西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成与多样性格局

为探讨西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成、多样性格局及其维持机制,该文对两种植被的树干附生苔藓植物进行了调查与分析。所选择的样地位于西双版纳勐腊县,每种植被类型选择10个20 m×20 m的样方,每个样方选择约10棵样树。结果表明:(1)本调查共记录到附生苔藓20科39属60种,其中热带雨林附生苔藓有19科33属48种,热带山地常绿阔叶林附生苔藓有9科14属19种,共有种6种。(2)热带雨林最优势科为平藓科,热带山地常绿阔叶林最优势科为锦藓科。(3)热带雨林比热带山地常绿阔叶林具有更高的物种丰富度、β多样性和γ多样性。(4)不同径级和不同树皮粗糙度宿主上附生苔藓植物的盖度差异显著,而物种丰富度差异不显著。(5)苔藓植物的生活型对植被类型有所偏好,表现为悬垂型、扇型、交织型、粗平铺型集聚于热带雨林,丛集型、垫状型、细平铺型集聚于热带山地常绿阔叶林。(6)直接排序结果表明,宿主特性特别是树皮粗糙度显著影响苔藓植物生活型的组成和分布。热带雨林比热带山地常绿阔叶林能提供更多样的微生境,因而孕育了物种更丰富的附生苔藓植物。考虑到不同的植被类型或宿主特性条件下孕育着不同生活型组成的苔藓植物,因此可将苔藓生活型作为今后森林监测的一项重要指标。     

哀牢山湿性常绿阔叶林树干4种附生苔藓植物的水分变化对光合、荧光参数的影响

以云南哀牢山湿性常绿阔叶林中的附生苔藓植物为对象,研究林内4种树干附生苔藓植物阿萨羽苔(Plagiochila assamica Steph.)、西南树平藓(Homaliodendron montagneanum(Müll.Hal.)M.Fleisch.)、刀叶树平藓(H.scalpellifolium(Mitt.)M.Fleisch.)、大羽藓(Thuidium cymbifolium(Dozy et Molk.)Dozy et Molk.)的吸水力、失水特征和脱水耐性,稳定碳同位素(δ~(13)C)值,以及水分变化对苔藓净光合速率和荧光参数的影响。结果显示:(1)4种附生苔藓植物均具有较强的吸水能力和较低持水力,脱水耐性较强,脱水后均可迅速复水;(2)4种附生苔藓植物的δ~(13)C值较低,其水分利用效率小于其他维管束植物;除大羽藓外,其余3种附生苔藓雨季的δ~(13)C值与水分利用效率均高于干季;(3)4种附生苔藓植物的净光合速率(Pn)随着含水量的降低而降低,在含水量小于60%~80%时,其最大光化学效率(Fv/Fm)急剧下降,反映出这些附生苔藓植物的光合、荧光特性对水分变化非常敏感。     

龙脑香热带雨林附生苔藓沿宿主垂直梯度的微生境偏好及其指示作用

该研究首次借助林冠塔吊调查了西双版纳国家级自然保护区龙脑香热带雨林样地内69棵树13个垂直高度上的附生苔藓植物,结果表明:目标样树上共记录到隶属于25科60属的90种附生苔藓,其中细鳞苔科物种数最多,占比达25.6%。13个垂直高度上共划分出三种生态类型:喜阳苔藓(散生巨树上45 m的区域),喜阴苔藓(乔木树干上15 m的区域),广布苔藓(广泛分布于宿主各个垂直高度上,生态位宽),并筛选出对微生境有特殊偏好的17种苔藓指示种(IndVal≥0.7,P0.05)。随宿主垂直高度的升高,扇型和交织型的苔藓占比降低,悬垂型苔藓占比先升高后降低,细平铺型和粗平铺型的苔藓占比升高。大气湿度、水汽压、胸径以及树皮粗糙度对附生苔藓生活型的分布偏好具有显著影响。总之,沿宿主垂直高度上的附生苔藓对微环境变化在生活型和形态结构上有着不同的响应方式,而同一种生态型的苔藓群落有相似的适应机制。因此,在森林林冠生境变化的监测和管理中,对微生境具有明显偏好的附生苔藓物种可作为有效的指示材料。     

