亚热带常绿阔叶林附生地衣凋落物的物种多样性和生物量的边缘效应

地衣是亚热带山地森林系统附生生物类群的重要组成部分之一,对环境变化极其敏感。为了更好地了解附生地衣对森林边缘效应的响应,我们在云南哀牢山地区原生山地常绿阔叶林中,分别在距林缘5m、25m、50m和 100m处设立样地,收集附生大型地衣的凋落物1年;分析附生地衣凋落物的物种多样性和生物量、功能群特征和组成结构对林缘深度变化的响应特征。研究结果显示,边缘效应能够显著提高林缘附生地衣群落的物种多样性和生物量;其发生的距离最深不超过25m。林缘-林内梯度上,不同地衣功能群的响应模式具有各自的特征。排序分析表明仅在5m样地与其他样地之间存在显著差异,指示种分析则发现仅5m样地具有指示种。哀牢山原生林中边缘效应促进林缘附生地衣生长和分布的现象,可能与当地高湿环境削减了地衣的高光损伤以及以叶状和灌状类群为主的地衣个体受到风力破坏的程度相对较低有关。     

哀牢山湿性常绿阔叶林和次生林木质物残体的组成与碳贮量

研究了哀牢山山地湿性常绿阔叶林和3个主要次生演替类型(栎类萌生林、滇山杨林和旱冬瓜林)木质物残体的组成和碳贮量特征.结果表明:哀牢山徐家坝地区原生森林(常绿阔叶林)木质物残体的碳贮量达36.56t.hm-2,树种组成以腾冲栲、木果石栎和景东石栎等为主,多数是直径≥10cm的大径级、处于中等分解状态的倒木.这主要是由于该区常年潮湿温凉,木质物残体中占优势的壳斗科倒木的材质坚硬,难以分解.3类次生林木质物残体的碳贮量在1.2～5.0t.hm-2,为栎类萌生林>滇山杨林>旱冬瓜林,表现出木质物残体碳贮量随群落演替进程而逐渐增加的趋势.     

Responses of the distribution of four epiphytic cyanolichens to habitat changes in subtropical forests

 蓝藻地衣是附生植物类群的重要组成部分, 在森林生态系统的环境监测和养分循环中发挥着重要作用。该研究在云南哀牢山亚热带森林系统的2种原生和6种次生森林群落中, 以粉缘绵毛衣(Leioderma sorediatum)、天蓝猫耳衣(Leptogium azureum)、网肺衣(Lobaria retigera)和双缘牛皮叶(Sticta duplolimbata) 4种常见蓝藻地衣为对象, 共设立120个样地, 调查了它们在3 600株树木0–2 m树干上的分布, 探讨其分布特征及与森林类群、宿主种类以及林龄等生境因子的关系。研究发现4种蓝藻地衣在森林群落间的分布模式明显不同。除双缘牛皮叶的盖度和频度在原生苔藓矮林中最高外, 其他3种蓝藻地衣的最高值均出现于次生林如厚皮香(Ternstroemia gymnanthera)林和滇山杨(Populus bonatii)林中; 而哀牢山地区广布的原生木果柯(Lithocarpus xylocarpus)林中, 4种蓝藻地衣极为少见。4种地衣都能生长于10多个树种上, 但明显表现出对厚皮香、滇山杨和硬壳柯(Lithocarpus hancei)等树种的偏好性, 以及对小花山茶(Camellia forrestii)等的排斥性。森林群落的林龄、胸径、最大胸径、林冠开阔度、基面积、树木密度和树种多样性等因子的变化均对4种附生蓝藻地衣的分布产生重要影响, 但在景观尺度上影响程度相对较小, 在不同森林群落内部却有各自的重要作用。其中, 林龄、林冠开阔度和宿主胸径是影响蓝藻地衣分布的最重要的生境因子。     

托木尔峰国家级自然保护区树附生地衣分布与环境关系的研究

为了查明新疆托木尔峰国家级自然保护区树附生地衣群落的空间分布规律,在保护区的不同海拔随机设计样点,进行树附生地衣的生态调查,以地衣盖度为指标应用典范对应分析揭示树附生地衣群落物种分布与环境因子之间的关系。结果表明,15个样点中共得出31种树附生地衣,地衣群落结构主要受海拔、湿度、郁闭度和光照强度的强烈影响,并随这些因素的变化表现出一定的空间差异,说明保护区附生地衣的群落变化和空间分布与这些环境因子高度相关。此外,某些地衣可能受全球暖化和气候干旱化的潜在威胁,增加森林盖度和生境湿度是保护地衣资源的有效措施。     

