a survey of the vegetation of glen coe, scotland, summer 2002

C O M M I S S I O N E D R E P O R T

For further information on this report please contact:

Dominic SargentScottish Natural HeritageThe Governor’s House The ParadeFORT WILLIAMPH33 6BATelephone: 01397 704716E-mail: [email protected]

This report should be quoted as:

Averis, A.B.G. and Averis, A.M. (2006). A survey of the vegetation of Glen Coe, Scotland,Summer 2002. Scottish Natural Heritage Commissioned Report No.207 (ROAME No. F02LD04).

This report, or any part of it, should not be reproduced without the permission of Scottish Natural Heritage.This permission will not be withheld unreasonably. The views expressed by the author(s) of this report shouldnot be taken as the views and policies of Scottish Natural Heritage.

© Scottish Natural Heritage 2006

Commissioned Report No. 207

A survey of the vegetation ofGlen Coe, Scotland, Summer 2002

(ROAME No. F02LD04)

Background

Glen Coe is of both national and international importance for a range of features, one of which is itsvegetation. The area has varied geology and covers a broad altitudinal range, giving rise to a wide diversityof habitats, including grassland, heaths, tall herb ledge communities, high altitude flushes, and rock andscree. The site also supports notable assemblages of mosses, liverworts and lichens and the non-vascularand vascular plant assemblages include a number of Red Data Rock and Nationally Scarce species.

The survey, which was a co-operative venture between Scottish Natural Heritage and the National Trust forScotland, was required in order to provide: an up-to-date overview of the habitat types present, theirdistribution and their extent; and detailed information on the site’s botanical interests.

Main findings

● 129 types of vegetation and habitat, 275 vascular plant species and 243 bryophate species (of which43 are oceanic types) were identified in the survey. Glen Coe therefore has a great diversity ofvegetation and habitat types, including several which are uncommon in Britain or Europe, and a richflora which includes several rare species. These combine to make the site one of the most botanicallyinteresting places in the British uplands.

● Glen Coe’s oceanic climate results in very good bryophyte-rich heaths and woods and fine examples ofblanket bog. The presence of both acid and basic rocks produces good examples of acidophilouswoods, heaths, bogs and snow-beds; these being juxtaposed with some of the best examples ofbasiphilous vegetation outside the Breadalbane hills.

● Most of the uncommon vascular species known from the site were relocated and new populationestimates provided. However, little specific management can be done for the rare montane plants otherthan keeping sheep numbers sufficiently low.

A survey of the vegetation of

Glen Coe, Scotland, Summer 2002

Commissioned Report No. 207 (ROAME No. F02LD04)Contractor: Ben and Alison AverisYear of publication: 2006

C O M M I S S I O N E D R E P O R T

Summary

For further information on this project contact:Dominic Sargent, Scottish Natural Heritage, The Governor’s House, The Parade, Forth William PH33 6BA

Tel: 01397 704716

For further information on the SNH Research & Technical Support Programme contact:Senior Management Unit Advisory Services, Scottish Natural Heritage, Great Glen House, Leachkin Road, Inverness IV3 8NW

Tel: 01463 725000 or [email protected]

Executive summar y

This project was commissioned by Scottish Natural Heritage in order to collate existing information on thevegetation and flora of the Glen Coe Site of Special Scientific Interest (SSSI) and the Glencoe Special Areaof Conservation (SAC), to survey and map the vegetation and to assess the populations of rare vascularplant species. This site is about 10km SSE of Fort William in the western Highlands of Scotland. It coversabout 5000 hectares and is mountainous, with an altitudinal range of 30–1150m a.s.l. It is variedgeologically, including limestone and various igneous rocks. Much of the lower ground has a cover of peat.Streams, cliffs and screes are very common. The climate is cool and wet (oceanic). This report begins witha background study of previous work on the vegetation and flora of this site, followed by the results of oursurvey.

We did the fieldwork for this survey in the periods 4–14 June, 8–13 July and 21–28 July 2002. Wemapped the vegetation at the scale of 1:10,000 and classified it according to the National VegetationClassification, to sub-community level wherever possible. We recorded information about the vegetation (listsof species with an indication of the quantity of each) in representative quadrats measuring 2m x 2m in mostvegetation types. We made a list of all plant species found during the survey, indicating the approximatequantity of each species as ‘rare’, ‘uncommon’, ‘common’ or ‘abundant’. We mapped and described thelocations of species of special interest. We searched for each of the rare vascular species known here, andrecorded the location, habitat and population size where we found them.

We recorded 129 types of vegetation and habitat, 275 vascular plant species and 243 bryophytes in thissurvey. Each vegetation type is described in this report. We found most of the uncommon vascular plantspecies known from this site: Athyrium distentifolium, Carex capillaris, C. saxatilis, Cerastium alpinum, C. arcticum, Cystopteris montana, Draba norvegica, Dryas octopetala, Juncus biglumis, Pinus sylvestris, Poa alpina, P. glauca, Salix myrsinites, Saxifraga cernua, S. rivularis, S. nivalis and Sibbaldia procumbens.The Glen Coe survey area has a great diversity of vegetation and habitat types including several which areuncommon in Britain or Europe, and a rich flora including several rare species. These combine to make thesite one of the most botanically interesting places in the British uplands. The most notable habitats here are:

● Snow-beds (U11, U12, U14, U18 and Pohlia ludwigii types).

● Basic igneous crags with tall herb vegetation (U15/17), flushes (M11/12) and nationally importantpopulations of some rare montane plants (possibly the largest British populations of Saxifraga cernuaand S. rivularis).

● Bryophyte-rich heaths (H20c/H21b) with internationally important assemblages of northern, sub-montaneand montane oceanic liverworts.

● Rocky, bryophyte-rich woodland (W17) including one of the richest sites for oceanic bryophytes in Britain.

● Internationally important examples of western, oceanic blanket bog (M17 and M18 with M1 and M3bog pools).

● Nationally important limestone area on the steep NE and N slopes of Meall Mór, with herb-richgrassland (CG10, CG11, CG12, U4F, U5c and M25c), flushes (M10 and M11), tall herb vegetation(U15 and U17), woodland (W9), Dryas heath (CG14) and Salix myrsinites scrub (W20).

