Nuovi micromiceti su Pandanacee

Abstract

Helianthemetea guttati communities are pioneer spring and early summer ephemeral grasslands, dominated by non-nitrophilous therophytes. In Continental Portugal, these communities have not yet been fully investigated, and thus the objectives of the present study are: (1) to identify community types in therophytic grasslands; (2) to recognize those communities that configure the European priority habitat 6220* (pseudo-steppe with grasses and annuals); (3) to establish environmental gradients underlying their spatial variation; (4) to assess how floristic composition is affected by land use factors. Vegetation sampling using phytosociological methodology was carried out on 80 grasslands. Modified Twinspan classification and canonical correspondence analysis (CCA) was applied for the classification and ordination of relevés whereas partial CCA (pCCA) and variation partitioning were used to assess the relative influence of individual land use factors. Some 270 species were identified across 11 community types whose floristic patterns were mainly explained by environmental gradients related to altitude and soil type while land use variables could only explain a small part of the floristic variation. Based on biogeography and the determination of diagnostic species, four phytosociological new associations and a new subassociation are proposed: Holco-Brachypodietum distachyi, Holco-Micropyretum tenellae, Micropyro-Anthoxanthetum aristati and Leontodonto-Vulpietum bromoidis vulpietosum membranaceae.     

Environmental factors and vegetation composition, Lefka Ori massif, Crete, S. Aegean

Aim The aim of this study was to explore the environmental factors that determine the spatial distribution of oro‐mediterranean and alti‐mediterranean plant communities in Crete. Location The paper provides a quantitative analysis of vegetation–environment relationships for two study areas within the Lefka Ori massif Crete, a proposed Natura 2000 site. Methods Eleven environmental variables were recorded: altitude, slope, aspect, percentage of bare rock, percentage of unvegetated ground, soil depth, pH, organic matter content and percentages of sand, silt and clay content. Classification of the vegetation was based on twinspan , while detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) were used to identify environmental gradients linked to community distribution. Results One hundred and twenty‐five species were recorded from 120 plots located within the two study areas. Forty‐seven of the recorded species are endemic, belonging to 35 families. Hemicryptophytes and chamaephytes were the most frequent, suggesting a typical oro‐mediterranean life form spectrum. The samples were classified into five main community types and one transitional. The main gradients, identified by CCA, were altitude and surface cover type in the North‐west site, while in the Central site the gradients were soil formation‐development and surface cover type. Main conclusions The use of classification in combination with ordination techniques resulted in a good discrimination between plant communities and a greater understanding of controlling environmental factors. The methodology adopted can be employed for improving baseline information on plant community ecology and distribution in Mediterranean mountain zones.     

Macrophyte community structure and species occurrence in relation to environmental determinants in the ephemeral aquatic habitats of Gavdos,Greece

The aims of this study were to explore the environmental factors that determine the distribution of plant communities in temporary rock pools and provide a quantitative analysis of vegetation–environment relationships for five study sites on the island of Gavdos, southwest of Crete, Greece. Data from 99 rock pools were collected and analysed using Two-Way Indicator Species Analysis (TWINSPAN), Detrended Correspondence Analysis (DCA) and Canonical Correspondence Analysis (CCA) to identify the principal communities and environmental gradients that are linked to community distribution. A total of 46 species belonging to 21 families were recorded within the study area. The dominant families were Labiatae, Gramineae and Compositae while therophytes and chamaephytes were the most frequent life forms. The samples were classified into six community types using TWINSPAN, which were also corroborated by CCA analysis. The principal gradients for vegetation distribution, identified by CCA, were associated with water storage and water retention ability, as expressed by pool perimeter and water depth. Generalised Additive Models (GAMs) were employed to identify responses of four dominant rock pool species to water depth. The resulting species response curves showed niche differentiation in the cases of Callitriche pulchra and Tillaea vaillantii and revealed competition between Zannichellia pedunculata and Chara vulgaris. The use of classification in combination with ordination techniques resulted in a good discrimination between plant communities. Generalised Additive Models are a powerful tool in investigating species response curves to environmental gradients. The methodology adopted can be employed for improving baseline information on plant community ecology and distribution in Mediterranean ephemeral pools. Handling editor: S. M. Thomaz     