不同水分条件下三种附生地衣的光合作用特性

附生地衣是哀牢山湿性常绿阔叶林生态系统中重要的结构性组分。通过对该区域山地森林中3种典型附生地衣平滑牛皮叶 (Sticta nylanderiana)、网肺衣 (Lobaria retigera) 和橄榄斑叶 (Cetrelia olivetorum)在不同水分条件下的光合光响应及荧光参数的测定分析,结果显示,附生地衣光补偿点 (LCP)、光饱和点 (LSP)较高,对强光适应能力较强。在水分胁迫 (含水量5%~10%) 条件下,3种附生地衣的最大净光合速率 (Pmax) 仅为17~50nmol·g-1·s-1。随着含水量的增加,地衣的最大净光合速率 (Pmax) 与暗呼吸速率 (Rday) 逐渐增大,LCP降低,而LSP随之提高,这表明3种附生地衣具备“阳生植物”的某些特性,从而能够在一定程度上适应野外光照较强的灌丛、向阳林冠等生境。地衣叶绿素光反应中心初始荧光参数 (F0) 和最大光化学效率 (Fv/Fm) 随含水量下降而显著降低,暗示其光反应中心对水分有很强的敏感性。水分条件的改善有助于附生地衣的光反应中心进入到较高的生理活性状态。     

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。     

亚热带森林系统4种附生蓝藻地衣的分布对生境变化的响应

蓝藻地衣是附生植物类群的重要组成部分, 在森林生态系统的环境监测和养分循环中发挥着重要作用。该研究在云南哀牢山亚热带森林系统的2种原生和6种次生森林群落中, 以粉缘绵毛衣(Leioderma sorediatum)、天蓝猫耳衣(Leptogium azureum)、网肺衣(Lobaria retigera)和双缘牛皮叶(Sticta duplolimbata) 4种常见蓝藻地衣为对象, 共设立120个样地, 调查了它们在3600株树木0-2 m树干上的分布, 探讨其分布特征及与森林类群、宿主种类以及林龄等生境因子的关系。研究发现4种蓝藻地衣在森林群落间的分布模式明显不同。除双缘牛皮叶的盖度和频度在原生苔藓矮林中最高外, 其他3种蓝藻地衣的最高值均出现于次生林如厚皮香(Ternstroemia gymnanthera)林和滇山杨(Populus bonatii)林中; 而哀牢山地区广布的原生木果柯(Lithocarpus xylocarpus)林中, 4种蓝藻地衣极为少见。4种地衣都能生长于10多个树种上, 但明显表现出对厚皮香、滇山杨和硬壳柯(Lithocarpus hancei)等树种的偏好性, 以及对小花山茶(Camellia forrestii)等的排斥性。森林群落的林龄、胸径、最大胸径、林冠开阔度、基面积、树木密度和树种多样性等因子的变化均对4种附生蓝藻地衣的分布产生重要影响, 但在景观尺度上影响程度相对较小, 在不同森林群落内部却有各自的重要作用。其中, 林龄、林冠开阔度和宿主胸径是影响蓝藻地衣分布的最重要的生境因子。     

北京东灵山主要森林植被中苔藓植物的物种多样性

通过对北京东灵山地区32个样地植被及苔藓植物的调查,对主要森林植被中苔藓植物物种多样性进行分析。结果表明:东灵山主要森林植被中共有苔藓植物16科33属79种,优势科为青藓科(Brachytheciaceae)和绢藓科(Entodontaceae)等。密叶绢藓(Entodoncompressus)、小牛舌藓(Anomodon minor)等为优势种。地面生苔藓植物物种丰富度以落叶阔叶混交林中最大,白桦林中最小,苔藓植物群落相似性以辽东栎林与落叶阔叶林之间最高,以山杨林与油松林之间最低。树附生苔藓植物物种丰富度以落叶阔叶混交林中最大,油松林中最小;苔藓植物群落相似性以油松林和山杨林之间最高,华北落叶松林与白桦林之间最低。地面生和树附生苔藓植物多样性与林型、树种及其所形成的林内微环境相关。     