Scottish Natural Heritage Commissioned Report No. 207 (ROAME No. F02LD04)

The Glen Coe hills are grazed by sheep and red deer, and also some roe deer, wild goats, cattle andmountain hares. Different animal species graze in different parts of the site, and the intensity of grazing alsovaries. The effects of grazing on the vegetation and flora are discussed in this report.

This report ends with appendices containing: (1) quadrat data from representative 2m x 2m quadrats indifferent vegetation types; (2) a list of all plant species which we found during the survey; (3) ‘Target Notes’for locations which we found to be of special botanical interest; (4) maps; (5) photographs of a selection ofvegetation types and individual plant species.


View looking WSW across the River Coe to the Three Sisters

Contents

Summar y

Executive summar y

1 INTRODUCTION 11.1 The study area 11.2 Geology and geomorphology 11.3 Climate 21.4 Nomenclature 2

2 BACKGROUND STUDY 32.1 Vegetation 32.2 Flora 4

2.2.1 Vascular plants 42.2.2 Bryophytes and lichens 6

3 SURVEY METHODS 83.1 Vegetation 83.2 Flora 9

4 DESCRIPTIONS OF VEGETATION TYPES 10

5 EVALUATION 625.1 General description of the site 625.2 Vegetation 62

5.2.1 General patterns of vegetation 625.2.2 Wider British and European distributions of vegetation types 645.2.3 Uncommon vegetation types 64

5.2.3.1 Qualifying EC Habitats Directive Annex I Habitats 705.2.3.2 Non-qualifying EC Annex I Habitats 73

5.3 Flora 755.3.1 Phytogeography of the flora 755.3.2 Uncommon plant species 78

5.3.2.1 Nationally scarce and nationally rare vascular plants 78

5.3.2.2 Oceanic bryophyte species 835.3.2.3 Uncommon bryophyte species 83

5.4 Summary of the total botanical interest 905.5 Relationships between vegetation and environment 93

5.5.1 Geology 935.5.2 Climate 94

5.6 Land-use and management 975.6.1 History of vegetation, land-use and management 975.6.2 Grazing 985.6.3 Human views and recreation 100

5.7 Assessment of the Upland Vegetation Survey vegetation mapof Glen Coe 101

5.8 Extra note on birds and animals seen during this survey 102


6 ACKNOWLEDGMENTS 103

7 REFERENCES 104

Appendix 1 Quadrat data 107

Appendix 2 List of al l plant species found in this sur vey 232

Appendix 3 Target notes 248

Appendix 4 Maps 265

Appendix 5 Photographs 331

Note: Appendices 1–5 have not been reproduced within this

report but may be consulted by contacting the Nominated

Officer whose contact details are given on the title page.

List of tablesTable 1 List of vegetation types recorded in this survey at Glen Coe 2002 58

Table 2 British distribution of NVC types recorded in this survey at 67Glen Coe in 2002, and European distribution of similar vegetation

Table 3 Number of species in each phytogeographical group, and their 77percentage representation of the total flora recorded in this surveyat Glen Coe in 2002

Table 4 Plant species recorded in this survey at Glen Coe in 2002 which 84are uncommon at least in most parts of Great Britain

Table 5 Locations where nationally rare and nationally scarce vascular plant 85species were found in this survey at Glen Coe in 2002

Table 6 Occurrence of vegetation types in relation to broad rock and soil 95type (limestone, basic igneous rock and acidic rock)


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1 INTRODUCTION

1.1 The study area

This project was commissioned by Scottish Natural Heritage. The purpose of the project was to collateexisting information on the vegetation and flora of the Glen Coe Site of Special Scientific Interest (SSSI) andthe Glencoe Special Area of Conservation (SAC), to survey and map the vegetation and to assess thepopulations of rare vascular plant species.

The site takes in the range of hills on the southern side of Glen Coe, from Buachaille Etive Mór (Photograph1) in the east to Meall Mór in the west (Photograph 2), together with part of the Aonach Eagach ridge onthe north side of the glen (Photograph 5). In the south-east the site boundary follows the River Etive as far asDalness, then runs north-west up Gleann Fhaolain, over the watershed and down the Fionn Ghleann to itsjunction with the Allt na Muidhe. Further west, it encircles Meall Mór. In the north the boundary follows theRiver Coupar and the main A82(T) road as far west as Achtriochtan, where it cuts north up to the summit ofMeall Dearg, runs along the crest of Aonach Eagach to Sgorr nam Fiannaidh, then drops down theClachaig gully to the Clachaig Inn. It then follows the River Coe as far as the northern tip of Meall Mór. The site measures about 15km from east to west and 5km from north to south.

The survey area covers about 5000ha. It consists of four well-defined parts: the conical hill of Meall Mór;the long ridge of Bidean nam Bian, its satellite peaks An t-Sron, Stob Coire nam Beith, Stob Coire nanLochan and Stob Coire Sgreamhach and the subsidiary ridges of Aonach Dubh, Gearr Aonach and BeinnFada (known as the Three Sisters – see photograph at end of Executive Summary); the precipitous slopes ofSgorr nam Fiannaidh and the Aonach Eagach; and the isolated ridges of Buachaille Etive Beag andBuachaille Etive Mór. The hills are steep-sided and rise from deep glens to narrow ridges and slender summits.The altitudinal range of the study area is from around 30m a.s.l. in the bottom of the glen to 1150m a.s.l.at the summit of Bidean nam Bian.

There are two farms in the glen – Achnambeithach and Achtriochtan. The land is grazed by sheep and reddeer and is popular with walkers and climbers.

1.2 Geology and geomorphology

The geology is complicated and varied. There is Dalradian limestone at the western end of the site andgranite at the eastern end. In between the two are exposures of Dalradian quartzite, phyllite, graphitic schist,slate and mica-schist, rhyolite, andesite and basaltic lavas and tuffs, Triassic sedimentary rocks and intrusionsof diorite and porphyrite (British Geological Survey 1979). On many cliffs the strata run vertically and theeroded dykes of softer material result in spectacular gullies and scree-slopes, separating immense buttressesand sensational jagged pinnacles.

Glen Coe is the type locality for the geomorphological phenomenon of Cauldron Subsidence. As a result ofa circular fault-line, a circular block of the earth’s crust sank down into the molten rock below it. Magma wassqueezed up around the subsiding block to form the sides of the cauldron. The whole area shows signs ofintense glacial activity and there are textbook examples of knife-edged ridges, pyramidal peaks, ice-erodedhollows or corries, truncated spurs, hanging valleys and classic U-shaped, over-deepened glens.


