Phytosociological and geographical survey of Mesohromion communities in Western Europe

A survey is given of the floristic composition and geographical distribution of limestone grasslands assigned to the Mesobromion ereeti, in Western Europe.This survey is based upon 800 vegetation relevés from various investigators throughout the area. Relevant tables have been summarized into a synoptic table with 30 columns. Each column represents a community described as an association or a subassociation.The synoptic table shows a division into a NW European group (Ireland, England, Denmark, and South Sweden) and a Central-W European group (Germany, The Netherlands, Belgium, France, and Switzerland). Associations belonging to the former have, as a rule, a smaller distribution area than the latter.In the latter group only two associations can be distinguished: Mesobrometum erecti and Antherico-Brometum ass. nov. The Mesobrometum is widespread and includes many subassociations. The Antherico-Brometum shows a transition towards the Xerobromion erecti.According to Barkman et al., (1976) the often used name Gentiano-Koelerietum has to be seen as a not-correct synonym for the Mesobrometum erecti.The Mesobromion grasslands can, in general, not be divided based upon their agricultural management system.An attempt is made to interpret the gradual transition in floristic composition of the limestone grasslands from NW to SE, with climatological data.Plant nomenclature follows Heukels & Van Ooststroom (1975) or, else, Clapham et al. (1968) for phanerogams and Vander Wijk et al. (1969) for bryophytes.I am much indebted to Prof. Dr. J. J. Barkman, who encouraged me to tackle the present study and stimulated me throughout its duration.     

Molecular markers provide evidence for a broad‐fronted recolonisation of the widespread calcareous grassland species Sanguisorba minor from southern and cryptic northern refugia

• Calcareous grasslands belong to the most species‐rich and endangered habitats in Europe. However, little is known about the origin of the species typically occurring in these grasslands. In this study we analysed the glacial and post‐glacial history of Sanguisorba minor, a typical plant species frequently occurring in calcareous grasslands.
• The study comprised 38 populations throughout the whole distribution range of the species across Europe. We used molecular markers (AFLP) and applied Bayesian cluster analysis as well as spatial principal components analysis (sPCA) to identify glacial refugia and post‐glacial migration routes to Central Europe.
• Our study revealed significant differences in the level of genetic variation and the occurrence of rare fragments within populations of S. minor and a distinct separation of eastern and western lineages. The analyses uncovered traditional southern but also cryptic northern refugia and point towards a broad fronted post‐glacial recolonisation.
• Based on these results we postulate that incomplete lineage sorting may have contributed to the detected pattern of genetic variation and that S. minor recolonised Central Europe post‐glacially from Iberia and northern glacial refugia in France, Belgium or Germany. Our results highlight the importance of refugial areas for the conservation of intraspecific variation in calcareous grassland species.

     

Observations on north-west european limostone grassland communities IV. Phytosociological notes on chalk grasslands in Denmark

This paper deals with phytosociological studies of Danish chalk grasslands. These grasslands should be reckoned to a new association, viz. Solidagini-Helictotrichetum pratensis, alliance Mesobromion, class Festuco-Brometea. Two geographically vicarious subassociations could be distinguished. Furthermore attention has been payed to the cryptogams occuring in these grasslands, the influence of dominance of an introduced plant species on the species diversity, and the changes during the last decades in species composition as well as in area of the chalk grasslands in Denmark.     

高通量测序分析甘肃高寒草原瑞香狼毒根内生菌群落结构特征研究

【背景】瑞香狼毒是我国天然草地退化的标志性植物之一,在入侵的过程中其根内微生物发挥着重要的作用,探究瑞香狼毒根内微生物群落对有益微生物资源的利用及生物防控瑞香狼毒具有重要意义。【目的】探究甘肃省祁连山东麓不同高寒草原毒性杂草瑞香狼毒根部内生真菌和细菌的组成、多样性等群落结构特征。【方法】通过Illumina MiSeq高通量测序技术对甘肃祁连山东麓不同高寒草原的瑞香狼毒根部内生菌组成及多样性进行分析。【结果】不同高寒草原瑞香狼毒根内子囊菌门(Ascomycota)在真菌中占优势地位;蓝细菌(Cyanobacteria)、变形菌门(Proteobacteria)、放线菌门(Actinobacteria)在根内生细菌中属于优势类群。G4样点的真菌和细菌OTU(OperationalTaxonomic Unit)数量和Shannon多样性指数均高于其他4个采样点。主坐标分析(Principal Coordinate Analysis,PCoA)表明,不同高寒草原瑞香狼毒根内生菌群落之间存在一定的差异。One-Way ANOVA分析结合Pearson相关性分析显示,不同样地瑞香狼毒真菌群落结构在属水平上相关性较小,而细菌在属水平存在一定的相关性。【结论】本研究为进一步开发利用瑞香狼毒内生真菌和细菌资源、阐明瑞香狼毒内生菌群的群落结构和生态功能提供理论参考。     