哀牢山湿性常绿阔叶林及不同类型次生植被的幼苗更新特征

采用"空间代替时间"的方法,对云南哀牢山徐家坝地区湿性常绿阔叶林及不同类型次生植被中木本植物幼苗的种类组成、数量及更新方式等进行了调查.结果表明:在所有调查样地中,共记录到木本植物幼苗46种,其中乔木31种、灌木15种.荒草地、滇山杨林、栎类萌生林、原生林4种不同植被类型的幼苗密度分别是(421.9±59.5)、(34.4±8.1)、(31.6±7.2)、(18.5±6.6)株·25 m-2.随着演替的进行,灌木的比例逐渐减小,乔木的比例则逐渐增加.在森林恢复演替过程中,处于演替初期的荒草地灌木萌生更新占优势,到了演替后期则以乔木萌生幼苗更新为主.随着群落演替进程的增加,幼苗密度逐渐减少,幼苗生活型和木本植物的幼苗更新方式均发生变化.     

阿尔泰山两河源自然保护区地面生地衣群落物种分布特征

用聚类分析和主成分分析法对阿尔泰山两河源自然保护区地面生地衣群落进行划分;根据优势地衣种类进行群落命名。应用典范对应分析法探讨了地面生地衣种类分布与环境间的关系。结果表明:阿尔泰山两河源自然保护区地面生地衣可分为以下4个群落:白腹地卷+硫石蕊+喇叭粉石蕊群落;茸珊瑚枝+长根地卷+黑瘿地卷群落;土星猫耳衣+软鹿蕊+膜地卷群落;雪岛衣+北方石蕊+绿皮地卷群落。各群落中的地面生地衣种类与盖度,在一定程度上反映了地面生地衣在阿尔泰山两河源保护区分布格局与环境的关系。典范对应分析结果表明,阿尔泰山地面生地衣群落物种分布受到土壤湿度、土壤pH、海拔高度、林冠层郁闭度和灌木层盖度等环境因素的影响,其中海拔、土壤湿度、林冠层郁闭度等对地衣物种分布的影响最大。     

Non-dominant trees significantly enhance species richness of epiphytic lichens in subtropical forests of southwest China

Host species has an important influence on the distribution of epiphytic lichens in forests. However, the importance of non-dominant trees in shaping lichen communities has been poorly studied owing to the relative rarity of individuals. The importance of dominant and non-dominant trees for distribution of epiphytic lichens was determined in eight subtropical forests in southwestern China. Dominant trees supported more abundant total and exclusive lichen species only in secondary forests. The occurrence of non-dominant trees promoted lichen diversity within forest types and influenced lichen communities on both tree groups. The effects of total tree species on lichen distribution largely resulted from the presence of non-dominant trees. Dominant and non-dominant trees supported distinct lichen assemblages within forest types, and ordination analyses showed a clear separation. Our study, therefore, reinforces the importance of non-dominant trees for conserving epiphytic lichens, and highlights that lichen assemblages are shaped by both dominant and non-dominant trees.     

Assessing the effects of forest management on epiphytic lichens in coppiced forests using different indicators

Abstract

Epiphytic lichens are one of the taxonomic groups most sensitive to forest management. Nevertheless, they have not yet been exhaustively included in the assessment of Sustainable Forest Management. This work aimed at evaluating the effects of forest management on epiphytic lichens in coppiced forests, exploring the spatial patterns of diversity and the composition of communities. Moreover, the goal was to compare the performance of four potential indicators for monitoring the effects of forest management on epiphytic lichens: total lichen diversity, species associated with intensive management, species associated with aged coppiced woodlands and Indicator Species Ratio (ISR). In humid Mediterranean Liguria, 50 sampling units were chosen in Castanea sativa and deciduous Quercus spp. forests subjected to different forest management practices: intensively managed coppice and aged coppice/high forest. The effect of forest management was evident in terms of species composition, since it was possible to find significantly associated species for each of the two management types. At each sampling site, the four indicators were calculated using Indicator Value Analysis and compared through correspondence analysis. The ISR was shown to be a more effective indicator, being independent of floristic composition and the occurrence of rare species.     