1.3 Climate

The climate is cool and oceanic, with cool summers and winters, heavy and frequent rain, and frequent windand low cloud. The following climate data were compiled in the 20th century. Average annual rainfall isover 250cm, and over 380cm on the high ground (Meteorological Office 1952), making this one of thewettest parts of Britain. There are on average at least 220 wet-days (1mm or more of rain) a year (Ratcliffe1968). Winters are cool, with a January–February mean daily temperature at sea-level of about 3.0–3.5°C(Meteorological Office 1975). Summers are fairly cool, with a July–August mean daily temperature at sea-level of about 14°C (Meteorological Office 1975). The annual temperature range is small by general Britishstandards. Snow falls on low ground on an average of about 20–25 days a year (Meteorological Office1952) and lies for an average of 30–40 days per year (Page 1982). Obviously there is more snow athigher altitudes and snow showers can occur in any month. Although there is little high plateau to gathersnow, the precipitation is so great and the hills so high that considerable drifts of snow – snow-beds –accumulate each winter in the high corries and in hollows on the ridges. These snow-beds persist well intosummer in most years and can lie until August. The mean annual windspeed is about 12–13mph(Meteorological Office 1952). Cloud obscures bright sunshine for an average of about 75% or more of theday (Page 1982).

Rain is moderately acidic by general Scottish standards (Fowler et al. 1982, Philips Modern School Atlas1992) and nitrate deposition has been found to be at a moderate concentration of about 0.6g N m2

(Pitcairn & Fowler 1989). Sulphur dioxide pollution in 1987 was estimated to be negligible (Hill et al. 1991).

1.4 Nomenclature

Nomenclature in this report follows Stace (1997) for vascular plants, Blockeel & Long (1998) for bryophytes,and Purvis et al. (1992) for lichens. Place names are those on the Ordnance Survey 1:10 000 maps.

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2 BACKGROUND STUDY

2.1 Vegetation

Donald McVean visited the Glen Coe hills in the late 1950s as part of his work with Derek Ratcliffe on thevegetation of the Scottish Highlands. Several quadrat samples from the site appear in McVean & Ratcliffe(1962), the resulting monograph. He recorded Vaccinium-Luzula vegetation (NVC U16) with montaneoceanic liverworts in it on Stob Coire Sgreamhach, Cryptogramma-Athyrium snow-bed U18 in Coire namBeitheach, Alchemilla-Sibbaldia vegetation U14 (no location was given), and Dryas heath CG14 andCarex-Saxifraga mire M11 on Meall Mór. Derek Ratcliffe visited the area in 1961 as part of his study onperegrines, and must have found time to look at the vegetation, for he wrote to the Nature Conservancy in1962 to say that Bidean nam Bian was worthy of SSSI status. He wrote a further eulogy in the NatureConservation Review (1977):

This mountain (1141m), forming the southern side of Glen Coe, is scenically magnificent andgeomorphologically classic, with abrupt truncated spurs separated by hanging valleys which drainfrom the high corries. It is formed largely of lavas and agglomerates of Old Red Sandstone age, butthey vary greatly in lithology. The great cliffs of these spurs are of hard, acidic rhyolite and are muchfrequented by climbers, but there are numerous outcrops of calcareous andesite, which give a richflora. Some of the lower ledges and ravine sides have vestiges of birch and rowan wood, but all themore accessible lower slopes have been heavily grazed by sheep and are covered with grasslandmixtures of Festuca-Agrostis and Nardus. Locally there is much bracken and Trichophorum, Molinia,Sphagnum and Potentilla erecta on damper ground. Calluna and Vaccinium myrtillus are generallypoor but grow luxuriantly on ungrazed ledges. In places there is species-rich Festuca-Agrostisgrassland, and the ‘Lost Valley’ has much Festuca-Thymus grassland with Racomitrium canescens onalluvial soils.

Higher on the slopes this submontane complex passes to good Vaccinium-Empetrum heath, and on theridges of the Three Sisters there is a great deal of the western variant of this type, in which the dwarfshrubs share dominance with Nardus and Racomitrium lanuginosum. Steep ground has the alpineFestuca-Vaccinium grass heath with Alchemilla alpina and Racomitrium, and this merges with patchyRacomitrium heath on the upper ridges. The summit areas are too small to have an extensivedevelopment of this last community.

The high corries have a good deal of species-poor Deschampsia cespitosa grassland, and there areboth Nardus and Nardus-Trichophorum snow-bed grasslands in more definite hollows. Other late snowcommunities include the Cryptogramma-Athyrium, Sibbaldia-Alchemilla, Rhytidiadelphus-Deschampsiaand Racomitrium-Kiaeria starkei types and in the highest corrie, rocky hollows and gullies have muchmixed bryophyte-dominated vegetation from dry to spring and flush types.

The calcareous cliffs, which occur from near the foot of the mountain to the actual summit, have a richmontane flora with most of the characteristic calcicoles of Breadalbane mica-schist hills, together witha number of rare and local species, notably Saxifraga cernua, S. rivularis, S. nivalis, Draba norvegica,Cystopteris montana, Juncus biglumis, Cerastium arcticum, C. alpinum and Poa alpina. Many of themore common species are in profusion and tall herb communities are well represented. Deschampsiaalpina is common on both acidic and basic rocks and many calcifuge alpines are abundant. From itswestern position this mountain has a large number of oceanic bryophytes, and some of the lowerledges have good fragments of the northern Atlantic liverwort community so widespread in the north-west Highlands. The deep, block-filled stream ravine of the Allt Coire Gabhail provides a wide rangeof bryophyte and fern habitats, and there are fine growths of Hymenophyllum wilsonii.

The adjacent but much lower hill Meall Mór (675m) has extensive outcrops of Dalradian limestone,with an abundance of calcicoles including Dryas, Salix myrsinites and Thalictrum minus, and there isa large area of rich grassland. These two hills are complementary ecologically and have been treatedas a single area.

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The vegetation of the Bidean nam Bian SSSI was surveyed by the Nature Conservancy Council’s UplandSurvey Team in 1981 and 1982. A further visit was made in 1987 to check the map and to add NVCmire communities. A vegetation map was produced as a series of transparent overlays to black-and-whitevertical aerial photogrpahs at a nominal scale of 1:25,000. No site description was ever written, but thedescription from the Nature Conservation Review (Ratcliffe 1977) was included in the file of maps andphotographs. The Upland Survey team, working before the National Vegetation Classification (NVC) wasproduced, used the simpler classification devised by Birks & Ratcliffe (1980). They found 29 types ofvegetation. These could be equated with up to 53 NVC communities or sub-communities.