Plant communities of the montane mesophilous grasslands (Polygono bistortae–Trisetion flavescentis alliance) in central Europe: Formalized classification and syntaxonomical revision

Abstract

The research conducted here presents a syntaxonomical revision of the montane mesophilous meadows of the Polygono bistortae-Trisetion flavescentis alliance in central Europe – the Slovak part of the western and eastern Carpathians. These typical semi-natural grasslands occur mainly as small islands over the calcareous bedrocks. Associations of this alliance have tight relationships to the Arrhenatherion, Mesobromion and Nardo-Agrostion alliances. Formal definitions based on combination of the species groups were used regarding the diversity and geographical range of the Polygono bistortae-Trisetion flavescentis alliance. Following the formal definitions from the eight associations reported previously for Slovakia, only four of them can be placed within the Polygono bistortae-Trisetion flavescentis: Campanulo glomeratae-Geranietum sylvatici, Geranio sylvatici-Trisetetum, Crepido mollis-Agrostietum capillaries and Geranio-Alchemilletum crinitae. A comparison of traditional and formalized classification, and the advantages/disadvantages of the formalized classification are discussed.     

Pattern analysis in Belgian limestone grasslands

Pattern analyses were made in several limestone grasslands of Belgium. Vegetation has been sampled in continuous transects. Monospecific patterns are defined by means of two non-parametric indices; multivariate analysis (reciprocal averaging) completed the analysis. The following kinds of pattern are described and explained: random, aggregate, clump, gap, stationary density variation, non-stationary density variation, gradient and many complex patterns. The multivariate analysis leads to a careful phytosociological study of the transects and to the definition of floristically homogeneous regions. Some particular points of pattern analysis, the problem of vegetation sampling and some perspectives are discussed.     

Role of topography and soils in grassland structuring at the landscape and community scales

The relative importance of abiotic factors in community assembly is debated and thought to be dependent on the scale. I investigated the relative role of topography and soils as structuring agents at the landscape and the community scales in 126 subalpine calcareous grasslands in the Pyrenees, in terms of species composition and abundance. I wished to know: (1) the role of abiotic factors in the organization of plant communities across the landscape; (2) how much of the variation in community distribution was accounted for by abiotic factors; and (3) how well their role applied to the distribution of dominant species at the landscape and the community scales. The hypothesis was: abiotic factors play an important role in community distribution in the landscape, but species interactions are more important within communities. Multivariate methods generated four communities, organized in two contrasting groups along the main vegetation axis, which explained 13% of the variation: mesic grasslands (Nardus stricta and Festuca nigrescens communities) and xeric grasslands (Carex humilis and Festuca gautieri communities). Mesic communities were more acidic and fertile than xeric communities. Changes in the abiotic environment, accounting for up to 80% of the variation in the vegetation, were smooth, while the transition between xeric and mesic grasslands was sharp in terms of species composition. The distribution in the landscape of the first main species from each community was closely related to abiotic factors, which modeled poorly the abundance of the main species at smaller scales. At the within-community scale, the explanatory power of biotic relationships was community dependent, producing the most significant models for plants highly dominant within their communities, such as N. stricta and F. gautieri. Contrary to current hypothesis, there was a shift from mainly positive relationships among dominant species in fertile mesic communities to mainly negative in infertile xeric ones.     

