Biological collections in an ever changing world: Herbaria as tools for biogeographical and environmental studies

Plant specimens stored in herbaria are being used as never before to document the impacts of global change on humans and nature. However, published statistics on the use of biological collections are rare, and ecologists lack quantitative data demonstrating the relevance to science of herbarium specimens. I found 382 studies with original data that used herbarium specimens to document biogeographical patterns or environmental changes. Most studies are less than 10 years old, and only 1.4% of the herbarium specimens worldwide have been used to answer biogeographical or environmental questions. The vast majority (82%) of papers dealt with vascular plants, but some studies also used bryophytes, lichens, seaweeds and fungi. The herbarium specimens were collected from all continents, but most of the studies used specimens from North America (40% of studies) or Europe (28%). Many types of researches (conservation, plant disease, plant invasion, pollution, etc.) can be conducted using herbarium specimens. Climate change, and especially phenological reconstructions, are clearly emerging research topics. By group, small herbaria (<100,000 specimens) are consulted as often as very large herbaria (>1,000,000 specimens) for biogeographical and environmental research, but in most cases, only large facilities provide specimens collected worldwide. The median number of specimens per study in papers using computerized collections (15,295) was much higher than for papers that did not include electronic data (226). The use of molecular analyses to investigate herbarium specimens is still relatively unexplored, at least from biogeographical and environmental points of view. Combined with recently developed procedures to correct biases, herbarium specimens might provide in the near future exciting additional spatio-temporal insights that are currently unimaginable.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

Arctic warming on two continents has consistent negative effects on lichen diversity and mixed effects on bryophyte diversity

Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well‐known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long‐term (9–16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N₂‐fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.     

Phylogeny and taxonomy of the ‘manna lichens’

We present ataxonomic revision of the ‘manna lichens’ based on morphological, chemical, ecological and molecular data. A large number of herbarium specimens and fresh collections were examined. Phylogenetic analyses were performed using nuclear ribosomal (nrITS, nrLSU) and mitochondrial small subunit (mtSSU) sequences. Some notable phenotypic characters were plotted on the phylogenetic tree, and the analysis reveals that some of these characters are useful for genus and species level distinction of certain ‘manna lichens.’ Phylogeny of the Megasporaceae was revised using a combined data set of nrLSU and mtSSU and performing parsimony and Bayesian analyses. Five genera (Aspicilia, Circinaria, Lobothallia, Megaspora and Sagedia) are recognized. Further, the relationships of five presumably closely related genera of ‘manna lichens’, namely Agrestia (vagrant), Aspicilia (crustose) Circinaria (crustose), Chlorangium (vagrant) and Sphaerothallia (vagrant) with different growth forms were analysed. The analyses revealed that ‘manna lichens’ do not form a monophyletic group but occur in different clades within the genus Circinaria. The genera Agrestia, Chlorangium and Sphaerothallia are assigned as new synonyms under the genus Circinaria and no vagrant or erratic species remain in the genus Aspicilia. The analyses also show that five new erratic, vagrant and crustose species can be recognized. In this study two ‘manna lichens’, viz. Circinaria rostamii sp. nov. (Azerbaijan, Iran and Turkey), and Circinaria gyrosa sp. nov. (Armenia, Azerbaijan, Iran, Turkey, Turkmenistan and Spain) are described as new to science. Three potentially new species with crustose and erratic forms need additional study. Aspicilia fruticolosofoliacea is reduced to synonymy under C. alpicola, and a lectotype is designated for C. aspera. Thirteen new combinations in Circinaria are presented. The phenomenon of vagrancy is briefly discussed, and the biogeography of the ‘manna lichens’ is outlined. Illustrations, distribution maps, and an identification key to the species are provided.     

中国云南西部地衣区系(英文)

对中国云南西部高山地区的大型地衣进行了调查。采集地衣标本400余份,鉴定为46属86种,其中40种在朝鲜半岛有报道。所有采集的标本现保藏于顺天大学韩国地衣研究所和中国科学院昆明植物研究所隐花植物标本馆。     

河南省湿地藓属的初步研究

在研究标本的基础上,根据配子体和孢子体的外部形态及其解剖结构和文献中对中国湿地藓属(Hyophila)系统位置的主要观点,鉴定出河南省分布有该属植物6种,其中花状湿地藓(H.nymaniana)、四川湿地藓(H.setschwanica)、匙叶湿地藓(H.spathulata)3种为河南省藓类新记录.编制了河南省湿地藓属植物分种检索表,对每个种的生境、分布及识别特征进行了概述.并对湿地藓属的系统演化位置以及该属植物在河南省的分布进行了讨论.     