Fenton (1991) drew up a 1:25,000 Phase One habitat map of the Glen Coe estate, based on the NCCmap described above, and his own field survey work in 1990–1991.

Wanda Fojt, working for the NCC Peatlands team, visited two mire sites in Glen Coe – one to the east ofLoch Atriochtan and one on the north-facing slope of Aonach Dubh. She recorded Carex-Pinguicula miresM10, Carex rostrata-Sphagnum mires M4 and herb-rich Juncus-Galium mires M23.

Robinson (1994) surveyed the structure and condition of several blocks of woodland in this area, makingbrief notes on ground flora and also making an assessment of current and potential tree and shrubregeneration of each surveyed block of woodland.

The National Trust for Scotland (1998) produced a Grazing Management Plan for the Glencoe SpecialArea of Conservation. This was done as part of the European Union ‘LIFE’ Upland Grazing Project, in whichthe NTS worked in partnership with the Institute of Terrestrial Ecology, Scottish Natural Heritage, MacaulayLand Use Research Institute and Scottish Wildlife Trust. The report contains assessments of the present andpotential effects of grazing on the different EC Annex I habitats within the Glencoe Special Area ofConservation (SAC). It identifies a potential difficulty in that the ideal intensity of grazing is moderate in the species-rich grasslands but lower in the other Annex I habitats. It proposes all-year grazing by 500 Scottish Blackface ewes in the species-rich grasslands on Meall Mór, exclosure of part of the upperslope on Meall Mór, and removal of sheep from higher-altitude areas with Annex I habitats in the mainBidean ridge area of the site and from low-altitude wet heath between the River Coe and the main A82(T)road. It also suggests summer grazing by cattle in the Lairig Eilde to reduce tussocky Molinia caerulea.

2.2 Flora

2.2.1 Vascular plants

Compared with Ben Lawers and the Angus glens, the botanical treasures of Glen Coe seem to have beendiscovered relatively late. For example, Saxifraga cernua was not recorded here until 1949 (Lusby & Wright1996), although it had been known from Ben Lawers since 1790 (Hooker 1821). However, the study areais now well-known for its rare plants of which there is a notable concentration in the upper part of the easternarm of Coire nam Beitheach, where base-rich andesite rocks are exposed at over 900m.

Many botanists have visited the upper eastern part of Coire nam Beitheach in search of rare montane plants,and have recorded their results in varying detail. Those who have left records are E Greig Hall in 1972,Hugh MacAllister’s party in 1974, Paul Harrold in 1977, David Horsfield in 1981 and 1987, Lynne Farrell

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in 1985, Ro Scott in 1986, Sandy Payne in 1990, Shepherd & Gillespie in 1991, and Viv Halcrow in1995. The Botanical Society of the British Isles (BSBI) visited Coire nam Beitheach on a field meeting in 1986,and the local recorders for the BSBI, Alf Slack and Bernard Thompson, have made numerous visits. DonaldMcVean visited the Glen Coe area in the mid to late 1950s and recorded quadrat samples in a number ofvegetation types, including the Dryas octopetala vegetation on Meall Mór, as part of a more extensivesurvey of Scottish upland vegetation (McVean & Ratcliffe 1962). In 1986, Phil Lusby, who was then workingon a PhD on the taxonomy of Cerastium arcticum, visited the lower parts of Coire nam Beitheach in orderto collect samples of this species that had been washed down the gullies by spating streams. Halcrow(1996) undertook a survey of several uncommon vascular plant species known to occur in Coire namBeitheach and on the NE slope of Meall Mór, and made suggestions for monitoring their populations.

Three Nationally Rare vascular plant species – recorded in fewer than 16 10km squares in Great Britainsince 1950 – have been recorded in the study area. These are Saxifraga cernua, S. rivularis and Carexlachenalii.

Saxifraga cernua is now known from only three sites in Britain: Ben Nevis, Bidean nam Bian and Ben Lawers.On Ben Lawers it has been much reduced by collecting and is now thought to be so endangered there thatstocks are being maintained in cultivation at the Royal Botanic Garden in Edinburgh. The species is an arctic-alpine and the Scottish hills are at the far western edge of its range. The climate is probably no longer idealand its survival must be considered in jeopardy. S. cernua flowers occasionally and the flowers are visitedby insects but it never sets fruit. Instead, it reproduces by the small red bulbils which are carried in the axilsof the leaves. This, however, means that because there is no sexual reproduction and genetic recombination,the offspring are genetically identical to the parent and this lack of diversity may make it difficult for the plantsto survive environmental changes.

Saxifraga rivularis is somewhat less rare. It is known from Bidean nam Bian and Ben Nevis together with a scattering of localities in the north-west Highlands and in the central and eastern Highlands. There are 18 recent 10km-square records. Unlike S. cernua, this species does set viable seed. It, too, is far morecommon in the hills of Scandinavia and the Alps. Seeds of Saxifraga rivularis were collected by a memberof staff of the Royal Botanic Garden, Edinburgh, in 1997 in order to maintain a seed-bank for the speciesat the Royal Botanic Gardens, Kew (Phil Lusby, personal communication).

In Glen Coe Saxifraga rivularis has been reported to be common among boulders and along the sides ofsprings and rills on the floor of the eastern arm of Coire nam Beitheach. It is also known from rock ledgesaround Church Door Buttress. S. cernua also occurs in this area on a ledge known as The Terrace whichruns east from a conspicuous gully called The Gash. In 1985 Lynne Farrell, the NCC expert on vascularplants, went up into upper Coire nam Beitheach with the local plant recorder Alf Slack. They found S. rivularis and S. cernua and estimated that the corrie held the largest British populations of both plants.

One of the most notable things about the records of the two rare saxifrages is how inconsistent they are. Notwo recorders seem to have found the same numbers of plants in the same places. This may be because thepopulations fluctuate and move. It may be that much of the work was done in bad weather, when navigatingwas difficult and plotting accurate locations impossible. It is not clear. The most unambiguous records arethose of Halcrow (1996), who marked her findings very clearly on a set of coloured photographs.