拉萨河谷草地群落的数量分类与排序

采用TWINSPAN数量分类和DCA、CCA排序的方法,对拉萨河谷草地23个样点进行统计分析。结果显示:(1)TWINSPAN数量分类将拉萨河谷草地群落划分成8种类型,拉萨河谷的草地群落分布呈现明显的垂直地带性分布格局。(2)TWINSPAN分类所划分的各群落在DCA排序图上都有各自的分布范围和界限,说明DCA排序能较好的反应各群落与其环境资源之间的关系,同时,TWINSPAN的分类结果也在排序图上得到较好的印证。(3)样点DCA排序的第一轴基本反映了海拔高度的变化梯度,第二轴基本反映了坡向的变化。(4)样点CCA排序表明,影响群落分布的主要环境因子是海拔,其次是坡向。CCA排序进一步阐明了拉萨河谷草地群落分布决定于海拔和坡向等环境因子,并间接验证了TWINSPAN的分类结果。(5)物种CCA排序和TWINSPAN分类结果表明:植物群落中物种的分布格局与植物群落类型的分布格局存在一定的相似性,物种的分布格局在很大程度上影响着群落的分布格局。     

协惯量分析与典范对应分析在植物群落排序中的应用比较

采用协惯量分析(PCA-CA COIA)和典范对应分析(CCA)两种排序方法, 对北京小龙门林场的黄檗 (Phellodendron amurense)群落进行了分析, 并用Spearman秩相关系数检验了对应排序轴的相关性。两种排序方法得出的结果基本一致, 两者的第一排序轴都反映了海拔高度和坡向对群落分布的影响, 而各自第二、第三排序轴所代表的环境意义有所差异, 并出现了交叉, 但是两者的前3个排序轴均反映了海拔、坡位、土壤厚度和凋落物层厚度的变化趋势, 说明在环境因子个数较少或共线性效应不明显的情况下, 协惯量分析也能达到CCA的分析效果, 并且在排序轴特征值解释量上高于典范对应分析。     

珠穆朗玛峰自然保护区小叶金露梅灌丛群落数量分类和排序及其多样性垂直格局

根据野外样方调查数据,采用双向种指示分析(TWINSPAN)和典范对应分析(CCA),对珠穆朗玛峰国家级自然保护区小叶金露梅灌丛群落进行分类和排序,并分析物种多样性沿海拔梯度的分布格局。结果表明:(1)该区域24个样地中,记载的维管束植物共有23科45属80种,出现频度较高的种有小叶金露梅(Potentilla parvifolia)、高山嵩草(Kobresia pygmaea)、木根香青(Anaphalis xylorhiza)、垫状点地梅(Androsace tapete)、藏沙蒿(Artemisia wellbyi)、垫状雪灵芝(Arenaria pulvinata)和柴胡红景天(Rhodiola bupleuroides)等。(2)经TWINSPAN等级分类将该区域小叶金露梅灌丛24个样地划分为10个群丛类型。(3)样地和物种CCA二维排序结果表明,海拔和坡位是影响该区域小叶金露梅灌丛群落和物种分布格局的主要环境因子。(4)该区域小叶金露梅灌丛群落物种丰富度、Shannon-Wiener指数和Simpson指数均随海拔升高呈下降的趋势,而Pielou指数呈上升的趋势。(5)样地中优势种小叶金露梅的盖度和高度沿海拔梯度呈显著下降趋势。     

河北小五台山国家级自然保护区森林群落与环境的关系

植物与环境之间的关系是一个复杂的演变过程,运用数量生态学方法探讨森林群落的物种组成、种群的生态特征、不同植物群落与环境之间的关系,有助于保护该区森林群落的稳定性和生物多样性。根据148个森林群落样方数据,选用双向指示种分析(TWINSPAN)和典范对应分析(CCA)方法,对河北小五台山国家级自然保护区森林群落进行分类和排序研究。结果表明:(1)TWINSPAN将该区的森林群落分为20个类型;(2)CCA排序结果较好地反映出群落分布格局与环境梯度的关系,各个森林群落类型在前两轴分异明显,在11个环境因子中,海拔、坡位、凋落层厚度、土壤导电率、土壤温度、土壤湿度、土壤厚度和干扰程度这8个环境因子对森林群落的分布起较大的作用,影响森林群落的分布格局,形成不同的植被类型。(3)乔木层优势种的CCA二维排序图所揭示的环境梯度与群落类型的分布有很大的相似性;t值双序图阐明了海拔、凋落层厚度、土壤温度、干扰程度等环境因子对森林群落乔木层优势种有着重要影响。采用TWINSPAN分类与CCA排序的方法,较好地解释了森林群落与环境因子的关系,为小五台山地区森林生态系统的科学管理和保护提供了理论依据,研究结果也为同类地区森林生态系统研究及保护提供参考和借鉴。     

Habitat use at fine spatial scale: how does patch clustering criteria explain the use of meadows by red deer?

