内外地温分布特征

地温是森林气候重要的环境因素之一,与植物生长密切相关,对地温的深入研究有助于揭示森林生态系统功能和评估森林环境效益。本文利用哀牢山亚热带常绿阔叶林林内和林外旷地(气象站)地温数据,分析发现:哀牢山亚热带常绿阔叶林林内外的地温均呈单峰型日变化,与林外相比,林内地温具有明显日变化的深度较浅(仅在20cm深度以上);林内平均地温具有显著日变化的深度在春季和秋季(20cm)要大于夏季和冬季(15cm);与林外相比,林内各层地温均小于同层的林外地温,并且林内地温的日变幅和年变幅也小于同深度的林外地温。总体来看,林外地温的年变幅和日变幅大于林内。     

秦岭中段不同坡向锐齿栎种子雨、土壤种子库与幼苗更新

以秦岭中段不同坡向(阳坡、半阳坡、半阴坡及阴坡)锐齿栎天然林为对象,对锐齿栎种子雨、种子库的数量、质量动态以及幼苗发育过程进行研究.结果表明: 锐齿栎种子雨一般从8月中旬开始,9月中下旬到10月上旬达到高峰期,11月中上旬结束;不同坡向的种子雨降落历程、发生时间和组成不同;半阳坡种子雨强度最大,为134.13粒·m^-2,其余大小依次为阳坡、半阴坡、阴坡;阳坡种子雨降落时间最早、持续时间最长,高峰期持续时间也最长,阴坡种子雨降落时间最晚、持续时间最短,高峰期持续时间也最短;种子活力及成熟种子占其种子雨的比例大小顺序为半阳坡>阳坡>半阴坡>阴坡.从种子雨降落结束到翌年8月,各坡向土壤种子库总储量大小顺序均为半阳坡>阳坡>半阴坡>阴坡;各坡向的土壤种子库中成熟、未成熟、被啃食种子数量和种子活力均随时间变化呈递减趋势,而霉烂种子数量增加;各坡向土壤种子库中的种子均主要集中在枯落物层,其次为0~2 cm土层,而在2~5 cm土层中只有极少量种子存在.4种坡向中锐齿栎幼苗的密度差异显著,半阳坡幼苗最多,其余依次是阳坡、半阴坡和阴坡.半阳坡更适合锐齿栎种子的萌发和幼苗生长.     

Patterns of plant sexual systems in subtropical evergreen broad-leaved forests in Ailao Mountains,SW China

Aims Sexual systems influence many components of the ecology and evolution of plant populations and have rarely been documented in subtropical evergreen broad-leaved forests (SEBLF). Here we report frequency distribution and ecological correlates of plant sexual systems in SEBLF at Ailao Mountains, and compare our results with that of tropical and cool temperate forests.Methods We examine the sexual systems of 703 species of woody angiosperms belonging to 103 families and 296 genera based on a comprehensive survey of SEBLF at Ailao Mountains Natural Reserve. Information of plant sexual systems and ecological traits were mainly based on published literatures and specimens as well as our field observations. The associations between plant sexual system and ecological traits were assessed with chi-square tests.Important findings Among these species, 60.2% were hermaphroditic, 15.8% were monoecious and 24.0% were dioecious. The percentage of dioecious sexual system among tree species (22.2%) in SEBLF was comparable to those of tropical tree floras, but much higher than those of temperate forests at North America. The percentage of monoecious sexual system among tree species (30.1%) in SEBLF was higher than that of tropical tree floras, but much lower than those of temperate forests at North America. Monoecy was significantly associated with the tree growth form and was relatively common in temperate genera. Dioecy was significantly associated with fleshy fruits and monoecy was significantly associated with dry fruit type in SEBLF. The high percentage of diclinous sexual systems (monoecy and dioecy) in SEBLF may be associated with the origin of the flora and the prevalence of relatively small inconspicuous flowers of the forests in the Oriental Region.     

哀牢山亚热带常绿阔叶林土壤含水量变化规律及其影响因子

利用ChinaFlux设置在哀牢山亚热带常绿阔叶林内的通量观测系统的土壤含水量数据,分析了哀牢山亚热带常绿阔叶林内土壤含水量的时间变化及其影响因子,以期了解哀牢山亚热带常绿阔叶林土壤含水量特征及其变化规律,为森林固碳能力、潜力和速率研究提供支撑.结果表明:(1)哀牢山亚热带常绿阔叶林不同深度土壤含水量大致呈单峰曲线分布,各层土壤含水量基本上随着土壤深度的增加而降低,波动范围也逐渐减小,最小值均在3月,最大值集中在7、8、9三个月.(2)0 ～ 100 cm各层深度的土壤含水量在17.7％～39.5％波动,其中100 cm深度的土壤含水量值最低、波动范围最小,总体来看,土壤含水量雨季高于旱季.(3)土壤含水量主要受降雨量、地温和相对湿度等要素的影响(P＜0.05),哀牢山亚热带常绿阔叶林5 cm深度的土壤含水量最大,约为38％.     