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Carex lachenalii, while less spectacular than the saxifrages, is also very rare. A species of high montanemires and wet, earthy gullies which are often snow-covered late into spring, it is short and inconspicuous.There are seven recent 10km-square records in the eastern Highlands and around the Ben Nevis range. It has been recorded on rock ledges on the ledges of the grassy and rocky slopes below The Terrace in theupper eastern part of Coire nam Beitheach but it has not been seen for some years (Halcrow 1996).

Several Nationally Scarce vascular plant species – recorded in 16–100 10km squares in Great Britain since1950 – have been recorded in the study area.

Athyrium distentifolium, Carex atrata, C. saxatilis, Cerastium alpinum, C. arcticum, Cystopteris montana,Draba norvegica, Juncus biglumis, J. castaneus, Poa alpina, P. glauca, Saxifraga nivalis, Sibbaldia procumbensand Veronica alpina have been found in Coire nam Beitheach. Carex atrata has not been seen since 1986when it was found by the BSBI on the crags of Stob Coire nam Beith (Halcrow 1996). J. castaneus was lastseen in 1995 (Halcrow 1996), when only two plants were seen in flushes in Coire nam Beitheach, and thespecies was not seen in its other recorded site on the crags of Stob Coire nam Beith. Veronica alpina wasrecorded in 1987 by Dave Horsfield, but no location was given. Sibbaldia procumbens has been recordedin good quantity on the northern side of the main Bidean ridge (e.g. Halcrow 1996).

Carex capillaris, Dryas octopetala, Potentilla crantzii and Salix myrsinites have been recorded on andaround limestone outcrops on the steep upper north-eastern slope of Meall Mór. They were all recorded hereas recently as 1995 (Halcrow 1996). C. capillaris was also found more widely on the north-eastern slopeof the hill (Halcrow 1996).

Salix myrsinites, an uncommon montane willow, has been the subject of debate as all of the montane willowsand the montane willow scrub community are rare in Britain. Like many of our montane species they aremore widespread on the European mainland where the winters are colder. They are at the edge of theirrange in Britain and are assumed to be relicts from a time when the climate was colder and drier than it istoday. Except in Glen Clova in the eastern Highlands – where the climate is more continental and more likethat of Scandinavia and the Alps than it is in the west – the willows tend to grow as isolated bushes or smallstands of scrub. The plants are dwarfed. They are palatable – especially the male plants which arepreferentially browsed – and are almost always confined to inaccessible ledges and steep ground wherethey are vulnerable to rock-falls and soil erosion. Halcrow (1996) saw no catkins on S. myrsinites on MeallMór and considered the species unlikely to spread because there was too much grazing here. Accordinglythe upper eastern face of the hill, where the willows occur, was fenced against sheep in 2000 with the aidof a grant from the Millenium Forest Trust. James Fenton suggested around that time (letter in SNH files) thatthere was likely to be a problem because although the willows are sensitive to grazing they also needpatches of bare ground in which to establish from seed. It is unfortunate that by excluding grazing thesurrounding vegetation might grow so tall and thick that the willows are unable to spread.

Pinus sylvestris, which is scarce as a native species, occurs in woodland along the river and in ravines inGlen Etive. There are three trees in the Clachaig gully and a single tree at the Meeting of Three Waters(Halcrow 1996).

2.2.2 Br yophytes and lichens

In 1954 the British Bryological Society visited Coire nam Beitheach and ascended Bidean nam Bian.Among their finds was the scarce oceanic liverwort Anastrophyllum donnianum with perianths (female sexual

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organs), a very rare sight indeed. They also found good populations of the snow-bed rarities Moerkia blyttii,Diplophyllum taxifolium, Lescuraea patens, Brachythecium glaciale, Mnium thomsonii, Marsupella boeckiiand Isopterygium muellerianum.

The British Bryological Society visited the Lost Valley and its boulder-field in 1986 and made a respectablelist of species.

Ben Averis visited the Lost Valley ravine and the woods on Gearr Aonach in 1988 and found a rich floraincluding several uncommon western species: the mosses Dicranodontium uncinatum, Glyphomitrium daviesii,Isothecium holtii, Paraleptodontium recurvifolium, Racomitrium ellipticum, Rhabdoweisia crenulata andTrichostomum hibernicum, and the liverworts Anastrepta orcadensis, Aphanolejeunea microscopica, Colura calyptrifolia, Douinia ovata, Frullania teneriffae, Gymnomitrion crenulatum, Harpalejeunea molleri,Herbertus aduncus, H. stramineus, Lepidozia pearsonii, Leptoscyphus cuneifolius, Mastigophora woodsii,Metzgeria leptoneura, Plagiochila carringtonii, P. exigua, Pleurozia purpurea, Radula aquilegia, Scapaniaornithopodioides and Sphenolobopsis pearsonii. This was part of a larger survey of bryophytes in westHighland woods (Averis 1991).

Alison Averis visited the woods on Gearr Aonach and in the Lost Valley boulder-field in 1992 and foundgood examples of the ‘Northern Atlantic Hepatic Mat’ community including Herbertus aduncus, Scapaniagracilis, S. ornithopodioides, Bazzania tricrenata, B. pearsonii, Plagiochila spinulosa, Pleurozia purpurea,Mastigophora woodsii, Mylia taylorii, Anastrepta orcadensis, Lepidozia pearsonii and, more unusually, thelow-altitude species Bazzania trilobata and Saccogyna viticulosa. She also found small patches of hepaticmat vegetation around rock outcrops on the northern side of Buachaille Etive Mór. This work was part of aPhD study (Averis 1994).

Fryday (1993) studied the lichens on the scree cones and fans in Glen Coe. These screes are some of thelargest in Scotland and a large number are packed into a small area. The lichen flora suggests that therehas been much reworking and re-distribution of the scree in the last 300 years or so, as well as movementin the recent past. The screes are derived from eroding igneous dykes.

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3 SURVEY METHODS

We did the fieldwork for this survey in the periods 4–14 June, 8–13 July and 21–28 July 2002. Work inthe first of these three periods was considerably hampered by low cloud and heavy rain, and by the amountof snow that was still lying in the upper corries.

