Koeleria macrantha (Ledeb.) Schultes (K. alpigena Domin, K. cristata (L.) Pers. pro parte, K. gracilis Pers., K. albescens auct. non DC.)

Abstract

A compactly tufted perennial; tufts stiff and spiky or more rarely soft and gently spreading, 4–60 (70) cm high. Culms slender, 1–3-noded, erect or sometimes slightly curved, downy towards the panicle or hairless. Leaves light grey-green to mid-green on the adaxial and mid-green to dark green on the abaxial surface. Blades curving abruptly to a hooded tip, up to 25 cm long and 1–3.5 mm wide, stiff or soft, usually flat in unstressed plants when fully expanded, ridged on adaxial surface. Both surfaces extremely variable in degree of hairiness, ranging from glabrous to densely hairy (hairs on margins up to 1.5 mm long). Abaxial surface generally more hairy than adaxial, lower sheaths sparsely to densely hairy. Ligules up to 1 mm long, membraneous, usually surrounded by hairs to 2 mm long. Culm leaves usually 2, upper to 7.5 cm long. Panicles spike-like, very dense and often interrupted in the lower part, narrowly oblong, sometimes tapering towards the top; branches spreading during flowering; 1–12 cm long and 2–10 mm wide, silvery-green or purplish, glistening; rhachillas short, usually densely hairy, but sometimes sparsely so; rhachis sparsely to densely hairy. Spikelets 3–6 mm long, 2–3 flowered; densely clustered on short pedicels, to 4 mm long on lower branches; spikelets oblong or wedge-shaped, compressed, breaking up at maturity beneath the lemmas; many remain undispersed on the culm until this falls. Spikelet number variable from around 20 to 200. Glumes persistent, pointed, sometimes with a short awn, and thin membraneous margins; silvery-green or purplish, glabrous to densely hairy; lower usually rather shorter than, but sometimes equal to the upper glume; upper glume oblong or elliptic-oblong, 4–5.5 mm long, 3-nerved; lower narrowly oblong, 1-nerved, 3–4.5 mm long. Lemmas pointed, keeled upwards, sometimes with a short awn, silvery-green or purplish, 3–5.5 mm long; glabrous to minutely hairy on keels and near tips, 3-nerved. Paleas as long as or slightly shorter than the lemmas, 3–5 mm long, 2-keeled, colourless and translucent; lodicules to 1 mm long and 2-toothed. Grain 2.5–3 mm, enclosed within the hardened lemma and palea. Mean caryopsis oven-dry weight 0.32 mg. Root system is a dense mat of gingery-brown fibrous roots, largely unbranched in the upper parts, more strongly branched in the lower; lateral roots often very crinkly. Short wiry rhizomes 1–1.5 mm diameter and to 7 cm long are sometimes present. Native. This is a complex taxon with much variation as a result of polyploidy, and with many ecotypes. The division of the macrantha group in Flora Europaea into nine or more species is based entirely on morphology, and it is considered that these may represent only minor taxa. The species macrantha at the present time is not further subdivided. 46 noted that in Britain Koeleria glauca had been separated from K. macrantha and some Floras produced since then have listed the two species for Britain. Work currently being undertaken by the present author questions this distinction. The distribution of Koeleria macrantha in the British Isles is shown in Fig. 1. The species is widely distributed on Carboniferous, Magnesian and Jurassic limestones, and on chalk, extending from the north of Scotland, including the Orkney Isles, the Outer and Inner Hebrides, and the Isles of Skye, Mull and Arran, through northern Britain and Wales, the Isle of Man and Anglesey to the south of England. It is also recorded from the Isle of Wight, Jersey, Guernsey and Alderney. In Wales it is abundant on the Lias of south Dyfed and on the calcareous substrata of Clwyd and Anglesey, and less abundantly on calcareous substrata elsewhere (4). In the north of Britain it is more common in coastal regions. The species is well distributed in the west of Ireland, less so in the east. The distribution of Koeleria macrantha in the British Isles. (O) Pre-1950; (●) 1950 onwards. Each dot represents at least one record in a 10-km square of the National Grid. Mapped by Mrs J.M. Croft, Biological Records Centre, Institute of Terrestrial Ecology, mainly from records made by members of the Botanical Society of