Bole epiphytic lichens as potential indicators of environmental change in subtropical forest ecosystems in southwest China

Lichen epiphytes are applied as excellent environmental indicators worldwide. However, very little is known about epiphytic lichen communities and their response to forest dynamics in subtropical China. This paper proposes the applications of the cover, diversity, and functional traits of epiphytic lichens to assess environmental changes associated with succession in subtropical forests of southwest China. Bole lichens were sampled from 120 plots of eight representative forest types in the Ailao Mountains. Total cover, species richness, diversity and community structure of bole lichens differed significantly among forest types, and the highest cover and diversity occurred in the Populus bonatii secondary forest (PBSF). Sixty-one indicator species were associated with particular forest types and more than 50% occurred in the PBSF. Both cover and diversity of most lichen functional groups varied regularly during forest succession. Lichen pioneer species were not displaced by competitively superior species as succession proceeds and cyanolichens were more prevalent in secondary forests. The results also highlight the importance of habitat variables such as canopy openness, host diversity, forest age, tree size, the size of the largest tree, tree density, and basal area on the lichen community. Consequently, our findings support the notion that epiphytic lichens, in terms of cover, diversity, species composition and functional traits can be used as effective indicators for large-scale and long-term forest monitoring. More importantly, the narrowly lobed foliose group was the best candidate indicator of environmental conditions in this region. The combined application of lichen indicator species and functional groups seemed to be a more reliable and more powerful method for monitoring forest dynamics in subtropical montane ecosystems.     

Spontaneous restoration of epiphytic lichen biota in managed forests planted on habitats typical for temperate deciduous forest

The study examines the changes of epiphytic lichen diversity in differently aged stands developing after clear cutting and pine planting on fertile habitats typical for deciduous forests. The study was conducted within one large complex consisting of pine, mixed pine-hornbeam and typical old oak-linden-hornbeam forests in northern Poland. Epiphytic lichens were recorded in 50 study plots randomly selected within 5 forest stand classes of a different structure and age, ranging from 80 to over 220 years. Altogether 143 lichen species were recorded, of which only 41 were entirely nonspecific, and were occurring in all the studied forest stand classes. Significant differences in lichen species richness between stand classes were found and the number of species increases with the forest age. Lichen species composition also differs and its changes progress towards restoration of lichen biota typical for deciduous forest consistent with the habitat. The age of the forest has the most significant effect on the biodiversity of lichen biota. Microhabitat space provided by oaks is highly desirable since it greatly enriches lichen biota in forests. Phorophyte specificity of particular lichens were assessed. Hornbeam and oak have the greatest number of species mostly confined to them and constitute a main refuge for lichens with a high conservation value. The changes of lichen biota are basically parallel with the changes of the forest stand structure. The selection of some parts within managed pine forests that should not be assigned for cutting in the future can be a simple procedure which helps to restore and preserve forest biodiversity.     

Species richness and vertical stratification of epiphytic lichens in subtropical primary and secondary forests in southwest China

An entire-tree investigation was conducted in two primary and three secondary forest types in the subtropical Ailao Mountains of southwest China to determine whether species richness and vertical stratification of epiphytic lichens responded to forest type and host attributes. Lichen species number, composition and cover differed significantly among forest types and tree species, while tree diameter and tree height had a modest influence. Epiphytic lichen species and functional groups showed clear vertical stratification. Epiphytic lichens were richer in canopies than on trunks and exhibited a great preference for the intermediate zones of trees, while five lichen groups showed distinct vertical diversification. The stratification patterns are clearly related to forest type and may reflect the microclimatic requirements of individual species, e.g. light availability and humidity.     

A survey method for assessing the richness of epiphytic lichens using growth forms

The species richness of epiphytic lichens is continuously decreasing by degradation and loss of habitat. Considering that taxonomic identification of all species is time and resource consuming, rapid assessment methods to extrapolate the total number of species are needed for practical conservation. This paper describes an alternative method using the correlation between lichens growth forms and species richness. The study was conducted in 406 forest stands located in Central Spain, covering a wide range of mediterranean-climate ecosystem regions, management intensity levels, canopy cover conditions, and tree sizes. The presence/absence of epiphytic lichens was determined in 6090 trees, which were dominated by oak species (Quercus ilex, Q. faginea, and Q. pyrenaica). In all type of forests, the diversity of growth forms was positively correlated with the total epiphytic lichen richness. In all cases, species richness increased in non-managed forest stands with dense canopies. Thus, we propose the use of lichen growth forms as a helpful surrogate of species richness to detect potentially conservation priority areas in the Mediterranean region.     