3.1 Vegetation

We classified the vegetation according to the National Vegetation Classification (NVC) (Rodwell 1991a,1991b, 1992, 1995, 2000), to sub-community level wherever possible. We classified some inaccessiblepatches on cliffs to community level only. It is generally not possible to delineate individual patches of singletypes of vegetation on a map at the scale of 1:10 000. It is, in any case, rare to find large areas of groundcovered by uniform swards of a single type of vegetation. It is most usual to find mosaics of two or moretypes – such as pools and wet channels within patches of blanket bog or small patches of grassland orbracken within areas of heathland. The mapped units on our vegetation maps represent mixtures orassemblages of several types of vegetation which occur consistently in a particular area, usually defined bythe topography or by the aspect of the slope. We labelled each mapped unit with the types of vegetationrecorded there, and showed the relative contribution of each type as an estimated percentage. It is hard todefine how accurate the mapped vegetation boundaries are. They are generally most accurate (within10–20m or so) for small or well-defined patches of vegetation whose edges can be most clearly related tofeatures such as streams, cliffs and roads on the Ordnance Survey map. The precise boundaries of the mixedmosaics of vegetation on a larger scale are less obvious in the field and therefore less easy to define: itwould be meaningless to give an estimated figure of accuracy for these boundaries.

We classed vegetation floristically intermediate between two NVC types as such: NVCx–NVCy.

Bad weather and the remote and difficult ground meant that mapping vegetation had to take priority overrecording quadrats, but we sampled most vegetation types by sets of representative quadrats. The word‘quadrat’ is used by botanists to mean a square-shaped area of ground defined for the purpose of recordinginformation about the vegetation. This square area may be marked (e.g. with wooden pegs) so that it canbe visited again in future to observe changes in the vegetation, or, as in the Glen Coe survey, it may beunmarked and defined only temporarily for the purpose of recording information about a typical orrepresentative example of a particular type of vegetation.

We recorded details about the vegetation in quadrats measuring 2m x 2m, except for the canopy layer(trees and shrubs) in woodland, for which we used a quadrat size of 10m x 10m (surrounding one or more2m x 2m quadrats of ground vegetation). Within each quadrat we recorded the approximate quantity ofthe aerial parts of each plant species seen, using the DOMIN scale:

1 = <4% cover (few individuals) 6 = 26–33% cover2 = <4% cover (several individuals) 7 = 34–50% cover3 = <4% cover (many individuals) 8 = 51–75% cover4 = 4–10% cover 9 = 76–90% cover5 = 11–25% cover 10 = 91–100% cover

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We recorded the location of each quadrat as an 8-figure Ordnance Survey grid reference. We alsorecorded the altitude (in metres above sea-level), slope aspect (N, NE, E etc.), slope gradient (gentle,moderate, steep etc.) and approximate vegetation height (in centimetres) for each quadrat. We aimed torecord at least five quadrats in each vegetation type, but difficulties of physical access combined with thelarge amount of time needed to map the vegetation and the very small extent of some vegetation types meantthat we could not record as many as five quadrats in many vegetation types. Tall-herb vegetation, willowscrub and Dryas octopetala heaths on cliffs were impossible to sample using quadrats because they wereinaccessible or were in tiny fragments covering less than a square metre. We sampled these types ofvegetation by making a list of all the species which we could see from safe vantage points.

3.2 Flora

We made a list of all plant species found during the survey. We also recorded the approximate quantity ofeach species as found in this survey. To do this we used the terms ‘rare’, ‘uncommon’, ‘common’ and‘abundant’. These equate with ‘rare’, ‘occasional’, ‘frequent’ and ‘abundant’, which are used by manybotanists. However, ‘occasional’ and ‘frequent’ are inappropriate for this purpose because they arespecifically to do with time and not quantity.

We mapped the locations of species of particular interest or other botanically noteworthy features, anddescribed them using ‘target notes’. We located as many as possible of the rare vascular species and madean estimate of population size for each one. These species are Saxifraga cernua, S. rivularis and Carexlachenallii, which are all nationally rare, and Cerastium arcticum, C. alpinum, Cystoperis montana, Poa glauca, P. alpina, Sibbaldia procumbens, Athyrium distentifolium, Draba norvegica, Juncus castaneus,J. biglumis, Saxifraga nivalis, Carex capillaris, C. saxatilis, C. atrata, Dryas octopetala, Salix myrsinites,Potentilla crantzii, Pinus sylvestris and Veronica alpina, all of which are nationally scarce. We used theannotated colour photographs prepared by Halcrow (1996) to locate the populations of the more notablespecies in upper Coire nam Beitheach.

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4 DESCRIPTIONS OF VEGETATION TYPES

We recorded 129 types of vegetation and habitat in this survey. These are listed in Table 1. Each of these vegetation and habitat types is described below. Table 2 shows the distribution of each NVC type inGreat Britain, and that of similar vegetation elsewhere in Europe. The quadrat data are given in Appendix 1.The 1:10,000 vegetation maps are in Appendix 4.

W3 Salix pentandra-Carex rostrata woodland

The small and inaccessible piece of woodland in the SE corner of Loch Achtriochtan, viewed throughbinoculars, appears to fit this NVC type. It has an open canopy of Salix cinerea growing in a deepwaterlogged sward of Carex rostrata, Juncus acutiflorus and tall herbs such as Lychnis flos-cuculi, Calthapalustris, Myosotis sp. (probably M. secunda) and Ranunculus flammula.

W3 occurs throughout the British uplands from north Wales northwards. It is widespread though rarelyextensive in the south-western Highands and in the Inner Hebrides.

W4 Betula pubescens-Molinia caerulea woodland

This is wet, grassy woodland with a canopy of birch and rowan or of Salix cinera and S. aurita. Under thetrees there is a green, grassy understorey of Molinia caerulea and other grasses such as Agrostis caninaand Holcus mollis. These are interleaved with ferns and herbs including Blechnum spicant, Dryopterisdilatata, Viola riviniana, Angelica sylvestris and a few trailing stems of Lonicera periclymenum. On the wetsoils there are spongy masses of bryophytes including Sphagnum palustre, S. fallax, Hylocomium splendens,Thuidium tamariscinum, Pleurozium schreberi, Hypnum jutlandicum and Scapania nemorea.

The W4 in Glen Coe does not fit well into any of the three sub-communities. Although in some patches thereare the ferns and honeysuckle of the Dryopteris-Rubus sub-community W4a, they are not really commonenough, and as this type of woodland occurs only in small patches it was classified to community level only.

W4 occurs in Glen Etive and in the Coire Gabhail woodland. It is common on wet soils throughout the northand west of Britain.