the British Isles. Koeleria macrantha is recorded throughout western, central and eastern Europe, but is noted only as a casual in Fennoscandia (Flora Europaea) although K. glauca occurs in Sweden (Skåne, Öland and Gotland) and Denmark. K. macrantha is absent from the Mediterranean islands of Sardinia, Sicily and Corsica, and the Azores; the species is present in Spain but not in Portugal. It occurs in the Baltic, Central, South-West, South-East and the Crimean divisions of Russia, being recorded from all regions in the Caucasus, Upper and Lower Volga, central and southern Urals, Carpathians, Moldavia, Black Sea, Lower Don and western and eastern Siberia to 128 °E, in the Tien Shan, Altai and Pamir mountains of the Central Asian states and in the southern parts of the Aral-Caspian area (14). 29 refer K. macrantha to the Eurosiberian Temperate floristic element. In Asia K. macrantha is recorded from Afghanistan, Iran, Lebanon, Syria, Turkey and Yemen, and in the Himalayan range from India, Kashmir, Nepal, Pakistan and Tibet. It is also recorded from the western Chinese provinces of Kansu, Shantung, Kiangsu and Hopeh, from the northern Chinese province of Shansi, from Japan, Korea, Manchuria and Mongolia. Koeleria macrantha is widespread in North America and is recorded from Arizona, Colorado, Kansas, Nebraska, North Dakota, Oregon, Utah, Washington and Wyoming (37); Louisiana, New Mexico and Mexico (30); California, Delaware, Indiana, Missouri, Ohio and Texas, and in Canada from Quebec to British Columbia (26). The species extends to 60 °N in Europe, to 62°N in Asia and to 55°N in North America. Koeleria macrantha is also recorded from central Africa – Ethiopia, Eritrea, Guinea and Cameroon, from the Transvaal and Natal areas of South Africa and from Swaziland (Vergl. Chor.). In England K. macrantha has an altitudinal range from sea level to 550 m (North Yorkshire), from sea level to 412 m in Scotland (eastern Highlands), from sea level to 457 m in Ireland (Donegal) (Alt. Range Brit. Pl.), and from sea level to around 185 m in Wales. It is found to 2700 m in the Alps (37), to 2200 m in the Pyrenees (45), to 2300 m in Turkey, to 2100 m in Afghanistan, to 3200 m in China and to 4400 m in Nepal. In North America the species is found to 3400 m and in Africa to 3700 m on Mount Kilimanjaro (Vergl. Chor.). Koeleria macrantha is exposed to average monthly maximum temperatures to around 39 °C in August in Spain, to around − 50 °C average monthly minimum temperatures in January in Siberia and to − 30 °C in January in European Russia (46), and to annual precipitation of around 2000 mm on the west coast of Scotland and in the Alps, and to conditions with no precipitation being recorded during July, August and September at Pagham in Afghanistan. No evidence in Britain of wind or insolation damage has been noted, nor from seaspray other than to make the leaves more glaucous. The species is subject to strong gales on northern coasts, such as on the islands of Canna and Sanday, Scotland (4) and to Föhn-swept slopes in Germany, and is found on steep dry slopes in the Jura Alps subject to strong insolation (14). Koeleria macrantha is found on both flat ground and steep slopes; it is common on slopes of 20–50 ° (Ecol. Atl.). In the British Isles the species shows no aspect preferences. In Europe it occurs rather less frequently on east-facing, and in Asia rather less frequently on west-facing slopes. It is a component of dry, semi-natural pastures, scree slopes, rock outcrops, old lead mines, limestone and quarry spoil (29) and also of limestone pavements. It occurs on sea cliffs and sand dunes, shelly or otherwise, particularly on partially fixed dunes, but is also present in the later successional closed turf, in both very short and tall vegetation to c. 60 cm; it is found frequently on coastal golf links and in the machair of the Outer Hebrides and Iona. The species is also found on pebble strands and shingle banks, on rocks present at the tops of beaches, and on sandy beaches where limestone runs into the sea, as in the Burren, Co. Clare; it is also recorded from limestone heaths here (37). The species has been recorded from heathy pastures in the Isle of Man, from peaty soil overlying limestone in Ireland (Bot. Irl.), from tall chalk heath