Species distribution modeling and molecular markers suggest longitudinal range shifts and cryptic northern refugia of the typical calcareous grassland species Hippocrepis comosa (horseshoe vetch)

Calcareous grasslands belong to the most diverse, endangered habitats in Europe, but there is still insufficient information about the origin of the plant species related to these grasslands. In order to illuminate this question, we chose for our study the representative grassland species Hippocrepis comosa (Horseshoe vetch). Based on species distribution modeling and molecular markers, we identified the glacial refugia and the postglacial migration routes of the species to Central Europe. We clearly demonstrate that H. comosa followed a latitudinal and due to its oceanity also a longitudinal gradient during the last glacial maximum (LGM), restricting the species to southern refugia situated on the Peninsulas of Iberia, the Balkans, and Italy during the last glaciation. However, we also found evidence for cryptic northern refugia in the UK, the Alps, and Central Germany. Both species distribution modeling and molecular markers underline that refugia of temperate, oceanic species such as H. comosa must not be exclusively located in southern but also in western of parts of Europe. The analysis showed a distinct separation of the southern refugia into a western cluster embracing Iberia and an eastern group including the Balkans and Italy, which determined the postglacial recolonization of Central Europe. At the end of the LGM, H. comosa seems to have expanded from the Iberian refugium, to Central and Northern Europe, including the UK, Belgium, and Germany.     

Vegetation analysis in colonies of an endangered rodent, the Cabrera vole (Microtus cabrerae), in southern Portugal

The Cabrera vole (Microtus cabrerae) is a threatened rodent endemic in the Iberian Peninsula with a patchy distribution and specific microhabitat requirements. This aim of this study was to document the composition of plant communities in habitats of Cabrera vole colonies in southern Portugal. Differences observed in plant species composition were also compared with vegetation structure, taxonomic and life form groups, species and group diversity, disturbance, topography and soil properties. Vegetation was sampled between March and July 2004, in 26 colonies occurring in five geographical areas. Grasses were the most abundant, common and diverse family in the colonies, and the perennial grass Agrostis castellana was present in 92% of colonies, with a mean cover of 16% of the site. Other frequently occurring species were Briza maxima (85%), Vulpia myuros (85%), Gaudinia fragilis (81%), Leontodon spp. (81%), Avena barbata (77%), Bromus hordeaceus (77%) and Tolpis barbata (77%). Colonies were classified in eight vegetation groups that included meadows, tall perennial grasslands, manured meadows with tall sedges, annual grasslands and ruderal and nitrophilous grasslands. Main gradients associated with composition differences were grass richness, annual and perennial grass cover, vegetation structure (herbaceous vegetation height), soil properties (texture and moisture), disturbance (ruderal species) and colony dimensions (area). Results suggest that the Cabrera vole is able to exploit a wide variety of grasslands, with a varying degree of ecological disturbance. Meadows and perennial grassland communities seem to be higher-quality microhabitats for voles.     

A comparative analysis of extreme thermophilic bacteria belonging to the genus Thermus

Several extreme thermophilic Gram negative bacteria found in a thermally polluted river in Belgium have been compared with Thermus strains isolated from widely distant geographical areas. This analysis has become possible after the design of a new culture medium (162).All strains examined (including the isolate successively denominated Flavobacterium thermophilum and Thermus thermophilus) were found to be morphologically identical with strain YT-1 of Thermus aquaticus. The cells are immotile, rod-like, strictly aerobic, catalase and oxidase positive. They produce amylase, hydrolyze gelatin and are confirmed to be highly sensitive towards penicillin.The nutritional pattern of all strains has been analysed extensively, by testing a broad spectrum of possible substrates.The strains display a uniform response to the microbiological tests applied and most probably belong to the same species: Thermus aquaticus.Abbreviations GC guanosine cytosine - ATCC American Type Culture Collection - DSM Deutsche Sammlung von Mikroorganismen     

Diversity of plant assemblages in isolated depressional wetlands from Central-Western Europe