Tree hollows can affect epiphyte species composition

Tree hollows often harbor animals and microorganisms, thereby storing nutritive resources derived from their biological activities. The outflows from tree hollows can create unique microenvironments, which may affect communities of epiphytic organisms on trunk surfaces below the hollows. In this study, we tested whether the species richness and composition of epiphytic bryophytes (liverworts and mosses) and lichens differ above and below tree hollows of Aria japonica and Cercidiphyllum japonicum in a Japanese temperate forest. The species richness of epiphytic bryophytes and lichens did not differ above and below hollows; however, the species composition of bryophytes differed significantly above and below hollows. Indicator species analyses showed that the moss species Anomodon tristis and the liverwort species Porella vernicosa were significantly more common below than above hollows, while the liverwort species Radula japonica and four lichen species, including Leptogium cyanescens, occurred more frequently above than below hollows. Our results highlight that tree hollows can produce unique microenvironments on trunk surfaces that potentially contribute to the maintenance of epiphytic diversity on a local scale.     

Spatial structure, rock type, and local environmental conditions drive moss and lichen distribution on calcareous boulders

Although mosses and lichens are a relevant component of the biota of rock habitats targeted for biodiversity conservation in Europe, the ecological factors driving their distribution are still poorly known. In this work, we examined the epilithic moss and lichen assemblages colonizing boulders of different types of calcareous rocks co-occurring in the same area in the Italian Alps. The goals were: (1) to evaluate if and to what extent different calcareous rocks host different assemblages; (2) to identify species associated to each rock type; (3) to quantify the relative importance of rock type, local environmental factors, and habitat spatial structure in explaining species distribution. Our results demonstrated that different calcareous rocks host different moss and lichen assemblages with some typical species, indicating that each rock type contributes to the total diversity of both mosses and lichens. Local environmental conditions influenced mosses and not lichens whose distribution is mainly associated to rock type. The patterns of both organism groups were also significantly related to habitat spatial structure, species assemblages tending to have a patchy distribution, which may reflect dispersal dynamics. Our results have implications for conservation: (1) each rock type may play a relevant role in maintaining the overall diversity contributing with unique assemblages and typical species; (2) the patchy distribution of both moss and lichen assemblages should warn from considering rock patches as a monotonous repetition of the same habitat across space.     

贵州乌江东风水库库区消落带苔藓植物区系分析

采用典型调查与路线调查相结合的方法,对贵州乌江东风水库库区消落带苔藓植物区系的物种组成、生活型、分布区类型和丰富性进行了调查和分析.结果表明:该区域共有苔藓植物18科58属101种,其中,藓类植物有16科56属99种,苔类植物有2科2属2种;优势科为灰藓科(Hypnaceae)、青藓科(Brachytheciaceae)和丛藓科(Pottiaceae),优势属为青藓属(Brahchythecium B.S.G.)、真藓属(Bryum Hedw.)和小曲尾藓属〔Dicranella(Müll.Hal.)Schimp.〕;单属科和单种属所占比例均较高,分别占该区域苔藓植物总科数和总属数的500%和638%.该区域分布的苔藓植物生活型可分为交织型、丛集型、垫状和平铺型4类,以丛集型种数最多(48种),占该区域苔藓植物总种数的475%.该区域分布的苔藓植物可划分为12个分布区类型,其中,温带成分种类最多,所占比例为815%;热带成分所占比例仅为185%;中国特有种所占比例也较高,为207%.该区域苔藓植物的丰富性综合系数(Si)为-03608,低于相邻的六冲河下游流域.综合分析结果表明:该区域的苔藓植物多样性较为丰富,多数种类具有较强的抗逆性,且丛集型苔藓种类最多,与库区消落带的特殊生境相适应;地理成分以东亚成分和北温带成分为主,总体属温带性质,且中国特有种较多,反映出该区域苔藓植物区系的特殊性和复杂性.     

Distribution and floristics of moss- and lichen-dominated soil crusts in a patterned Callitris glaucophylla woodland in eastern Australia