at Lullington Heath, Sussex (30), from limestone heath in Somerset and Wales (26), and from sandy heath near Branderburg, near Lossiemouth, Scotland, where tufts of K. macrantha were growing up through the middle of mats of Calluna vulgaris and Empetrum nigrum. Koeleria macrantha is recorded from ungrazed, partially shaded moist soil overlying Magnesian limestone in Derbyshire (45), from dry woodland in Pakistan, under Juniper scrub in Nepal, in deep shaded ravines in Nepal, and from wet marshy land in Kashmir. It is also recorded from open woods in North America, and extensively from prairies (35). Koeleria macrantha is widespread on calcareous brown earths, brown earths and rendzinas overlying chalk and limestone in England, and is less widespread in Wales and Scotland. It is recorded from moderately base-rich brown forest soils in Scotland (28), from partially stabilized aeolian deposits in Scotland (9) and on almost pure sand, with few shell fragments, at many places around the coast of the British Isles. It is reported from gleys of moderate to high base status in Scotland (10), and from incipient podzols in England (7). The species is recorded from peaty or sandy peat substrata along the coasts of western Ireland (79), and from dry eskers in Offalay and Galway (Ir. Pfl.). It is found on soils derived from basalt, dolerite (4), serpentine (57), slate, hornfels and phyllite in Scotland, and has been observed by the author on a granite cliff, north of Aberdeen. It is found over serpentine in Cornwall, and on ‘sandstone cliffs’ in Hampshire (72). Soil analyses (methods according to Chem. Anal.) for sites supporting K. macrantha gave a range for exchangeable calcium and magnesium, extracted with M ammonium acetate (pH 9.0), from 130 to 6500 mg kg−1 and from 560 to 3880 mg kg−1, respectively, while potassium ranged from 110 to 540 mg kg−1. Phosphate phosphorus, extracted with 0.5 m sodium bicarbonate (pH 8.5), ranged from 1.2 to 32 mg kg−1, and total nitrogen from 0.028 to 1.10%, to a high 1.8% from a site on basalt. The low values for all elements were from sand dune sites. Koeleria macrantha is recorded as occurring in soils of pH between 4.5 and 8.0 and very rarely between pH 3.0 and 4.5 (Ecol. Atl.). On chalk it is found at pH values up to 8.4 (14). The species grows well in minimal soil depths on tops of walls, in limestone pavement hollows and in rock crevices, with only 2–5 cm soil, but it is also found on much deeper soils (to 60 cm +), particularly on sand dunes. In Continental Europe, in addition to soils overlying calcareous rocks, K. macrantha occurs in gritty sandy soil over porphyr (40) and over gypsum (47) in Germany, in soils over serpentine in Greece, with very high nickel and chromium contents – 4400 and 880 µg g−1, respectively (39), and over schists in the Czech Republic (54). The species is recorded from dry, acid soils in Germany (24), on sand dunes in Holland and north-west Germany, and in chalk-free, humus-rich surface soils over chalk also in north-west Germany (14). It is further recorded from zinc-rich soils, containing between 0.5% and 8.7% zinc, in the neighbourhood of Aachen, Germany and also from Belgium (14). Koeleria macrantha is recorded from gneiss and biotite muscovite in the Karakorum Mountains of north Pakistan (32). The leaf sheaths of K. macrantha are white, papery and moderately persistent, as are the leaves, decomposing slowly, and so adding to the litter layer. Koeleria macrantha is predominantly a species of calcareous grasslands and occurs with a high constancy in the following communities defined by the National Vegetation Classification (61, 62). CG1 Festuca ovina–Carlina vulgaris grassland is found on freely (often excessively) draining rendzina soils with a high base status, on steep and rocky, but stable, slopes on hard limestones, usually with a southerly to westerly aspect and so with a tendency to summer droughting. Often heavily grazed by sheep and rabbits, this community is confined to Devonian limestone in Devon and to Carboniferous limestone in Wales and the Mendips. Koeleria macrantha has an overall constancy of III in CG1 but attains a constancy of V in the subcommunities of Helianthemum canum and Festuca rubra–Scilla verna and a constancy of IV in the K. macrantha