云南树干附生苔藓生物量沿纬度及海拔梯度的变化特征

为探讨不同森林群落树干附生苔藓的生物量及其组成沿纬度、海拔梯度的变化格局,采用样地调查法,对云南热带(勐腊县)、亚热带(镇沅县)及亚高山(丽江玉龙县)不同海拔不同森林类型中树干附生苔藓的总生物量、优势组分生物量和不同生活型组分生物量的分布格局进行调查研究。结果表明:(1)随着纬度、海拔的升高,3种代表性森林类型单位面积树干附生苔藓植物的生物量上升,以丽江亚高山针叶林最高(3.57 g·400 cm~(-2));镇沅亚热带常绿阔叶林次之,为2.00 g·400 cm~(-2);勐腊热带森林最低,仅为0.53 g·400 cm~(-2)。(2)3种森林类型树干附生苔藓植物的优势组分中无相同物种。(3)随着纬度、海拔的上升,单位面积附生苔藓植物平均生物量上升,优势种类的相对生物量占比有所下降,勐腊热带森林、镇沅哀牢山亚热带常绿阔叶林和丽江玉龙雪山亚高山针叶林树干附生苔藓优势组分的生物量分别占总生物量的92.3%、75.6%和85.2%。(4)随着纬度和海拔的上升,3种森林类型树干附生苔藓植物群落不同生活型组分的生物量存在明显的分异规律。细平铺型、扇形和丛集型分别为勐腊热带森林、镇沅哀牢山亚热带常绿阔叶林和丽江玉龙雪山亚高山针叶林的优势附生苔藓生活型。     

边缘效应对山地湿性常绿阔叶林附生植物的影响

作为热带、亚热带地区森林生态系统中重要组成成分之一的附生植物,对环境变化尤为敏感。为了解附生植物对森林生境破碎化的响应,本研究选择哀牢山北段徐家坝山地湿性常绿阔叶林因人为干扰而形成的四处森林边缘为研究对象,分别在距离林缘10、20、40和80m的位置设立样带,采用树干取样法调查各样带内距地面0-2m范围附生植物的物种组成、生物量和附生苔藓植物的生活型特征。结果显示：边缘效应对单位面积附生植物的生物量和附生苔藓植物盖度均具有极显著的影响（P〈0．001）,其中附生苔藓比附生蕨类对边缘效应更为敏感。边缘效应对附生苔藓植物的作用强度在各生活型之间存在着差异,其中对丛集型（P=0．014）和交织型（P=0．030）的附生苔藓植物的影响最显著。综合附生植物在各样带的分布特征,我们认为单位面积上附生植物的生物量和附生苔藓植物的盖度可作为指示森林边缘生境的重要指标。黄牛毛藓（Ditrichum pallidum）、角状刺枝藓（Wijkia hornschuchi）和格氏剪叶苔（Herbertus giraldianus）的分布具有指示林缘生境的意义,而尖叶羽藓（Thuidium philibertii）、大羽藓（zcymbifolium）、多疣麻羽藓（Claopodium pellucinerve）、波叶金枝藓（Pseudotrismegistia undulata）和扭叶藓（Trachypus bicolor）的分布局限于林内,具有一定的保护价值。     

哀牢山山地森林不同附生地衣功能群的水分关系和光合生理特征

附生地衣是热带和亚热带山地森林生态系统中重要的结构性组分, 在生物多样性保护、环境监测、养分循环中发挥着重要作用。附生地衣按共生藻、生活型和繁殖策略的不同可划分为不同的功能群, 不同附生地衣功能群的分布格局存在较大的差异, 然而其生理生态机制仍不清楚。该研究以我国西南地区哀牢山亚热带山地森林中的附生地衣优势类群为研究对象, 对该地区蓝藻地衣、阔叶地衣、狭叶地衣及枝状地衣4种功能群的8种附生地衣的水分关系、光合生理特征等进行了测定分析, 结果显示: 不同功能群附生地衣的持水力和失水速率均存在差异, 其中蓝藻地衣具有较高的最大水分含量, 而枝状地衣的失水速率较快; 过高和过低的水分含量都会抑制附生地衣的光合作用, 但抑制程度有所差异; 蓝藻地衣的光合作用最适水分含量比较高, 表明它们的光合生理活动对水分条件要求较高, 所以它们偏好潮湿的生境, 同时蓝藻地衣的光补偿点比较低但光饱和点却不低, 反映出它们具有较宽的光强适应范围, 所以蓝藻地衣能够同时分布于强光和弱光生境中; 枝状地衣的光合最适水分含量较低, 表明它们的光合生理活动对水分条件要求不是很高, 能够适应较为干旱的环境, 同时枝状地衣的光补偿点和光饱和点都很高, 说明它们的光合生理活动对光照条件要求比较高, 所以它们广泛分布于强光生境中; 阔叶和狭叶地衣的光补偿点比较高, 说明它们更适应有充足光照条件的生境。     