A closed depression is defined as a geomorphologic element where a sediment depository is encircled by hillslopes. Despite the fact that closed depressions are often the only stagnant water points in many European landscapes, few ecological researches on their plant assemblages have been done. The main goal of this study was to give first results of the environmental factors responsible for the vegetation composition, richness and rarity in the closed depressions of the Lorraine biogeographical district (Belgium, France and grand-duché de Luxembourg). We surveyed for plant presence 85 forest and 77 grassland closed depressions. For each site, wetland area, local environmental factors and regional connectivities registered. For each species, the Ellenberg values were compiled. To investigate the main source of variation in species composition and in species richness (including richness in rare species), Non-metric Multidimensional Scaling analyses and Generalized Linear Models were respectively used. Species pools in forest (forest and preforest species) and in grassland (bog plants, pioneers, helophytes) were quite different. In both landscapes, a gradient from plants typical of basic high-productive soils to plants typical of acid low-productive soils reflects a shared successional gradient. The accumulation of organic matter allowed the establishment of Sphagnum spp., which slowly acidified the soil and thus acted as ecosystem engineers for the arrival of bog plants. Moreover, the species composition was additionally driven by the plant light tolerance in forests and by the plant water requirements in grasslands. Mechanisms of species accumulation (increase in species richness) were different in forests and in grasslands: respectively related to the plant light tolerance and to the wetland area. At the regional level, the averaged soil productivity was negatively related to the richness in rare species. Indeed, perennial highly-competitive plants such as Glyceria spp., Iris pseudacorus or Urtica dioica impeded the establishment of smaller and rarer species. At the habitat level, isolated closed depressions (due to either low connectivity or low grazing pressure) have more habitat rare species, giving evidences of dispersal limitation in plant assemblages of closed depressions.     

Recovery of species diversity and composition of braconid parasitic wasps after reforestation of degraded grasslands in lowland East Kalimantan

Human-induced disturbances and wildfires can transform areas of tropical rainforest into Imperata-dominated grassland, but it may be possible that recovery of biodiversity is facilitated by reforestation with fast-growing trees. We compared the assemblages of braconid wasps as parasitoids of taxonomically diverse groups of insects among Imperata grasslands, young and mature plantations of Acacia mangium, young secondary forests after wildfires, and old secondary forests in the lowland of East Kalimantan. The abundance and species richness of braconids, which had declined in Imperata grasslands, somewhat increased in Acacia plantations, and also the species composition of braconids in Acacia plantations was transitional between Imperata grasslands and old secondary forests. Parasitoids of detritivores and wood borers increased markedly after plantation, while those of herbivores showed a distinct turnover of species all over the range from grasslands to old secondary forests. The plantation of A. mangium had most likely facilitated the recovery of the diversity of host forest and their parasitoids, but the recovery was just at the rudimentary stage even in mature plantations. Monitoring of parasitic wasps would be useful to test the continuous recovery of forest biodiversity in plantation stands.     

A functional analysis of vegetation dynamics in abandoned and restored limestone grasslands

Question: How do various plant functional groups and types are related to regeneration of limestone grasslands and to temporal changes in cover of trees and shrubs? Which mechanisms are responsible for changes in differently treated grasslands? Location: Southern Poland, 50° 01'N, 19° 50'E. Methods: A 12‐year restoration experiment after the clearing of a 35‐yr‐old secondary pine wood developed on limestone grassland, and a parallel observation of succession in neighbouring abandoned grassland were carried out. Changes in cover of species with different functional traits and established strategies in relation to time and cover of shrub layer were analysed using multiple linear regression. Results: Over 12 years the composition and cover of species changed in the studied grasslands, but significant differences still occurred between the old grassland and the grassland restored in former wood gaps and that developed in former closed wood. Despite the very close proximity of the old grassland, the composition of restored grasslands was still much determined by initial conditions. In these grasslands, the cover of Festuco‐Brometea species was significantly lower than in the old grassland. In all sites, the cover of species with large leaves and with leaves distributed regularly along the stem, and the cover of CS strategists increased. With increasing shrub cover in restored grasslands, the cover of annuals and biennials, tall species, and R and CS strategists decreased. Conclusions: Limited availability of seeds seems the principal reason for the weak regeneration of xerothermic limestone grasslands. Trees and shrubs inhibit the developed of both early successional species and perennials occurring in established communities. They significantly hinder regeneration of grasslands and should be cut every five or six years.     