subcommunity. Constant species, as defined by the NVC, for CG1 and the following communities are given in Table 1. CG2 Festuca ovina–Avenula pratensis grassland consists of a rich mixture of grasses and dicotyledons in a closed sward and is traditionally grazed by sheep and rabbits. It occurs most frequently in relatively dry and warm lowland climates on free-draining calcareous soils derived from calcareous bedrocks; often prone to summer droughting. Koeleria macrantha has a constancy of V in this community. CG3 Bromus erectus grassland achieves maximum extent on lightly grazed or ungrazed grasslands over chalk and oolite in the south-eastern areas of Britain, and K. macrantha has a constancy of III in this community. CG4 Brachypodium pinnatum grassland is also mainly associated with lightly or ungrazed calcareous swards in south-eastern Britain, but is more generally found on the cooler and damper areas of the chalk and oolite than is the Bromus erectus grassland, which is more Continental in character. Koeleria macrantha has an overall constancy of only I in this community, but of III in the Avenula pratensis–Thymus praecox subcommunity. CG5 Bromus erectus–Brachypodium pinnatum grassland occurs where B. pinnatum is favoured by amelioration of extreme Continental conditions, but not so greatly as to exclude Bromus erectus. The community is most characteristic on the calcareous, base-rich soils of the oolite on the north-western fringe of southern, lowland limestone. Koeleria macrantha achieves a frequency of III in the typical subcommunity, but an overall frequency of II. CG6 Avenula pubescens grasslands are found on a variety of gently sloping lowland limestones, mainly in the south of England. The soils are deep and moist, though mostly free-draining alluvial rendzinas, or calcareous brown earths, and in some cases more mesophytic soils occur on flat limestones. This community is lightly grazed by cattle; on sloping limestones it is grazed by rabbits. Koeleria macrantha has a constancy of III in CG6. CG7 Festuca ovina–Hieracium pilosella–Thymus praecox/pulegioides grassland is the characteristic vegetation type of thin, stony soils over chalk, or less frequently on south-facing gentle slopes of Carboniferous limestone. The soils are dry and impoverished; this is reflected in the species composition, with grasses generally exhibiting reduced vigour and herbaceous dicotyledons playing a predominant role. The short, open nature of the vegetation is maintained by heavy grazing, mainly by rabbits. Koeleria macrantha has an overall constancy of III, a constancy of IV in the K. macrantha subcommunity and V in the Cladonia spp. subcommunity. CG8 Sesleria albicans–Scabiosa columbaria grassland is found only on free-draining calcareous, steep slopes of Magnesian limestone in Durham, principally on rendzinas rich in calcium and magnesium carbonates. The climate is cool and dry, and the community is a plagioclimax vegetation maintained by grazing of domestic animals and rabbits. Koeleria macrantha has a constancy of IV in this community. CG9 Sesleria albicans–Galium sterneri grassland is found in the northern Pennine submontane or montane climate, over shallow, freely draining but moist, calcareous lithomorphic soils on drift-free Carbonifeous limestone exposures. It forms an important part of upland farm hill-pasture and as such is frequently grazed, mainly by sheep. In CG9 K. macrantha achieves a constancy of IV. CG13 Dryas octopetala–Carex flacca heath occurs in the cool oceanic lowlands of north-west Scotland, on shallow rendzinas over the calcareous rocks of the Durness limestone, and on calcareous shell sand blown from the shore. The community is mainly ungrazed on the mainland but is quite heavily grazed by sheep and rabbits on Skye. Koeleria macrantha has a constancy of III in the Salix repens–Empetrum nigrum subcommunity and an overall constancy of II in CG13. Koeleria macrantha occurs with a lower constancy in the calcareous grassland community of Festuca ovina–Agrostis capillaris (CG10). The species is also found at a low constancy in the mesotrophic grassland community of Cynosurus cristatus–Centaurea nigra (MG5) and in the calcifuge sward of Festuca ovina–Agrostis capillaris–Rumex acetosella grassland (U1). Koeleria