The distribution and abundance of soil crust lichens and bryophytes was examined in a patterned Callitris glaucophylla woodland in eastern Australia. Twenty-one lichen species and 26 bryophyte species were collected within thirty quadrats along a sequence of runoff, interception and runoff zones. Crust cover was significantly greatest in the interception zones (79.0 %), followed by the runoff zones (24.0 %), and lowest in the groved, runon zones (6.6 %). Lichens and bryophytes were distributed across all geomorphic zones, and, although there were significantly more moss species in the interception zones (mean = 9.1) compared with either the runoff (4.2) or runon (3.2) zones, the number of lichen species did not vary between zones. Ordination of a reduced data set of 32 species revealed a separation of taxa into distinct groups corresponding to the three geomorphic zones. Canonical correspondence analysis (CCA) of the 32 species and thirteen environmental variables revealed that the most important factors associated with the distribution of species were sheet and scarp erosion, soil stability and coherence, litter cover and crust cover. Surface cracking, microtopography and plant cover were of intermediate importance. The CCA biplot revealed that the timbered runon zones (groves) were dominated by `shade-tolerant' mosses Fissidens vittatus and Barbula hornschuchiana, whilst the heavily eroded runoff zones supported sparse populations of `erosion tolerant' lichens (Endocarpon rogersii) and mosses (Bryum argenteum and Didymodon torquatus). Interception zones supported a rich suite of `crust forming' mosses and lichens capable of tolerating moderate inundation by overland flow. Two other groups of taxa were identified by this analysis: the `pioneer' group, comprising mainly nitrogen-fixing lichens which occupy the zone of active erosion at the lower edge of the groves, and the `opportunists' dominated by liverworts, occupying the shallow depressions or bays at the margins of the groves and the interception zones. This study confirms that the non-vascular lichens and bryophytes in these arid soil crusts, are, like the vascular plants, strongly patterned according to geomorphic zone, being most strongly associated with soil surface and erosional features.     

广西藓类植物新记录的报道

通过对广西西南部邻近中越边境山地的多次野外采集获得1 952份藓类标本,经鉴定和查阅以往广西的相关文献资料,报道了广西藓类植物新记录47种,隶属25科36属。文中列出了新记录种及其标本信息,结合文献对每个种的分布进行了更新。该研究结果不仅丰富了广西苔藓植物的物种多样性,同时也为广西乃至中国苔藓植物的区系研究和资源开发提供了新的数据资料。     

Historical herbarium specimens in plant molecular systematics — an example from the fern genus <Emphasis Type="Italic">Lindsaea</Emphasis> (Lindsaeaceae)

We extracted, amplified and sequenced DNA from historical herbarium specimens and silica-dried samples of the fern genus Lindsaea in order to study the sequencing success between the two kinds of samples. High quality sequences were obtained from 57% of the herbarium samples. The specimens age was found to be of little importance for sequencing success when less than 75 years, but the colour of a specimen was found more indicative of sequencing success. Shorter DNA fragments were sequenced successfully twice as often as longer fragments from the herbarium material; in relatively recently collected silica-dried material longer sequences were obtained almost as frequently as short ones. No obvious differences in sequencing success between material originating from different herbaria was observed. We conclude that by using specifically designed DNA extraction protocols and by sequencing short DNA fragments from carefully selected specimens, herbarium material and type specimens can be successfully used in molecular systematics. Typical material or specimens from the type locality (topotypes) should be preferred, when placing a species in a phylogeny.     

浙江省清凉峰自然保护区苔藓植物区系成分研究

通过对2013~2014年于清凉峰自然保护区采集的1 500余份苔藓植物标本进行鉴定和文献研究,统计分析了清凉峰自然保护区苔藓植物种类组成,区系地理成分及特点,并对该地区与相邻8个地区藓类植物的丰富度系数比较和相似性系数比较,以明确该地区苔藓植物不同的生态分布类型,揭示浙江清凉峰保护区苔藓植物多样性的变化规律。结果表明:(1)浙江省清凉峰国家自然保护区苔藓植物共有62科143属337种(包括3个变种),其中苔类植物20科29属49种,藓类植物42科114属288种。(2)藓类植物的优势科9个、优势属16个。(3)该地区苔藓植物区系成分主要以东亚成分为主(占总种数41.84%),温带成分次之(占总种数28.57%),热带成分再次之(占总种数20.74%)。(4)浙江清凉峰自然保护区苔藓植物区系与浙江省境内的金华山、大盘山、天目山和凤阳山的亲缘关系较为密切。(5)浙江清凉峰自然保护区苔藓植物生态分布类型复杂多样,以石生类型(占总标本数的47.09%)最多,其次为岩面土生(占总标本数的27.68%)和土生(占总标本数的12.85%)类型。     

贵州普定喀斯特石生藓类生态分布与环境因子的相关分析

苔藓植物与环境的相关性一直是苔藓生态学的研究热点,研究喀斯特石生苔藓分布与环境的关系,为深入石生苔藓植物生态学研究提供理论依据。选择贵州普定石漠化区域石生环境中的5个大样方25个小样方125个采集点进行调查,采集苔藓植物并测定收集了相关的环境因子数据。结果发现:(1)样方共采集到标本267份,经鉴定有石生藓类54种。其中,丛藓科、灰藓科为优势科(属数≥3),曲柄藓属、扭口藓属、真藓属等为优势属(种数≥3),丛藓、细牛毛藓、美灰藓、绿羽藓为优势种(次数≥3)。(2)运用典范对应分析(CCA)分析了影响该地区石生藓类分布的主要环境因素,发现石生藓类分布与坡度呈显著的负相关;与其它环境因子相关的重要性依次为岩面倾角、海拔、土壤中全钾含量、土壤厚度和pH。因此,研究环境因子对石生藓类分布的关系可以为石漠化综合治理以及石面绿化植物的筛选提供参考。     