Large mammalian herbivores are keystone species in different ecosystems. To mediate the effects of large mammalian herbivores on ecosystems, it is crucial to understand their habitat selection pattern. At finer scales, herbivore patch selection depends strongly on plant community traits and therefore its understanding is constrained by patch definition criteria. Our aim was to assess which criteria for patch definition best explained use of meadows by wild, free-ranging, red deer (Cervus elaphus) in a study area in Northeast Portugal. We used two clustering criteria types based on floristic composition and gross forage classes, respectively. For the floristic criteria, phytosociological approach was used to classify plant communities, and its objectivity evaluated with a mathematical clustering of the floristic relevés. Cover of dominant plant species was tested as a proxy for the phytosociological method. For the gross forage classes, the graminoids/forbs ratio and the percentage cover of legumes were used. For assessing deer relative use of meadows we used faecal accumulation rates. Patches clustered according to floristic classification better explained selection of patches by deer. Plant community classifications based on phytosociology, or proxies of this, used for characterizing meadow patches resulted useful to understand herbivore selection pattern at fine scales and thus potentially suitable to assist wildlife management decisions.     

Spatial distribution pattern of the riparian vegetation in a basin in the NW Spain

The riparian vegetation of a basin in the NW Spain was studied to establish its spatial variation pattern and to relate floristic and structural differences in the community to environmental factors. Eighty-seven sampling units in 43 sampling stations were used. Samples were classified in 5 groups using Two Way Indicator Species Analysis (TWINSPAN). Three groups represented reaches with riparian wood along their banks: Mediterranean alderwoods and shrubby willow woods. The remaining two corresponded to floodplains with vegetation colonizing moderately eutrophicated deposits of gravel. Structural characteristics of richness and diversity differentiated the alder woods. In these, the shaded environment created by the woody species limited herbaceous vegetation development. This was dominated by Carex acuta subsp. broteriana. Classification and CCA ordination results were compared. The TWINSPAN groups could be recognized in the CCA graph. The ordination was related to a pollution gradient associated with altitude. This pollution gradient involved bank ruderalization, incorporation of nitrophilous species and a decrease in the vegetation quality. The influence of the lithological features on vegetation was also evident.     

秦岭山地天然次生林群落MRT数量分类、CCA排序及多样性垂直格局

采用多元回归树(MRT)对秦岭山地天然次生林群落进行数量分类,采用典范对应分析(CCA)进行排序,分析了秦岭山地天然次生林群落物种多样性沿海拔梯度的变化规律。结果表明:(1) 275个样方共有种子植物195种,隶属61科128属。乔、灌、草3个层次物种多样性变化沿海拔梯度的变化趋势基本一致,呈单峰模型;(2)经交叉验证认为秦岭山地天然次生林群落可分为2类,Ⅰ冬瓜杨(Populus purdomii)+陇东海棠(Malus kansuensis)+蛇莓(Duchesnea indica)群落,Ⅱ锐齿槲栎(Quercus aliena var. acuteserrata)+黄栌(Cotinus coggygria)+茜草(Rubia cordifolia)群落;(3) CCA排序结果揭示了群落生境的分布范围,反映出生态轴的排序意义,较好地反映秦岭山地天然次生林群落与环境因子的关系,其结果表明,海拔、坡向、凋落层厚度和干扰情况4个变量对该地区次生林群落的分布有较大的影响。     