Removal of secondary compounds increases invertebrate abundance in lichens

We investigated how lichen carbon-based secondary compounds (CBSCs) affect abundance of invertebrates in five lichen species growing on the forest floor (Cladonia rangiferina, Cladonia stellaris) or on tree trunks (Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea). To do this, CBSCs were removed by rinsing lichen thalli in acetone (which has no adverse effects on the lichens) and the lichens were re-transplanted in their natural habitat. After 4 months there was higher abundance of mites, springtails and spiders in the three epiphytic lichens that had their CBSC concentrations reduced. The increase in predatory spiders following CBSC reduction suggests that the compounds have multitrophic consequences. The acetone treatment reduced the number of nematodes in four of the lichen species. Given that lichens serve as important habitats for a diverse range of invertebrates, increased knowledge of how lichen CBSCs may regulate their abundance helps us to better understand the role that lichens and their defence compounds play in structuring forest food webs.     

Patterns and drivers of lichen species composition in a NW-European lowland deciduous woodland complex

We surveyed lichens in an extensive area of lowland deciduous oak and beech dominated forest under wood pasture management within the New Forest National Park, southern England. This provided the basis for an investigation of factors affecting the species density and composition of epiphytic lichen communities. Fifteen 1 hectare plots were established in the same sites as a parallel invertebrate survey, of which nine were in old growth forest and six in oak plantations of between c. 150 and 300 years old. In each plot 12 trees were identified for sampling and lichens were sampled on four aspects of each trunk. Results at the plot level showed that species density was significantly higher in the old growth woodland plots from those in plantations. Plot age had a significant effect on species on oak but this was not significant for lichen communities on beech. The species density of lichens associated with Trentepohliaceae photobionts on oaks showed a linear positive relationship with increasing plot age while the species density of species with other coccal green Chlorophyta decreased with age of the plot. A major part of the lichenised fungi with Trentepohliaceae photobionts includes indicator and notable species used in conservation evaluation and this component is most affected by fragmentation and isolation of forest sites. The results emphasise the importance of long term connectivity and ecological continuity in this extensive mosaic of lowland deciduous forest.     

Forest edge-interior differentiation in the epiphytic lichen diversity of the forest steppe in the Khangai Mountains,Mongolia

Aims The effects of traditional land use by mobile livestock keepers on biodiversity in forest steppe ecotones are insufficiently studied. Epiphytes are an important part of forest plant diversity. Here we analyze differences in the diversity and composition of the epiphytic lichen vegetation between the edge and the interior of Siberian larch forests in the Khangai Mountains, western Mongolia, which are highly subdivided into patches. We asked whether the epiphytic lichen vegetation at the forest edge differs significantly from that in the interior, whether the edge is inhabited by more nitrophilous species than the interior and whether the density of nomad camps around the forest affects epiphytic lichen diversity.Methods Cover percentages of epiphytic lichen species were recorded from 20 trees per plot on 6 plots in the interior and 6 plots at the edge of Larix sibirica forests. The position of nomad summer camps was surveyed using Global Positioning System. Data were analyzed with pairwise significance tests, analysis of similarities, nonmetric multidimensional scaling and canonical correspondence analysis.Important findings The composition of the epiphytic lichen vegetation clearly differed between the two habitats, with more species being more frequent at the edge than in the interior. However, there was no difference in species richness (α-diversity). The epiphyte vegetation at the edge was more uniform and characterized by lower variation of tree-level α-diversity and lower β-diversity than in the interior. At the edge, only nitrophytic lichens were dominant, whereas in the interior, nitrophytes and acidophytes were among the dominant species. This pattern is probably attributable to the spatial heterogeneity of the intensity of forest grazing and was shown to be influenced by the density of nomad summer camps in the vicinity of the forests. Tree-level α-diversity increased with stem diameter, but high-diameter trees were rare. The results suggest that the present level of forest patchiness and the effect of forest grazing increases the diversity of epiphytic lichens on the landscape level, while logging of high-diameter trees reduces lichen diversity.