贵州猴耳天坑地下森林苔藓植物多样性特征研究

为了探索苔藓在地下森林中的多样性特征,采用梯度划分法,对贵州猴耳天坑（深度280 m,坑口直径300 m）地下森林进行实地调查研究。结果表明：（1）猴耳天坑地下森林共有苔藓植物71种,包含5种不同生长基质苔藓,其中树附生苔藓物种最为丰富有41种,其余基质苔藓物种数量依次为：土附生>石面生>腐木生>叶附生。（2）地下森林苔藓生活型共计10种,喜阴暗潮湿环境的生活型种类达88%,而适应于强光干旱环境的生活型种类仅12%。（3）对地下森林不同梯度苔藓植物物种丰富度指数分析表明,坡中最高（8.461 7）,依次是坡底（7.502 0）和坡顶（6.978 5）,而苔藓植物Pielou均匀度指数的大小依次为：坡顶（0.945 4）>坡底（0.907 2）>坡中（0.844 5）。（4）β多样性指数分析表明,地下森林不同梯度群落间的相异性都介于（0.75~1.00）之间,底部与顶部的相异系数高达0.960 8。（5）地下森林因其独特的地形地貌和水热条件,不仅各个梯度群落间异质性较高,而且各个群落间相异性也都较高,结构分异明显,生境复杂。（6）RDA分析表明,光照、湿度和温度是影响天坑地下森林苔藓物种分布的主要因子,其中光照度影响最大。（7）坑内与坑外物种多样性的对比研究表明,天坑地下森林苔藓植物物种多样性高,是喀斯特石漠化地区物种的天然避难所。     

长白山暗针叶林苔藓植物生物量的研究

在长白山北坡暗针叶林对地面和树附生苔藓植物的生物量进行了测定.地面生苔藓采取样带调查取样法测定,树附生苔藓应用McCune方法对树干和树枝的附生苔藓生物量都做了细致的测定.结果表明,长白山暗针叶林中的苔藓植物分布很不均匀,随海拔变化差异很大,海拔1100m最低,仅为543kg·hm^-2;海拔1250m最高,达5097kg·hm^-2.苔藓植物生物量的变化对生境有很好的指示作用,特别是塔藓和拟垂枝藓的生物量随海拔的变化与森林系统的群落学特点有一定的相关性:在海拔1100～1700m,塔藓的生物量与臭冷杉的重要值变化趋势相近,随海拔升高而减少;拟垂枝藓的生物量与鱼鳞云杉重要值的变化趋势相似,随海拔升高而增加.此外,生物量随海拔的变化表明了不同苔藓植物对环境条件要求的差异,拟垂枝藓比塔藓水分条件要求更高.因此,生物量的研究在植物生理上也有一定的指示作用.     

喀斯特山地高校苔藓植物多样性及环境响应——以贵州大学为例

以贵州大学喀斯特山地校园为对象,调查其苔藓植物物种多样性及其生境特征,探讨苔藓植物生长与环境的关系。结果表明,研究区域内苔藓植物共12科27属51种。优势科是丛藓科(Pottiaceae)、青藓科(Brachytheciaceae)和灰藓科(Hypnaceae);主要生活型为丛集型、交织型和垫型;不同功能区域的苔藓植物生物多样性从高到低依次为行政办公区、休闲游憩区、教学科研区、体育运动区、宿舍休息区;不同生境物种丰富度从高到低依次是疏林草坪、林下、草坪环境、灌丛花池、道路;人为干扰程度与光照、湿度是影响苔藓植物多样性组成的重要因子。研究区域内苔藓植物多样性较低,多为土生藓类。在苔藓生长的微生境中,人为干扰低、空气湿度大的林地、疏林草坪更利于苔藓生长。     