新疆博格达山地面生地衣群落数量分类及物种多样性

根据对新疆博格达山地面生地衣群落27个样点(20 m×20 m)调查的数据,以各地衣种的盖度为指标结合双向指示种分析方法(TWINSPAN)和除趋势对应分析法(DCA)对博格达山地面生地衣群落进行数量分类并分析了群落结构特征及其多样性和相似性。采用典范对应分析法(CCA)对各群落的物种分布格局与环境因子的关系进行了探讨。结果表明,TWINSPAN分析和DCA排序将分布在博格达山的37种地面生地衣分为以下5个群丛。群丛1:膜地卷(Peltigera membranacea)+伴藓大孢蜈蚣衣(Physconia muscigena)群丛;群丛2:地卷(Peltigera rufescens)+伴藓大孢蜈蚣衣(Physconia muscigena)群丛;群丛3:裂边地卷(Peltigera degenii)+平盘软地卷(Peltigera elisabethae)群丛;群丛4:喇叭石蕊(Cladonia pyxidata)+犬地卷(Peltigera canina)群丛;群丛5:盾鳞衣(Placidium squamulosum)+缠结茸枝衣(Seirophora contortuplicata)群丛。物种多样性以群丛4最大为3.427;群丛1最小为0.672;群丛1和群丛4间的相似性最高为0.889,群丛3和群丛4相似性系数最低为0.607。CCA排序结果反映,该地区地面生地衣的分布受到森林植被郁闭度、人为干扰、光照强度的影响,而地表植被盖度和土壤pH等因素的影响不大。     

嵊州市公益林群落数量分类、排序及环境解析

为探讨群落分布和环境间的内在关系,采用二元指示种分析(TWINSPAN)和典范对应分析(CCA)法对嵊州市公益林植物群落94 块固定监测样地(20 m×20 m)进行数量分类与排序.结果表明,采用TWINSPAN 可将94 个公益林监测样地分成14 组,其中以马尾松为建群种的群丛最多,是该区域主要的优势群落类型.CCA 排序结果较好地反映了植物群落与环境因子的关系,其中坡向和坡位是植物群落分布的主导环境因子.此方法可有效解释植物群落分布格局与环境之间的内在联系,为区域公益林分类经营管理提供科学依据.     

群落二维格局分析的两种方法

群落二维空间格局研究能够更好地揭示群落的特征,但在分析方法上有较大的困难。用垂直相交的两条样带在两个方向上同时取样的二维取样法,获得数据,用DCA排序与格局分析相结合的方法,得到群落不同格局规模斑块的长、宽及面积,实现二维格局研究。在亚高山草甸群落格局研究中应用表明,两个方法均是有效的。     

Disturbance and the community structure of stream invertebrates: patch-specific effects and the role of refugia

1. We have previously shown that the impact of spates on stream invertebrates may differ among patches separated by distances of metres or less. Here we analyse the species-specific flood responses of larval chironomids and adult and near mature copepods living in different patch types. Four patch types (with eight replicates of each) were compared: the sandy mid-channel, fine sediments around dams, coarse sediments around dams, and dam debris. Additionally, since some fine sediment patches had been shown previously to act as flow refugia while others did not, we also examined species-specific responses in refugium vs. non-refugium fine sediment patches. Detrended correspondence analysis was used to test for changes in assemblage structure (species composition and relative abundance). 2. Species richness was not altered in a predictable manner by floods; the least stable patch types (mid-channel and coarse patches) did not necessarily show reduced species richness during the spate. 3. As indicated by the spread of DCA ordination scores, there was generally a high degree of overlap in the species composition among the four patch types. Nevertheless, copepod species composition and relative abundance were more similar among patch types during the spate than pre-spate. Spates may induce a re-distribution of copepod species among the patch types. Chironomid species composition and relative abundance were no more similar among patch types during the spate than pre- or post-spate. 4. For both chironomids and copepods, species composition and relative abundance (as assessed by DCA ordination scores) in refugium patches changed more in response to the spate than in the non-refugium patches. An influx of individuals from just a few species for each group was responsible for the change in assemblage structure. Thus, despite the fact that our past work has shown that refugia may confer enhanced resistance and resilience of copepod and chironomid assemblages in terms of total faunal abundances, the present work suggests that resistance and resilience of the species composition of the community apparently are no greater in refugium patches than in non-refugium patches.     