哀牢山亚热带常绿阔叶林土壤温度时空分布对模拟增温的响应

为把握森林不同深度的土壤温度对区域气候变暖的响应,评估气候变暖对亚热带森林土壤呼吸的影响,利用在哀牢山亚热带常绿阔叶林中设置的土壤增温和土壤呼吸人工控制实验,对2011—2013年的对照样地和增温样地不同深度的土壤温度实测数据进行了分析,结果表明:区域气候变暖导致的温度升高不改变土壤温度的年变化和日变化特征;干季的增温效应大于雨季;同一深度增温效应具有一定的年变化,但日变化不明显;增温效应在土壤表层较大,1—4月明显,2月最高,增温可达3℃;5 cm深度的平均增温效应在2℃左右,且年变化较小;随着深度的增加,增温效应呈现对数降低趋势;如果以0.5℃为增温效应的阈值,推算可得出:干季的影响深度为3.82 m,雨季可达12.04 m,年均为6.58 m。     

围封沙质草甸土壤种子库与幼苗库的特征

采用野外定点取样和萌发实验相结合的方法,研究了科尔沁沙地围封沙质草甸土壤种子库和幼苗库的特征.结果表明:土壤种子库总密度为13854±1651有效种子数/m2,而幼苗库的总密度为808±164株/m2,仅为土壤种子库总密度的6%,土壤种子库与幼苗库总密度之间存在显著的差异 F=36.0,n=219,P<0.001 .土壤种子库与幼苗库的植物种数分别为37和44,其中一、二年生草本植物分别占种子库与幼苗库植物种数的62%和50%,而多年生草本植物所占的比例分别为24%和41%,灌木所占的比例很小,分别为14%和9%.土壤种子库与幼苗库种类组成上的相似性系数为0.667,表现出较小的异质性.土壤种子库密度与幼苗库密度之间存在显著的相关性,其间关系可用三次曲线来描述.     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

川西亚高山65年人工云杉林种子雨、种子库和幼苗定居研究

川西亚高山人工针叶林已成为亚高山森林的重要组成部分, 它们是否具有持续的自然更新能力, 是决定川西亚高山针叶林群落演替方向和维持该区针叶林大面积存在的基础。以川西米亚罗亚高山人工云杉(Picea asperata)林(65 a)为研究对象, 对种子雨量年际变化、土壤种子库动态、种子萌发和幼苗定居等更新过程的关键环节进行了连续7年(2002-2008年)的野外观测, 以研究人工云杉林更新潜力及影响其更新的限制因素。结果表明: 该区云杉林种子雨一般从每年的10月初开始下落, 一直到翌年的1月底或2月初结束; 云杉种子散落存在明显的大小年现象, 种子散落周期为4年, 且大小年之间种子产量差异极大; 云杉种子从下落到土壤到种子完全失去活力不到1年时间, 属于Thompson和Grime定义的第II类土壤种子库类型。腐烂死亡和动物取食是土壤种子库损耗的主要因素, 而种子通过萌发真正转化为幼苗的比例非常低, 仅占2002年下落种子总量的3.6%。种子萌发后, 环境筛的作用导致云杉幼苗大量死亡, 尤其是在种子萌发后的一个生长季节内, 其死亡率高达78%。凋落物和苔藓是构成人工云杉林下地表的两种主要地被物类型, 二者占所有调查幼苗数量的93%左右; 两种地被物类型上0-2 cm层幼苗存活率最高, 分别占存活幼苗总数的76.07%和86.72%, 随地被物厚度增加, 幼苗存活率呈明显下降趋势, 而幼苗死亡率呈明显升高的趋势, 表明林下地被物厚度也是影响云杉幼苗定居的重要因素。两种地被物对幼苗生长的影响不同, 除株高之外, 分布在苔藓上的云杉幼苗生长参数(地径、分枝数、干重以及干重年增长率)明显高于分布于凋落物上的幼苗, 表明苔藓地被物更有利于云杉幼苗定居。尽管该区大量云杉种子下落, 但由于种子的高损耗率、幼苗的低输出率以及萌发幼苗的高死亡率, 使得人工云杉林下种子通过萌发转为实生幼苗的数量非常少, 最终真正能补充到云杉种群的个体数量非常有限。     