喀斯特石漠化区典型生境下石生苔藓的固土持水作用

为探究喀斯特石漠化区典型生境下石生苔藓植物的固土持水能力及影响因素,在其石生苔藓分布和形态特征基础上,深入探讨了不同生境下石生苔藓的固土持水效应。结果表明：（1）生境对石生苔藓植物的固土持水能力影响显著。草地中的宽叶真藓（Bryum funkii）固土量最高,为8.85×10³ kg/hm²,裸地中的美灰藓（Eurohypnum leptothallum）持水量最高,其最高持水量是自身干重的14倍。石生苔藓植物在裸地和草地中表现出较高的固土率,而在乔木林与草地下具有较好的持水率。（2）苔藓类型对固土持水能力存在显著性的影响。4种石生苔藓植物的固土率高低为宽叶真藓>尖叶对齿藓原变种（Didymodon constrictus var.constrictus）>卷叶湿地藓（Hyophila involuta）>美灰藓,且差异显著;4种苔藓植物的持水率强弱为美灰藓>宽叶真藓>卷叶湿地藓>尖叶对齿藓原变种。（3）石生苔藓植物的固土持水能力受自身功能性状和立地环境的综合影响,苔藓植物的固土率与干重存在显著的正相关关系,其持水量与干重和生物量间具有显著的正相关关系。综上,从固土持水和生态修复的角度出发,应加强裸地和草地两种生境的石漠化治理力度;可考虑将宽叶真藓和美灰藓作为喀斯特岩面生态恢复的先锋苔藓,其能有效解决喀斯特区石漠化大面积基岩裸露问题,提高喀斯特区水土保持效益。     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.     

Identifying indicator species for bryophyte conservation in fragmented forests

It is important to conserve bryophyte diversity in fragmented forests, due to the vulnerability of this group to environmental change. In this study, the utility of bryophyte functional groups (taxonomic classes, substrate classes, and life-forms) was examined as indicators for planning urban area conservation of bryophyte diversity. The study sites comprised 27 fragmented forests in Kyoto City, Japan. Using linear regression models, it was found that the species richness of almost all functional groups was positively correlated with patch size. Furthermore, two types of bryophyte life-form (fans and thalloid mats) were significantly correlated with environmental factors considered important for conservation planning. The species richness of fan bryophytes was positively correlated with the presence of natural forest patches and was negatively correlated with distance from mountainous areas. Similarly, the species richness of thalloid mat bryophytes was negatively correlated with maintenance practices. These results may be explained by the vulnerability of these two bryophyte groups to environmentally caused drought stress, accompanied by decreasing patch size, maintenance practices, disturbance, and/or loss of natural vegetation. Considering that drought stress represents a major threat to bryophyte diversity in fragmented forests, the species richness of hygrophilous life-forms (e.g., fans and thalloid mats) may be used as an indicator of fragmented forests that are less affected by drought stress, and these species should be preferentially conserved to maintain high levels of bryophyte diversity.     

Bryophytes in perennially moist forests of Papua New Guinea: ecological orientation and predictions of disturbance effects

Water relations of bryophytes must be understood along at least four dimensions: hydration/dehydration frequency; hydration duration; dehydration duration; degree of water loss. All these are biomass-dependent functions such that large colonies of bryophytes may maintain hydration longer than smaller colonies. The spread of a bryophyte colony allows lateral movement of capillary water; contiguous clones will thus allow lateral conduction of that water. In contrast, separated tufts or cushions may store water but will contribute very little to its transfer over the surface of the phorophyte.
Many of the tropical rainforests of New Guinea have a larger mass of epiphytic bryophytes than any temperate forest. The primary effect of disturbance in these forests is a reduction in bryomass, presumably due to desiccation because of increased insolation and wind movement. The reduction of bryomass will decrease water and mineral retention in the bryophyte clones; and the reduction in the size of those clones will reduce their contiguity and thus interfere with bryophyte-mediatcd water and mineral transfer. Disturbance may alter bryophyte species frequency but, under present agricultural and forestry practices in New Guinea, I have seen no evidence of significant loss of bryophyte species.     

影响广东黑石顶树附生苔藓分布的环境因子

通过对广东省黑石顶自然保护区内5个2500 m2样地内树附生苔藓的调查及有关环境因子的测定,研究了树附生苔藓的分布格局及其与环境因子的关系.树附生苔藓在不同高度的分布存在一定梯度,20 cm高处树附生苔藓的种类数与盖度均大于60 cm及更高处,且其群落优势种的数量组成与后者存在较大差异.不同树种附生苔藓盖度和种数差异较大.基于附生苔藓植物的盖度进行DCA排序及聚类分析将树种分成四组,马尾松(Pinus massoniana)因具有两种特有的网藓(巴西网藓Syrrhopodon prolifer和鞘刺网藓S.armatus)单独一组,福建青冈(Quercus chungii)亦与其余各种的差异均较大,形成一组,其余阔叶树种根据其所处的森林类型分成两组,针阔叶混交林内的阔叶树种和次生阔叶林内的阔叶树种各形成一组.对环境因子及树皮含水量和pH的分析显示,垂直梯度上空气湿度的差异可能是造成附生苔藓在不同高度分布差异的主要影响因子之一,不同树种附生苔藓的差异在一定程度上受树皮pH的影响,而与树皮含水量无关.同一树种上树附生苔藓的分布又在一定程度上受森林类型的影响.     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