Relationships between macroinvertebrate communities and environmental variables in metal-contaminated streams in south-west England

1. Macroinvertebrate species and information on thirty-nine variables were recorded at forty-six sites on twelve Cornish streams affected to varying extents by past metalliferous mining. 2. Relationships between macroinvertebrate communities and environmental variables were examined using canonical correspondence analysis (CCA). Copper was the strongest correlate with Axis 1 of the analysis, suggesting that it may have a major role in determining community structure. There were also strong correlations between Axis 1 and aluminium, alkalinity, pH, dissolved organic matter and algal cover, and between Axis 2 and discharge. 3. CCA distinguished four site groupings. In spite of seasonal changes in position on the ordination, as revealed through CCA and detrended correspondence analysis (DCA), sites retained their group membership. There were differences in the proportions of the major taxa within the four CCA-derived groups. In the group with the highest copper concentration, Ephemeroptera were absent, but triclads and chironomids were abundant, with Orthocladiinae the dominant group. 4. Changes in the occurrence of species along the metal gradient reflected changes in sensitivity. Tolerance' plots of selected species based on the CCA site ordination diagram were used to identify environmental thresholds. 5. As well as direct toxic effects of copper and aluminium on invertebrate communities, co-precipitation of these two metals could be an important physical factor. Further modifying influences resulting from interactions between toxic metals and pH, alkalinity, hardness and dissolved organic matter contributed to the complexity of conditions affecting the faunal community in metal-contaminated streams. 6. The most severely contaminated sites, with mean copper concentration exceeding 500 μgl^?1, were all characterized by a reduced community dominated by the flatworm Phagocata vitta, the chironomids Chaetodadius melaleucus and Eukiefferiella claripennis and the net-spinning caddis Plectrocnemia conspersa. The necessity for species identification is demonstrated in relation to variation in metal tolerance.     

Diagnostic analysis of conservation zones using remote sensing and GIS techniques in wet evergreen forests of the Western Ghats – An ecological hotspot, Tamil Nadu, India

This study highlights spatial characterization of evergreen forests of the Western Ghats – an ecological hotspot in Tamil Nadu, India – using remote sensing and GIS-based analysis in conjunction with ground-based phytosociological data. The evergreen forests of Tamil Nadu are distributed in four distinct hill ranges, Nilgiri, Anamalai, Palni and Tirunelveli, having different topographic, bioclimatic and disturbance levels. The evergreen forests in these four hill ranges are characterized for their uniqueness in terms of patch characteristics and phytosociology. A vegetation type map was prepared using IRS LISS III satellite data and was used to study the patch characteristics in terms of patch size, number, shape, porosity and landcover diversity (LD). The phytosociological characteristics, namely species richness, diversity, similarity and community assemblages, were studied using ground data collected from 95 sample points of 0.1 ha size. Patch size and number revealed distinct intactness and disturbance levels in these four hill ranges. Evergreen forests in the Tirunelveli hills comprising 216.09 km² are distributed in 306 patches, and in the Palni hills, with 285 km², forests are distributed in 1029 patches, indicating a high level of fragmentation. LD, indicating the spatial heterogeneity of landcover, was very high in the Nilgiri hills and low in the Tirunelveli hills. The spatial analysis helped to delineate homogenous large patches of evergreen forest, which can be adopted for appropriate conservation strategies. A total of 342 tree species belonging to 4490 stems were evaluated for phytosociology. Only 15–28% of similarity in terms of species distribution was found across the hill ranges. Conjunctive analysis of patch characteristics and species distribution showed high species richness in less fragmented evergreen forests and vice versa. The study identified the areas of prioritization in terms of ecorestoration and conservation based on patch and phytosociological characteristics.     

山西太岳山脱皮榆群落的生态梯度分析及环境解释

为了解释山西太岳山脱皮榆(Ulmus lamellosa)群落中物种的分布情况与该群落环境因子之间的相互关系,采用TWINSPAN数量分类和典范对应分析(CCA)与环境因子的变量分离进行讨论。结果表明,TWINSPAN将60个调查样方划分为7种群丛类型,体现了该脱皮榆群落主要以乔木脱皮榆和草本披针叶苔草(Carex lanceolata)为优势种。7种群丛类型与CCA排序结果一致,CCA排序第1轴主要体现了坡位和海拔;坡向与第2排序轴存在显著相关性。Monte Carlo检验结果表明,影响脱皮榆群落物种分布最主要的环境因子是海拔。在环境分离变量解释方面,环境因子解释了39.60%,空间因子解释了7.95%,空间因子与环境因子交互作用解释部分占10.89%。而其中不能解释的部分占41.56%。在该研究区,海拔对植物的分布有较好的解释力,其次是坡位和坡向。     