W7b Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland, Carex remota-Cirsiumpalustre sub-community

This is wet woodland with a canopy of alder, birch and hazel, and a short to moderately lush ground layercontaining many grasses and herbs. In the ground layer there are no clearly dominant species: thevegetation is a more even mix of many species such as Agrostis canina, Cynosurus cristatus, Deschampsiacespitosa, Holcus lanatus, Juncus acutiflorus, J. bulbosus, J. effusus, Lysimachia nemorum, Oxalis acetosella,Poa annua, Prunella vulgaris, Ranunculus acris, R. flammula, R. repens and Viola riviniana. Although thereare plenty of bryophytes in the ground layer they are not very conspicuous: the commonest species are themosses Brachythecium plumosum, Calliergonella cuspidata, Hypnum cupressiforme, Plagiomnium undulatum,Rhytidiadelphus squarrosus and Thuidium tamariscinum, and the liverwort Pellia epiphylla.

The vegetation is not a very clear fit for this sub-community, but it is nearer to W7b than to the other twosub-communities of W7. It occurs as small patches among grassy W11b woodland and drier, herb-rich

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W9b woodland on the steep north-eastern slope of Meall Mór. Much of the flora of the W7 woodland isshared with the associated W9 and W11 woodland types, and also with the nearby base-enrichedgrasslands and mires.

W7b is widespread at low altitudes in the British uplands and around the upland fringes.

W7c Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland, Deschampsia cespitosasub-community

This vegetation is quite different in both flora and habitat from the W7b described above. A canopy of mature, large alders grows over a dense, lush sward of Deschampsia cespitosa about 60–100cm tall.Other plants are few and sparse among the tall tussocks of D. cespitosa. Species seen here include Agrostis capillaris, Holcus lanatus, H. mollis, Juncus effusus, Ranunculus repens, Rumex acetosa, the mossesAtrichum undulatum, Eurhynchium praelongum, Rhytidiadelphus squarrosus and Thuidium tamariscinum, andthe liverwort Lophocolea bidentata. Ash seedlings occur very sparsely.

This vegetation is quite extensive on the flat valley floor near the River Coe, in the far north-west of the site.It is quite homogeneous; within the woodland the vegetation does not vary much (see Photograph 12). The woodland opens out here and there into glades with lush grassland and rush mire.

As with W7b, W7c is widespread at low altitudes in the British uplands and around the upland fringes.

W9b Fraxinus excelsior-Sorbus aucuparia-Mercurialis perennis woodland, Crepis paludosasub-community

This is woodland of damp, neutral or basic soils. The canopy is quite variable, with mixtures of birch, rowan,ash, hazel and alder. Under this is a rich array of grasses, ferns and tall mesotrophic herbs such asMercurialis perennis, Angelica sylvestris, Ranunculus acris, Crepis paludosa, Filipendula ulmaria, Valerianaofficinalis, Geranium sylvaticum, Anthoxanthum odoratum, Brachypodium sylvaticum, Dryopteris dilatata, D. filix-mas, Phegopteris connectilis and Oreopteris limbosperma. Among these are many smaller plants suchas Viola riviniana, Linum catharticum, Saxifraga aizoides, Selaginella selaginoides, Prunella vulgaris,Chrysosplenium oppositifolium, Primula vulgaris, Asplenium viride and A. trichomanes. There are mats andpatches of bryophytes including Hylocomium splendens, Ctenidium molluscum, Rhytidiadelphus triquetrus,Calliergonella cuspidata, Eurhynchium striatum, E. praelongum, Mnium stellare and Plagiomnium undulatumand, on rocks, Neckera crispa, Tortella tortuosa, Orthothecium rufescens, Anoectangium aestivum,Hymenostylium insigne and Preissia quadrata.

W9b is quite common on the rock outcrops and in the stream gullies on Meall Mór (see Photograph 11),where it forms mosaics with Quercus-Betula-Dicranum woodland W17. W9b also occurs in the Allt CoireGabhail ravine.

It is widespread on the more basic soils throughout the Highlands and the Inner Hebrides.

W11a Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Dr yopteris dilatatasub-community

There are small patches of W11a in the woods of the Allt Coire Gabhail ravine: on steep rocky ground outof the reach of grazing animals. Under a canopy of birch and rowan there is a green grassy ground layer

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made up of Holcus mollis, H. lanatus, Anthoxanthum odoratum and Agrostis capillaris. This is dotted withBlechnum spicant, Viola riviniana, Potentilla erecta, Galium saxatile, Oxalis acetosella and Primula vulgaris,set with clumps of Dryopteris filix-mas, D. dilatata, Oreopteris limbosperma and Phegopteris connectilis andentangled with long trails of Lonicera periclymenum. Among these plants are clumps and patches ofbryophytes such as Thuidium tamariscinum, Hylocomium splendens, Rhytidiadelphus squarrosus, R. loreus,Hypnum jutlandicum and Lophocolea bidentata.

W11a occurs only where the climate is not too cold and where there is little or no grazing. It is mostcommon in the south-west Highlands and in the uplands of England and Wales.

W11b Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Blechnum spicantsub-community

A dense canopy of birch, in some places with ash, alder or willows and an understorey of rowan, growsin a green and grassy underlayer. This can be overgrown with bracken, but is more typically like a short,well-grazed grassland consisting of mixtures of Agrostis capillaris, Anthoxanthum odoratum, Holcus mollis,Carex binervis and Luzula multiflora. It is speckled with small herbs such as Potentilla erecta, Galium saxatile,Prunella vulgaris, Primula vulgaris, Oxalis acetosella and Viola riviniana, and the fern Blechnum spicant.There are some thick swards of bluebells Hyacinthoides non-scripta, giving a smoky-blue haze of theirfragrant flowers under the trees in spring. There are plenty of bryophytes too: mostly large robust mossessuch as Hylocomium splendens, H. brevirostre, Thuidium tamariscinum, Rhytidiadelphus squarrosus, R. loreus, Polytrichum formosum and P. commune.

There are patches of W11b along the rocky banks of the River Coe, on steep rocky banks on Meall Mórand in the Allt Coire Gabhail ravine.

W11b is common throughout the north-western British uplands, where it is one of the most common types ofwoodland on well-drained light soils where there is moderate to heavy grazing.