The soil seed bank and understorey regeneration in Eucalyptus regnans forest,Victoria

Abstract The soil seed bank and its relation to the extant vegetation in a Eucalyptus regnans F. Muell. forest in the Central Highlands of Victoria were examined. The average seed density was 430 germinable seeds m^?2 to a depth of 2 cm. There was a polynomial regression relationship between the density and species richness of seeds in soil and forest age (0. 6–54 years). Species richness was not significantly different among soil depths (0- 2 , 2- 5 , 5–10 and 10–20 cm) in the forest stand of 54 years old. More seeds germinated from the 5–10 cm depth than from the other depths. Forbs accounted for 73% of the total germinable seeds and there was no germination of E. regnans. The number of species, particularly woody plant species, germinating from the soil seed bank were significantly lower than in the extant vegetation. However, almost all species present in the soil seed bank were present in the vegetation. The soil seed bank provides an important source for the rapid regeneration of understorey vegetation following clear-cutting and slash-burning in the E. regnans forest. The rapid understorey establishment may play an important role in protecting soil from erosion, in nutrient conservation, replacement and redistribution. The soil seed bank may also be a necessary source of maintaining genetic diversity in the forest over the long term.     

Similarities in the impact of three large invasive plant species on soil seed bank communities

Invasions by alien plant species may substantially alter soil seed bank communities. While decreases in seed bank species richness, diversity, and composition as a consequence of plant invasions have been reported, the characteristics of seed banks associated with different invasive species have not been compared in any detail. Here, we describe changes in the characteristics of soil seed banks invaded by three large herbaceous invasive plants, Fallopia japonica, Gunnera tinctoria, and Heracleum mantegazzianum. The study was carried out at the spatial scales of site and plot, to reduce variability in seed bank data. Information on seed bank persistence was inferred from seed depth (0–5, 5–10, and 10–15 cm) and from time of sampling (May and October). Despite differences in the reproductive strategy and geographic distribution of these invaders, as well as in the standing vegetation and habitat types examined, the seed banks of invaded areas were similar in composition and in the relative abundance of different species. Invaded seed banks were dominated by seeds of a few agricultural weed species and/or rushes, suggesting that common features of the invaders, including a large standing biomass, extensive litter production, and the formation of mono-species stands may result in comparable selection pressures that favors traits that are largely genera or species-specific. These findings have a direct relevance for the development of strategies aimed at restoring previously-invaded sites while also improving our understanding of the long-term implications of plant invasions.     

哀牢山不同类型亚热带森林地表凋落物及土壤节肢动物群落特征

分别于2005年4月（干热季）、6月（雨季）和12月（干热季）,采用样线法对哀牢山中山湿性常绿阔叶林、山顶苔藓矮林和滇山杨次生林地表凋落物及其中的土壤节肢动物群落进行了调查.结果表明： 凋落物现存总量、C储量和C/N在干、湿季均表现为苔藓矮林＞滇山杨林＞常绿阔叶林,N储量差异不大.蜱螨目和弹尾目是3林地凋落物层节肢动物群落的优势类群,双翅目幼虫、鞘翅目、蚁类和同翅目为常见类群,3林地凋落物层土壤节肢动物群落相似性系数极高.3林地凋落物层土壤节肢动物群落密度(ind·m^-2)没有显著差异,但相对密度(ind·g^-1)呈现出常绿阔叶林和滇山杨林显著高于苔藓矮林;土壤节肢动物密度季节变化呈现干季（4月和12月）显著高于雨季（6月）,群落香农多样性指数差异不显著.干热季(4月)3林地枯枝落叶现存量与其土壤节肢动物群落和主要类群密度呈显著正相关关系,而在干冷季(12月)凋落物总现存量与其土壤节肢动物群落及蜱螨目相对密度则呈显著负相关关系;弹尾目和鞘翅目密度与地表凋落物层的N储量也有显著的正相关关系.哀牢山亚热带森林凋落物及其土壤节肢动物群落的发展与森林植被结构密切相关, 其群落个体数量和多样性受森林地表凋落物的调控, 但林内其他环境因素如温湿度对森林凋落物层土壤动物个体数的季节变动也有明显影响.     