川西北高寒地区多年生禾本科人工草地土壤线虫群落动态

为了查明青藏高原地区建植多年生禾本科人工草地对土壤线虫群落的影响及土壤线虫群落随人工草地建植年限增加的变化趋势,以川西北红原县2018年建植的垂穗披碱草样地(Elymus nutans)和老芒麦样地(Elymus sibiricus)为研究对象,以天然草地(Natural grassland)为对照,于2018—2020每年9月对土壤线虫群落、植被和土壤因子等进行连续监测。研究共分离到土壤线虫38470条,隶属于2纲8目65科172属,平均密度2482条/100g干土;食细菌线虫是研究样地的主要营养类群。垂穗披碱草和老芒麦样地的土壤线虫群落组成结构与天然草地间存在明显差异,但差异性随建植年限增加逐渐降低。线虫群落密度和多样性指数在垂穗披碱草、老芒麦和天然草地间均无显著差异。随建植年限增加,土壤线虫群落密度显著上升,Pielou指数显著下降,丰富度和Shannon指数无显著变化;植物寄生线虫密度和比例在第2年显著增加,并成为优势营养类群。线虫群落生态指数在不同草地类型间无显著差异,但垂穗披碱草和老芒麦样地的线虫基础指数随建植年限增加先增加,在第3年时显著降低(P<0.05)。冗余分...     

Invasion history, habitat preferences and phenology of the invasive ladybird Harmonia axyridis in Belgium

Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe, H. axyridis is considered to be an invasive alien species because of its potential to disrupt native ladybird communities. Since 1999, the Belgian Ladybird Working Group mapped all Belgian Coccinellidae and recorded data on substratum plants and habitat. The first feral H. axyridis population in Belgium was recorded in 2001, but the expansion rate is decreasing because it now colonised the whole country. Recorded occupancy in Belgium showed an average rate of increase of 189% between 2002 and 2006. In Belgium, H. axyridis occurred in a wide range of habitats, including those of high conservation value. However, habitat and land cover analysis showed that H. axyridis is more frequently found in urbanised landscapes than in semi-natural landscapes. A niche overlap analysis based on plant use data showed that the potential to affect native species is higher for generalist, deciduous and coniferous tree ladybird species than for heathland and wetland specialist species. Phenology data showed that H. axyridis is able to reproduce later in the year than native species. Based on recorded distribution, ecology and phenology, we discuss the potential of H. axyridis to disrupt native ladybird assemblages in Belgium.     

Observations on North-West European limestone grassland communities: Phytosociological and ecological notes on chalk grasslands of Southern England

Summary In 1970 chalk grasslands were studied in southern England, according to the principles of the French-Swiss School. The communities described belong to the associationCirsio-Brometum Shimwell 1971, emend., allianceMesobromion erecti (Br. Bl. & Moor 1938) Oberd. 1957, orderBrometalia W. Koch (1926 n.n.) Br.-Bl. 1936, of the classFestuco-Brometea Br.-Bl. 1936, emend. Tüx. 1961. Within theCirsio-Brometum three new sub-associations are distinguished: 1) subass. nov.anthoxanthetosum odorati; 2) subass. nov.Cornetosum sanguineae; and 3) subass. nov.campanuletosum rotundifoliae.Special attention is paid to the relationship that exists between the management of these communities and the seral stages that lead to the development of serub following the decreasing influence of grazing. Ulex europaeus scrub occurring on calcareous soil was also studied. The cryptogams of the chalk grasslands, mainly Musci, were investigated in detail.Plant nomenclature follows Heukels & van Ooststroom (1970) or for taxa not mentioned by these, Clapham et al. (1968) for phanerograms and Margadant (1959) and van der Wijk et al. (1969) for bryophytes.Contribution to the Symposium on Plant species and Plant communities, held at Nijmegen, 11 12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff.     

Phylogeny and Evolutionary History of the Ground Squirrels (Rodentia: Marmotinae)

Although ground squirrels (Spermophilus) and prairie dogs (Cynomys) are among the most intensively studied groups of mammals with respect to their ecology and behavior, a well-resolved phylogeny has not been available to provide a framework for comparative and historical analyses. We used complete mitochondrial cytochrome b sequences to construct a phylogeny that includes all 43 currently recognized species in the two genera, as well as representatives of two closely related genera (Marmota and Ammospermophilus). In addition, divergence times for ground squirrel lineages were estimated using Bayesian techniques that do not assume a molecular clock. All methods of phylogenetic analysis recovered the same major clades, and showed the genus Spermophilus to be paraphyletic with respect to both Marmota and Cynomys. Not only is the phylogeny at odds with previous hypotheses of ground squirrel relationships, but it suggests that convergence in morphology has been a common theme in ground squirrel evolution. A well-supported basal clade, including Ammospermophilus and two species in the subgenus Otospermophilus, diverged from all other ground squirrels an estimated 17.5 million years ago. Between 10 and 14 million years ago, a relatively rapid diversification gave rise to lineages leading to marmots and to several distinct groups of ground squirrels. The Eurasian ground squirrels diverged from their North American relatives during this period, far earlier than previously hypothesized. This period of diversification corresponded to warming climate and spread of grasslands in western North America and Eurasia. Close geographic proximity of related forms suggests that most species evolved in or near their current ranges.     