Effects of Forest Use on Aphyllophoraceous Fungal Community Structure in Sarawak, Malaysia

Aphyllophoraceous fungi are expected to reflect changes in the environmental conditions caused by forest use. To reveal the effects of forest uses on the fungal community structure, we performed a 3‐month survey of aphyllophoraceous species in five forest types (undisturbed primary forest, isolated patches of primary forest, old and young fallow forest, and rubber plantations) in Sarawak, Malaysia in 2005. We used a canonical correspondence analysis (CCA) to reveal the relationships between fungal community composition and the environmental variables (canopy openness, soil water potential, amount and composition of coarse woody debris, litter mass, basal area, plant species composition). A total of 155 samples from 67 species were collected during the study period. The fungal species density represented by the number of species in a transect differed significantly among forest types. The fungal species density increased significantly with increasing number of pieces of coarse woody debris (CWD), but decreased significantly with increasing the scores of second axis of principal component analysis (PCA) for plant species composition. In the CCA ordination, automatic forward selection revealed that only the number of pieces of CWD significantly affected the fungal species composition. The occurrences of Flabellophora licmophora, Coriolopsis retropicta, Microporus vernicipes, and Amauroderma subrugosum were positively correlated with the number of pieces of CWD. Our study clearly demonstrated that forest use negatively affected aphyllophoraceous fungal diversity and suggest that the quantity of CWD would be an important determinant of fungal diversity and composition.     

Syntaxonomy and phytogeographical differentiation of the Fagus woods in the Southwest Balkan Peninsula

Abstract. Up to now the syntaxonomy of the Fagus woods of the southern Balkans is far less known than that of beech communities in the northwestern part of the Balkan Peninsula and in central and western Europe. A set of 607 phytosociological relevés from southeastern Serbia, the Former Yugoslav Republic of Macedonia, and northern and central Greece was subjected to numerical classification and ordination. These results were used for a syntaxonomic revision of Fagus, Fagus‐Abies and Fagus‐Acer woodland communities of the southwestern Balkans. The Doronico columnae‐Fagenion (seven associations from Serbia, the F.Y.R.O.M., and high altitudes in northern Greece) and the Doronico orientalis‐Fagenion (four associations from northern and central Greece) were distinguished. The results of ordination by Canonical Variates Analysis (CVA) showed a very distinct phytogeographical pattern of differentiation between community types. The floristic composition of Fagus woods changes markedly along the north‐south gradient. The share of Subbalkan, Balkan, Submediterranean and Mediterranean species increases significantly towards the south, while the Central‐European, European, Euro‐Siberian, Euro‐Asiatic and Circumpolar species decrease in the same direction. The syntaxonomic differentiation of the Fagus woodlands and their classification into regional suballiances reflects this gradient. With the rise in altitude, the number of Balkan, Mediterranean, Submediterranean, European and Euro‐Asiatic species declines.     

Floristic relationship between scrubland and grassland patches in the Mediterranean landscape of the Iberian Peninsula

In the rural Mediterranean landscape, mosaics of patches of sclerophyllous scrubland and semi-natural grasslands are frequent. The plant communities of these patches, which are physiognomically easy to recognise, are very heterogeneous. The objective of this paper is to determine whether the patches of scrub-grassland represent an integrated response unit of the vegetation with regard to the physical environment (climatic, geographical and edaphic factors) and human use, or whether, on the contrary, this is an independent response. In order to do this a total of 50 sampling sites where scrubland and grassland patches were in contact were studied along a 370 km E-W mesoclimatic gradient from central Spain to Portugal (Iberian Peninsula). Two distinct zones, the east and the west halves of the study area, were identified according to the plant communities. Within each of these zones, each type of patch responded to the previously mentioned factors, differentially and independently. This determined a general lack of inter-community association or floristic correlation between scrub and grassland plant communities in contact. The spatial association is a random process probably related to the particular human management realised on every scrubland and grassland patch. The scrub–grassland pattern, which is characteristic of the Mediterranean landscape, does not represent an integrated response unit of the vegetation to a given environment, but rather the sum of the independent responses to the environment of the two patches in contact.Nomenclature for taxa: T. G. Tutin et al. 1694–1980. Flora Europaea Cambridge University Press, Cambridge.