Characteristics of floor litter and soil arthropod community in different types ot subtropical forest in Ailao Mountain of Yunnan, Southwest China

By using line transect method, an investigation was conducted on the floor litter and soil arthropod community in a mid mountain wet evergreen broad-leaved forest, a mossy dwarf forest, and a Populus bonatii forest in Ailao Mountain of Yunnan in April (dry and hot season), June (rainy season), and December (dry and cold season), 2005. In both dry and rainy seasons, the existing floor litter mass, C storage, and C/N ratio in the three forests all increased in the order of mossy dwarf forest > P. bonatii forest > evergreen broad-leaved forest, but the N storage had less difference. In the floor litter layer of the forests, Acari and Collembola were the dominant groups of soil arthropod community, while Diptera larvae, Coleoptera, ants, and Homoptera were the common groups. The Sorenson coefficients of soil arthropod community in the three forests were extremely great. No significant differences were observed in the soil arthropod density (ind x m(-2)) in the floor litter layer among the three forests, but the relative density (ind x g(-1)) of soil arthropods was higher in the evergreen broad-leaved forest and P. bonatii forest than in the mossy dwarf forest. In the three forests, the density of soil arthropods was significantly higher in dry season than in rainy season, but the Shannon diversity index had less difference. There were significant positive correlations between the existing floor litter mass and the individual density (ind x m(-2)) and dominant groups of soil arthropod communities in dry and hot season (April), but negative correlations between the existing floor litter mass and the relative density (ind x g(-1)) of soil arthropod communities and Acari in dry and cold season (December). The individual densities of Collembola and Coleoptera also had positive correlations with the N storage of the existing floor litter mass in the three forests. It was considered that the floor litter and the development of soil arthropod community in the litter layer of the subtropical forests in Ailao Mountain had a close relation with the vegetation structure of the forests, and the individual density and the diversity of the soil arthropod community were controlled by the floor litter, whereas the environmental factors such as temperature and moisture in the forests also had obvious effects on the seasonal dynamics of the individual density of the soil arthropods.     

Woody debris stocks in different secondary and primary forests in the subtropical Ailao Mountains, southwest China

Woody debris (WD), including coarse woody debris (CWD) and fine woody debris (FWD), is an essential structural and functional component of many ecosystems, particularly in montane forests. CWD is considered to be the major part in forest WD and it is primarily composed of logs, snags, stumps and large branches, while FWD mainly consists of small twigs. Attributes of dead woody material may change in accordance with trends in stand dynamics. The primary forest (primary montane moist evergreen broad-leaved forest) in Ailao Mountain National Nature Reserve (NNR) preserves the largest tract of natural vegetation in China. The Alnus nepalensis (D. Don) association, Populus bonatii (Levl.) association and secondary Lithocarpus association represent the secondary and chronological types following human disturbance by fires and logging under different intensity. The mass and composition of coarse woody debris (CWD, ≥10 cm in diameter) and fine woody debris (FWD, 2.5–10 cm in diameter) were inventoried in a primary forest and its three secondary counterparts. Estimates of total mass of woody debris across secondary types to primary forest ranged from 2.4 to 74.9 Mg ha⁻¹. The lowest value was found in the A. nepalensis association and the highest values were in the primary forest of which logs are the considerable differences. The ratios of CWD to FWD were low in the secondary types (about 1–4) but high in the primary forest (above 15). Our results suggested that for the recovery of woody debris in the secondary forest, it might last longer than the age of the oldest successional stage studied. Yang Lipan and Ma Wenzhang contributed equally to this work.     

哀牢山中山湿性常绿阔叶林土壤氮转化的海拔效应

采用树脂芯法将哀牢山中山湿性常绿阔叶林内土壤分别移植到中海拔的次生林和低海拔的人工林下培养,并以原地培养为对照,对土壤氮素转化的海拔效应进行了研究.土壤氮素的净矿化速率、净硝化速率和淋溶速率受季节和海拔的影响极为显著(P＜0.01).海拔的影响在雨季前期最显著,高海拔土壤的净矿化和净硝化速率分别为-5.81和-4.18mg N·kg-1·60d-1,移植到中、低海拔培养后,净矿化速率分别为20.92和44.15 mgN·kg-1· 60 d-1,净硝化速率分别为17.07和20.38 mgN· kg-1 ·60d-1,淋溶量也分别增加了0.37倍和2.77倍.由于雨季中后期反硝化作用增加导致净矿化和净硝化速率降低,导致高、中海拔培养的土壤净氮矿化速率在雨季中期达到最高值,雨季后期降低.由此可推断,未来的气候变暖很可能会加快哀牢山中山湿性常绿阔叶林土壤氮素的转化速率和气态损失量.     