NEW DEPSIDES FROM LICHENS: MICROCHEMICAL METHODOLOGIES APPLIED TO THE STUDY OF NEW NATURAL PRODUCTS DISCOVERED IN HERBARIUM SPECIMENS

A high-performance liquid chromatographic method originally devised to provide a reproducible measure of retention is used to determine chemical structures of new compounds in lichens. The method relates retention to a standard homologous series so that retention indices of members of other homologous series are linear with carbon number. When two or more homologs are known, retention of new members of the same series can be accurately predicted. Combining this method with other microchemical techniques allows the determination of structures of lichen depsides in small samples of crustose species for which the total material is meager and known only from the herbarium. The conditions are sufficiently mild that chemical decompositions of labile depsides are avoided. Eight new orcinol-type p-depsides with seven- or nine-carbon oxidized sidechains on one or both rings are reported from an endemic Japanese species, Haematomma pachycarpum. The nine-carbon sidechain is the longest ever found in lichen depsides. These new compounds are higher homologs of the known lichen products confluentic, 2‘-O-methylperlatolic, 4-O-methylolivetoric, and 2‘-C-methylmicrophyllinic acids. In addition, an isocoumarin with a nine-carbon sidechain occurs in some specimens. Haematomma pachycarpum has the most complex secondary-product chemistry thus far reported for any lichen species. The significance of the technique used here lies in providing a way to characterize new compounds in rare or endangered species, thereby completing data sets for phylogenetic reconstructions.     

Multi‐scale patterns of moss and lichen richness on the Antarctic Peninsula

Mosses and lichens are the dominant macrophytes of the Antarctic terrestrial ecosystem. Using occurrence data from existing databases and additional published records, we analyzed patterns of moss and lichen species diversity on the Antarctic Peninsula at both a regional scale (1°latitudinal bands) and a local scale (52 and 56 individual snow‐ and ice‐free coastal areas for mosses and lichens, respectively) to test hypothesized relationships between species diversity and environmental factors, and to identify locations whose diversity may be particularly poorly represented by existing collections and online databases. We found significant heterogeneity in sampling frequency, number of records collected, and number of species found among analysis units at the two spatial scales, and estimated species richness using projected species accumulation curves to account for potential biases stemming from sample heterogeneity. Our estimates of moss and lichen richness for the entire Antarctic Peninsula region were within 20% of the total number of known species. Area, latitude, spatial isolation, mean summer temperature, and penguin colony size were considered as potential covariates of estimated species richness. Moss richness was correlated with isolation and latitude at the local scale, while lichen richness was correlated with summer mean temperature and, for 17 sites where penguins where present with <20 000 breeding pairs, penguin colony size. At the regional scale, moss richness was correlated with temperature and latitude. Lichen richness, by contrast, was not significantly correlated with any of the variables considered at the regional scale. With the exception of temperature, which explained 91% of the variation in regional moss diversity, explained variance was very low. Our results show that patterns of moss and lichen biodiversity are highly scale‐dependent and largely unexplained by the biogeographic variables found important in other systems.     

Comparison of Herbarium Label Data and Published Medicinal Use: Herbaria as an Underutilized Source of Ethnobotanical Information

The use of herbarium specimens as vouchers to support ethnobotanical surveys is well established. However, herbaria may be underutilized resources for ethnobotanical research that depends on the analysis of large datasets compiled across multiple sites. Here, we compare two medicinal use datasets, one sourced from published papers and the other from online herbaria to determine whether herbarium and published data are comparable and to what extent herbarium specimens add new data and fill gaps in our knowledge of geographical extent of plant use. Using Brazilian legumes as a case study, we compiled 1400 use reports from 105 publications and 15 Brazilian herbaria. Of the 319 species in 107 genera with cited medicinal uses, 165 (51%) were recorded only in the literature and 55 (17%) only on herbarium labels. Mode of application, plant part used, or therapeutic use was less often documented by herbarium specimen labels (17% with information) than publications (70%). However, medicinal use of 21 of the 128 species known from only one report in the literature was substantiated from independently collected herbarium specimens, and 58 new therapeutic applications, 25 new plant parts, and 16 new modes of application were added for species known from the literature. Thus, when literature reports are few or information-poor, herbarium data can both validate and augment these reports. Herbarium data can also provide insights into the history and geographical extent of use that are not captured in publications.