Indirect effects of hiking trails on the community structure and diversity of trunk-epiphytic bryophytes in an old-growth fir forest

Abstract

Forest hiking trails may influence local microclimate and biodiversity, but the effects on community structure and diversity of epiphytic bryophytes on trees are currently unknown. Epiphytic bryophytes on 82 Abies faxoniana Rehder & Wilson tree trunks (41 along the hiking trail edge and 41 controls in the forest interior) were investigated at four heights from the ground (10, 50, 120, and 180 cm). At each site, air temperature and humidity were monitored for 1 year. The light radiation levels and air temperature were higher, and the canopy leaf area index and air humidity lower at the trail edge, indicating deterioration in microclimate, resulting from the trail establishment. The epiphytic bryophyte species richness, community cover, and mean cover of dendroid and pendent growth forms on trunks were significantly lower at the trail edge than the control site, suggesting that trail construction caused these reductions. One marked effect of the presence of the trail was the increase in some sun-loving species and decrease in shade-tolerant species. Moreover, the trail also slightly influenced species richness and epiphytic bryophyte cover at both community and species population levels along the height gradient. Comprehensive analyses showed that microclimate deterioration was mainly driven by the trail establishment, and that the change in micro-climate along the trail, rather than any host traits, played an important role in the declining epiphytic bryophyte community structure and diversity at the trail edge, confirming the initial hypothesis that the presence of a raised boardwalk (hiking trail) indirectly influences epiphytic bryophyte community and diversity by altering the microclimate.     

紫金山国家森林公园苔藓植物多样性及其与环境关系研究

为深入了解紫金山国家森林公园苔藓植物资源,对南京紫金山展开苔藓植物野外调查,应用典范对应分析(CCA)研究苔藓植物分布与环境间关系。结果表明,调查中共记录苔藓植物23科37属65种,青藓科(Brachytheciaceae)、丛藓科(Pottiaceae)、羽藓科(Thuidiaceae)和绢藓科(Entodontaceae)为优势科(≥6种),绢藓属(Entodon)、青藓属(Brachythecium)、匐灯藓属(Plagiomnium)和真藓属(Bryum)为优势属(≥4种);苔藓植物生境以土生为主,树生较少;苔藓植物生活型以交织型和丛集型为主,占比分别达46.15%和35.38%;苔藓植物区系以温带和东亚成分为主,区系成分呈现明显过渡性;CCA排序结果表明,湿度、坡向和海拔是影响该区域苔藓植物分布的主要环境因子。     

Bryophyte phytomass in tropical ecosystems

Knowledge about bryophyte production and phytomass in the tropics is poor and has been derived principally from studies of epiphytic bryophytes. Such work has been undertaken in Transvaal, Tanzania, Venezuela, Peru and Borneo. Except for the studies in Peru and Borneo, the vegetation types studied and the methods used have been too variable to allow many comparisons or generalizations to be made. However, in general it can be stated that phytomass of epiphytic bryophytes in equatorial latitudes increases from the tropical lowlands to the foreslline. This striking phenomenon has been hypothetically explained by gradients of factors such as precipitation, humidity, temperature and desiccation, or combinations of these. Recent experiments on the gas exchange of tropical-montane bryophytes have revealed that the rate of net assimilation decreases dramatically above 25^oC and that high temperature combined with low light intensities, as realized in the tropical lowland forest, results in high rates of respiration and insufficient net photosynthesis. Experiments with temperate bryophytes show the same results. If can be assumed that tropical lowland species have special physiological adaptations to survive these unfavourable conditions, such as a specialized phytochrome system or effective storage of photosynthetic products. Additional factors could include the relatively high nutrient supply provided by abundant precipitation in tropical montane forests, and damage to cells by strong desiccation in the tropical lowlands.