Estimation of epiphytic biomass and nutrient pools in the subtropical montane cloud forest in the Ailao Mountains,south-western China

Canopy organic matter (COM) composed of epiphyte community exists as a complex subsystem in the montane cloud forest (MCF). We estimated the biomass and nutrient pools of COM in a subtropical MCF in the Ailao Mountain National Nature Reserve (NNR). The diameter at breast height (DBH) of host trees was the best parameter in the estimation of epiphytic biomass, compared to the height (H) of host trees and the combination of both parameters. The effect of host tree species was negligible in the estimation. Therefore, it was concluded that the DBH data of all trees of a forest regardless of species was a suitable index for the non-destructive estimation of epiphytic biomass in field surveys, especially for investigations of larger study areas. According to species-specific equations, the total COM biomass was estimated to be 2,261 ± 537 (SD) kg ha⁻¹, which was dominated by bryophytes (73.6%) and canopy humus (13.9%). The greatest proportion of COM was located on the inner branches (52.2%), followed by outer branches, trunks, the understory, and branch junctions. The nutrient pools (mean ± SD, kg ha⁻¹) of the COM were: N = 37.9 ± 9.0, P = 1.97 ± 0.47, K = 9.6 ± 2.3, Ca = 9.6 ± 2.3, Mg = 2.64 ± 0.63 and Na = 0.25 ± 0.06. Assessments of epiphytic biomass and nutrient capital supported the idea that the canopy subsystem holds a substantial pool of nutrients, especially when compared to the labile components of the forest ecosystem.     

Forest plant communities of the Baohua Mountains,eastern China

Abstract. In the forest of the Baohua Reserve in the Baohua Mountains, eastern China, six plant communities and 11 sub‐communities were described using the Braun‐Blanquet method. These communities can be assigned to four already described Chinese vegetation types. Curtis’ Compositional Index (CI) and Shannon's diversity index were used to determine the successional stage and species diversity of the communities. The community of Brachystachyum densiflorum and Pinus massoniana and the subcommunity of Fraxinus chinensis and Quercus acutissima with Pueraria lobata had the lowest CI and would be an initial successional stage; the community of Phoebe sheareri and Cyclobalanopsis glauca had the highest CI and would be the final stage. The community of Brachystachyum densiflorum and Pinus massoniana and the sub‐community of Fraxinus chinensis and Quercus acutissima with Pueraria lobata, both pioneer successional stages, had the lowest species diversity. The climax communities had the highest diversity.     

Restoration of ditch bank plant species richness: The potential of the soil seed bank

Abstract. Small‐scale landscape elements, such as ditch banks, play an important role in preserving plant species richness in agricultural landscapes. In this study, we investigate whether the seed bank might be useful for restoring the above‐ground plant species richness. We studied the vegetation and seed bank composition at six species‐rich and six species‐poor ditch banks, where agri‐environment schemes are running to maintain and enhance ditch bank plant diversity. We show that the number of species in the seed bank was low, regardless of the number of species in the established vegetation. Moreover, the seed bank was always dissimilar to the established vegetation. Target species for nature conservation were occasionally present in the seed bank at both species‐poor and species‐rich sites, but rarely so if the species was absent from the established vegetation. We conclude that the potential use of the seed bank for restoration of ditch banks is minimal. At present, plant species richness seems to be largely controlled by germination opportunities; high biomass and competition appear to hamper germination at species‐poor sites. We recommend continued nutrient reduction at such sites. Soil disturbance measures and deliberate sowing should also be considered.     

贡嘎山不同林龄峨眉冷杉种子雨及土壤种子库

本文采用种子雨收集器、土壤种子库筛选、室内萌发实验及野外实地调查等手段,研究了贡嘎山海螺沟内不同林龄和海拔分布上下界限的6个峨眉冷杉(Abies fabri)林的土壤种子库、种子雨的组成和时空分布特征.结果表明:土壤种子库分布随土壤深度增加而减少,主要集中在凋落物层;在同一海拔,随着林龄增加,土壤种子库和种子雨大小先增加再降低,成熟林达到最大值,且其完整种子比例最高,整体质量最好;不同海拔,林龄相近的成熟林调查结果显示,海拔分布上下界限的峨眉冷杉林土壤种子库和种子雨显著小于海拔分布中段的成熟林,这可能与海拔引起的水热条件差异有关;峨眉冷杉林的最佳结实阶段在成熟林,其种子质量好,且散布后在林下保存得最好;树种生物学特性和林下微生境可能是主要原因.