Contributions to the sociology and chorology of contrasting plant communities in the southern part of the ‘Wienerwald’ (Austria)

Five plant communities contrasting in successional status and human impact from the southern part of the ‘Wienerwald’ (Austria) are analyzed using vegetation relevés, spectra of area types and a newly proposed disjunction quotient. A climax community (Asperulo-Fagetum), a subclimax community (Querco-Carpinetum s.l.), an anthropogenous substitute community (Mesobromion) and two natural, non-climax permanent communities (Euphorbio saxatilis-Pinetum nigrae and Fumano-Stipetum eriocaulis) are recognized. The disjunction quotient is defined as the number of partial (discontinous) areas divided by the size of the total area of distribution of a species. In particular, the average disjunction quotients of the species in the first two communities reflect relatively table environments only slightly influenced by man, with many ancient, stable taxa. These communities are characterized by species with well-delimited, stable distribution areas. The species in the Mesobromion community have very low average disjunction quotients as its component species are widely and continuously distributed and are often promoted by man. In contrast to these communities, the species linked to the natural permanent, non-climax communities of extreme habitats, have high distribution quotients i.e. small, disconinuous areas; this illustrates the relic character of these plant communities and of the eastern edge of the Alps is a whole. Using the highly variable disjunction quotient of all species and communities examined, the concepts of climax and permanent communities (of different origin) are discussed with regard to European conditions.     

Ecological amplitude in Silene nutans in relation to allozyme variation at the western margin of its distribution

Abstract. At the western border of its geographical range, Silene nutans (Caryophyllaceae) has evolved two groups of parapatric populations showing distinct allozyme patterns and apparently occurring on different bedrock types. This study tests the hypothesis that these groups represent edaphic ecotypes. With this in view, the ecological amplitude of 36 populations of Silene nutans from Belgium was investigated and their synecology specified using vegetation composition and soil parameters; Ellenberg indicator values were used for ecological interpretation of the vegetation analyses. The results provide evidence that allozyme and habitat variations are correlated, the two groups occurring on contrasting soil with distinct vegetation types. One group is restricted to alkaline soils and typically occurs in open calcareous grasslands, fringes and scrub. The other group is characterized by wider autecological and synecological ranges with a bimodal pH- distribution, occurring on two kinds of siliceous soils: (1) neutral soils, supporting vegetation with a high species richness comprising many mesophilous species, and (2) CaCO₃ -free bedrocks characterized by acidophilous, species-poor, open dry grasslands and woodlands. It also appears that factors not related to soil chemistry, such as microclimate, are involved in the ecological specialization of the two ecotypes. It is argued that these ecotypes exemplify adaptive radiation and parapatric speciation at the margin of a species distribution area.     

Fine-scale spatial distribution of leaves and shoots of two chalk grassland perennials

The fine-scale spatial distribution of leaves and shoots of Brachypodium pinnatum and Carex flacca, two rhizomatous graminoids, was investigated in two chalk grasslands in South Limburg (The Netherlands). The objective was to examine whether leaves and shoots of Brachypodium, a dominant species, had a regular distribution on a small scale, as has been suggested for other clonal species that form high-density stands. Patterns were compared to Carex, which is never found to be as abundant as Brachypodium.The number of shoots and leaf contacts were counted in small quadrats, grouped in a grid. Using Moran's I analysis for autocorrelation, it appeared that leaves and shoots of both species were arranged in clumps, and that these clumps were randomly distributed across the soil surface. Shoot clumps in Carex were smaller in diameter and not as pronounced as those in Brachypodium.In most cases, patterns of leaves and shoots were positively correlated, indicating that leaves were predominantly positioned above and around the groups of quadrats where the shoots were attached. However, in dense stands of Brachypodium the positions of leaf clumps were not correlated to those of shoot clumps. This is a result of the tall growth form of this species and its high shoot densities, and it is suggested that this will be a characteristic of any species that dominates a dense stand.