macrantha is also present with low constancy in the heathland community of Calluna vulgaris–Scilla verna (H7), and occurs with a higher constancy in: H6 Erica vagans–Ulex europaeus heath, a subshrub vegetation confined to the Lizard in Cornwall, on freely draining brown earth poor in calcium, but usually base-rich. The community survives most extensively over serpentine. Constant species are: Agrostis canina ssp. montana (V), Carex flacca (III), Erica cinerea (V), E. vagans (V), Filipendula vulgaris (IV), Ulex europaeus (V), U. galli (IV) and Viola riviniana (IV). Koeleria macrantha achieves a constancy of V in the Festuca ovina subcommunity and an overall constancy of II. The species is also recorded from several maritime communities. It is found with a constancy of IV in the Arenaria serpyllifolia–Sedum acre subcommunity and an overall constancy of III in the OV39 Asplenium trichomanes–Asplenium ruta-muraria crevice community, on limestone. Constant species for this community are Asplenium trichomanes (V), Asplenium ruta-muraria (V), Porella platyphilla (IV) and Homalothecium sericeum (IV). Koeleria macrantha occurs at low frequency in the following communities: OV 34 Allium schoenoprasum–Plantago maritima community, restricted to areas of seasonally damp soils over serpentine on the Lizard cliff margins; OV37 Festuca ovina–Minuartia verna in limestone areas contaminated with heavy metal spoil; MC5 Armeria maritima–Cerastium diffusum maritime therophyte community; MC9 Festuca rubra–Holcus lanatus cliff grassland; MC10 Festuca rubra–Plantago species maritime grassland; MC11 Festuca rubra–Daucus carota maritime grassland. Koeleria macrantha is also found at low constancy in the following dune communities: SD8 Festuca rubra–Galium verum fixed dune community; SD9 Ammophila arenaria–Arrhenatherum elatius dune grassland; SD10 Carex arenaria dune community and SD12 Carex arenaria–Festuca ovina–Agrostis capillaris dune grassland. Koeleria macrantha has also been recorded by the author from an extensive area of shingle mixed with sand (c. 2 m O.D.) in Suffolk. Accompanying species were Achillea millefolium, Artemisia maritima, Bromus hordeaceus ssp. hordeaceus,Catapodium marinum, Hordeum marinum, Lolium perenne, Lotus corniculatus, Ononis repens, Papaver rhoeas, Plantago coronopus, Puccinellia maritima and Reseda lutea. Koeleria macrantha occurred frequently in this community, and also in a community recorded from a raised beach (c. 3 m O.D.) also in Suffolk, with sparse, very short vegetation over almost pure sand, with very few shell fragments. The accompanying species here were: Agrostis capillaris, Dactylis glomerata, Elymus pycnanthus, Festuca ovina, Hordeum marinum, Hypochaeris radicata, Lolium perenne, Lotus corniculatus, Medicago lupulina, Plantago coronopus, P. lanceolata, P. maritima and Sedum acre. In Scotland, 9 recorded K. macrantha with a high constancy from two coastal dune communities. The Euphrasio–Festucetum arenariae association occurs on immature, brown calcareous soils over fixed dunes, with gentle to moderate slopes, particularly on the Caithness coast; the differential species are Bellis perennis, Euphrasia spp., Gentianella campestris, Ranunculus acris and Viola tricolor ssp. curtisii. The Astragalo–Festucetum association also occurs on fixed dunes with gentle to moderate slopes on the drier eastern coast of Scotland; the differential species are Astragalus danicus, Festuca ovina, F. rubra, Galium verum and Tortula ruralis. Koeleria macrantha has a constancy of IV in both these dune associations. 9 also described K. macrantha from an ultra-basic grassland community in Ayrshire. The Plantago maritima–Sieglingia decumbens community is found on freely drained, brown forest soils with a high magnesium content, derived from ultra-basic rocks. Species with a high constancy are Achillea millefolium, Festuca ovina, Trifolium repens, Viola riviniana and K. macrantha, with a constancy of IV. The species has been recorded by the author from a flat, sandy foreshore at Auchenmalg Bay, Galloway, in a tall (60 cm +), open community dominated by Raphanus maritimus. Koeleria macrantha occurred frequently in this community, which also included Anthyllis vulneraria, Arrhenatherum elatius, Cochlearia officinalis, Hieracium pilosella, Plantago lanceolata, Poa pratensis, Rumex acetosella, Tripleurospermum maritimum and Vicia hirsuta. In Ireland, 79 lists K. macrantha as a component of the Plantaginetum coronopo–maritima association. This dwarf coastal community, which is subject to frequent strong winds and sea-spray, is found on peaty or sandy-peat substrata on flat or not very steep cliff tops in western Ireland. The vegetation is heavily and persistently grazed by sheep and rabbits. On Malahide Island, north of Dublin, K. macrantha was recorded by 52 from the Festuco–Galietum maritimi association, which is found on middle dunes of grass-dominated, ungrazed or very lightly grazed, vegetation. Differential species are Anthoxanthum odoratum, Briza media, Dicranum scoparium, Euphrasia occidentalis, Koeleria macrantha, Potentilla sterilis, Pteridium aquilinum, Rosa pimpinellifolia, Thymus praecox and Trifolium repens. Koeleria macrantha is a component of the association Camptothecium–Asperuletum, characterized by Asperula cynanchica, Neotinea maculata and Gentiana verna, and described from stable, landward dunes in Galway (Ir. Pfl.) and also from Co. Clare by 37. The association Violo curtisii – Tortuletum ruraliformis occurs on dry stabilized coastal dunes and is characterized by Centaurium erythraea, Cerastium diffusum, Koeleria macrantha and Phleum arenarium. Originally described from Kerry and Co. Mayo (Ir. Pfl.) it has also been recorded from Co. Clare (37). Koeleria macrantha occurs in the Antennarietum hibernicae association (Ir. Pfl.) found in hollows between dunes on the coast of Co. Clare and also in dry eskers in Offalay and Galway. The soils have a high humus content and are not very calcium-rich. Characteristic species are Antennaria hibernica, Anthyllis vulneraria, Asperula cynanchica, Carex arenaria, C. caryophyllea, Carlina vulgaris, Danthonia decumbens, Festuca rubra, Gentiana campestris, Lotus corniculatus, Plantago lanceolata, Polygala vulgaris, Ranunculus bulbosus and Thymus pulegioides. This association was also recorded from Co. Clare by 65. 37 described K. macrantha from the Hyperico–Dryadetum association from the Burren, Co. Clare. This is a shrub-dominated community found on leached organic soil over bare limestone. Characteristic species are Calluna vulgaris, Danthonia decumbens, Dryas octopetala, Empetrum nigrum, Hylocomium splendens, Hypericum pulchrum, Hypnum cupressiforme var. ericetorum, Neckera crispa, Polygala vulgaris and Rosa pimpinellifolia. Koeleria macrantha is also found in the mesotrophic grassland Cynosurus cristatus–Centaurea nigra (MG5) which is the typical grassland of well-drained pastures over limestone, usually grazed by cattle in Ireland. In Continental Europe, K. macrantha is an order character of the Brometalia erecti and occurs in a number of associations belonging to this order throughout western Europe; these occur on dry steep slopes throughout the Alpine range, in dry rocky places in France and Belgium, the deeper moister soils of the Danube and Rhine valleys, and also on north-facing, more humid slopes of the Alps. Koeleria macrantha is also a component of some of the associations of the order Festucetalia valesiacae in various parts of Central Europe, including the Jura, Nahe Valley, Rheinhesse and the Bohemian Mountains. The species is also found in some associations of the class Arrhenatheretalia in the driest parts of the range. A range of communities containing K. macrantha, with a frequency of II and above, is given in Tables 2 and 3 to illustrate its geographical and ecological range. 15 measured shoot thrust in various herbaceous species and found that those species which were dominants in their usual habitats generated the greatest thrust. Koeleria macrantha with fine shoots demonstrated less thrust and much of this was directed horizontally through the canopy. They concluded that its fine aerial shoots might confer a high ability for ‘fine-scale’ foraging for light between the dominant species, a strategy in keeping with its usual intermediate status in lightly grazed or ungrazed grasslands. Koeleria macrantha also occupies an intermediate status in the prairies of the North American Midwest, and although its rapid recovery after severe droughts initially results in a monoculture, it is not a good competitor and is gradually relegated to its intermediate role by more vigorous species such as Andropogon spp., Poa pratensis and Stipa spp. (1). Koeleria macrantha exhibits a more dominant role in its coastal habitats on cliff tops, shingle and sand dunes where vegetation is often shorter, though often ungrazed; its flattened, tufted habit is an advantage here. 29 recorded K. macrantha as being notably absent from heavily disturbed situations and where there is intense competition, and 7 noted that it was rather less frequent on steep slopes in Derbyshire with very shallow soil and loose limestone fragments, where the dominant species was Festuca ovina, than on slopes with greater stability and less loose material and with Festuca ovina and F. rubra as co-dominants. Koeleria macrantha is classed as a good forage grass in North America (35), but 66 comments that it is considered of little agricultural value in Europe because it develops few leaves and these soon become tough. However, in long vegetation on deeper soils it tends to have longer, softer leaves than when growing on, for example, rock outcrops over shallow soils. 77 observed that K. macrantha increased in a sward over a period of 34 years, after being enclosed against domestic animals and rabbits. Other abundant species in the enclosure were Festuca ovina and Avenula pratensis which also increased; these are species also frequently found with K. macrantha in grazed calcareous grassland. Watt commented that these three species were much more vigorous inside, than outside, the enclosure. Nevertheless, K. macrantha is recorded as a component of a woodland replacement plagioclimax community in central Perthshire, Scotland, which is maintained principally by sheep grazing, but is also grazed by cattle, red deer, rabbits and hares (28), and the species has a high constancy (IV) in the NVC plagioclimax community of CG8 (Sesleria albicans–Scabiosa columbaria grassland) also maintained by the grazing of domestic animals and rabbits. However, K. macrantha does not seem able to withstand continual heavy grazing. 16 reported that the effects of overgrazing were to produce a decline in its abundance, vigour and productivity, and a simulated grazing experiment by the author, in which clipping was carried out at three height regimes, every 2 weeks for 10 weeks, resulted in the greatest production of shoot material when the plants were only lightly clipped (at 15 cm above soil level). An experiment carried out by 11 extending for 4 months during the summer with clipping every 14 days showed that K. macrantha increased the diameter of its root mass and stele as a result of the clipping, to a greater degree than Bromus inermis and Poa pratensis. 44 observed that burning stimulated inflorescence production in K. macrantha in the summer of the year following a spring burn, but by the following year no significant stimulus to flowering remained, while 16 noted that burning led to reduced yield of K. macrantha for a period of 3–5 years in Canadian prairies subject to moderate grazing. However, 1 reported that where fire was used as a means of weed control in the prairies of Nebraska, K. macrantha became temporarily widespread. Koeleria macrantha is very tolerant of trampling and has been noted, for example, at one site as being the only species present, providing almost 100% cover, apart from a small amount of Festuca rubra and Poa annua. It has been observed on many paths and tracks, particularly at coastal sites, but also on inland limestones, as very flattened, stiff little tufts, flowering on very short culms (see V (b)). Koeleria macrantha forms small tufts in cliff faces, on walls and in limestone pavement hollows. It is usually found as scattered individuals or more rarely with a number of tillers joined by short rhizomes, in short grazed and ungrazed turf, but on paths and tracks which are not too heavily trampled it often forms almost a monoculture, as it does sometimes on steep roadside banks, especially on sandy soils. In moderately tall vegetation to around 60 cm it is again usually found as scattered individuals of straggling appearance, rather than as tufts. Koeleria macrantha never produces very large tussocks and plants cultivated in a fertile garden soil produ