W17a Quercus petraea-Betula pubescens-Dicranum majus woodland, Isothecium myosuroides-Diplophyllum albicans sub-community

This woodland has a canopy of birch. This canopy is quite thin and open on the cliffs, but in the ravinesand on larger ledges it is more dense, with a shrub layer of rowan. W17a has a dark, dense, heathyground layer of Calluna vulgaris with Vaccinium myrtillus and in some places Erica cinerea and E. tetralix.There is a sprinkling of small plants under the shrubs, including Blechnum spicant, Deschampsia flexuosa,Galium saxatile, Oxalis acetosella and Melampyrum pratense. A rich array of bryophytes grows inconspicuous mats and patches over the ground, over fallen tree-branches and rocks and the bases of trees,and clothing the living branches of the trees. Most of the species are large and common ones such as themosses Breutelia chrysocoma, Isothecium myosuroides, Rhytidiadelphus loreus, Pleurozium schreberi,Hylocomium splendens, H. umbratum, Hypnum jutlandicum, Dicranum scoparium, D. majus and Sphagnumquinquefarium and the liverworts Diplophyllum albicans, Scapania gracilis and Plagiochila spinulosa. There are also many more exacting western or oceanic plants such as the moss Dicranodontium uncinatumand the liverworts Herbertus aduncus (Photograph 48), Bazzania tricrenata, B. trilobata, Mylia taylorii,Anastrepta orcadensis, Lepidozia pearsonii, Harpelejeunea molleri, Lejeunea patens, Plagiochila punctata,Mastigophora woodsii and, at higher altitudes, the more montane Bazzania pearsonii, Scapaniaornithopodioides and Plagiochila carringtonii.

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W17a is widespread in the study area. Most of the woodland on the steep slopes and cliffs belongs to this sub-community, as does much of the woodland in the Allt Coire Gabhail ravine and boulder-field (see Photograph 28).

W17a is widespread in the western British uplands from south-west England northwards, though south of theHighlands it is generally poorer in bryophytes than here in Glen Coe.

W17b Quercus petraea-Betula pubescens-Dicranum majus woodland, Typical sub-community

This is woodland with a canopy of birch and rowan, and a heathy, lightly grazed ground layer with muchCalluna vulgaris and Vaccinium myrtillus. The canopy varies from fairly tall trees over 10m in height toyoung, scrubby examples only 4–5m tall. The ground vegetation is also variable in height: from 10–15cmto well over 50cm. Heather and blaeberry are the two commonest vascular species: together they form adwarf shrub layer covering about 30–90% of the ground surface. Mosses are very common, growingbeneath the dwarf shrubs and filling the spaces in between. The commonest moss species here areHylocomium splendens, Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphus loreus, Sphagnumquinquefarium and Thuidium tamariscinum. Other vascular species and bryophytes are generally sparse:they include Agrostis canina, Deschampsia flexuosa, Potentilla erecta, Dryopteris filix-mas, Pteridiumaquilinum, Empetrum nigrum, Melampyrum pratense, the mosses Ptilium cristacastrensis (Photograph 55),Plagiothecium undulatum, Dicranum majus, D. scoparium and Breutelia chrysocoma, and the liverwortsAnastrepta orcadensis and Barbilophozia floerkei.

W17b occurs in complex mosaics with other woodland types on steep to moderate slopes in lower CoireGabhail and along the nearby part of the River Coe. Soils in these places are evidently acidic and welldrained. W17b is widespread in the British uplands.

W17c Quercus petraea-Betula pubescens-Dicranum majus woodland, Anthoxanthum odoratum-Agrostis capil laris sub-community

This is similar to the W17b woodland described above, but is not as heathy. It has little or no Callunavulgaris, and only small amounts of Vaccinium myrtillus. The grasses Agrostis canina, Anthoxanthumodoratum and Deschampsia flexuosa form a patchy sward together with Potentilla erecta, Oxalis acetosella,Blechnum spicant and scattered bracken fronds. The vegetation therefore has a greener, more grazed andgrassy appearance than W17b. It looks rather like W11, but the moss carpets are much more extensiveand there are fewer dicotyledonous herbs. The moss carpets contain much the same range of species as dothose in W17b.

Small patches of W17c occur among W17a and W17b in the Allt Coire Gabhail-River Coe woodlands,and also in the small blocks of woodland along Glen Etive.

W17c is widespread in upland Britain.

W18d Pinus sylvestris-Hylocomium splendens woodland, Sphagnum capillifolium/quinquefariumsub-community

There are patches of this kind of native pine woodland in mosaic with birch woodland (W17a) on a steepslope on the north side of the River Etive, at the south-eastern edge of the survey area. The W18d here has

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a canopy of Pinus sylvestris with some birch and rowan. The woodland was too steep and inaccessible for detailed examination, but views through binoculars showed the ground vegetation to contain muchCalluna vulgaris, with Vaccinium myrtillus, Molinia caerulea, Deschampsia flexuosa, Agrostis sp. (probablyA. canina), Solidago virgaurea and mosses including Sphagnum capillifolium. It is this last species, togetherwith Molinia, which places the vegetation in this sub-community – the main type of damp native pinewoodland in the Highlands, and the commonest pinewood type in the west. A fence has been erectedaround this W17a/W18d birch-pine woodland in Glen Etive. The exclosure is now ungrazed. It includessome open wet heath just upslope of the pinewood: there is young birch and pine regeneration here.

W20 Salix lapponum-Luzula sylvatica scrub

The patches of montane willow scrub on the crags on the upper slopes of Meall Mór are no more thanfragments; tiny patches; all that remains of what once, in a cooler and drier climate, might have been alarge area of scrub. Most of the willows are just isolated bushes. Some are surrounded by a few smallerindividuals, and there are some denser thickets on the larger crags. Nearly all are inaccessible. A fewbushes on the lower ledges were reached by determined scrambling, but most of the crags, including thosewith the larger clumps of willows, had to be viewed through binoculars from safe ground nearby. Thewillows cling to the dripping limestone ledges, growing in lush meadows of herbs and grasses such asDeschampsia cespitosa, Festuca vivipara, Anthoxanthum odoratum, Thymus polytrichus, Ranunculus acris,Valeriana officinalis, Luzula sylvatica, Sedum rosea, Saussurea alpina, Oxyria digyna, Saxifraga aizoides,S. oppositifolia, Silene acaulis, Selaginella selaginoides, Carex pulicaris, C. panicea, C. capillaris, C. flacca,C. hostiana, Geranium sylvaticum, Linum catharticum, Alchemilla alpina, A. glabra, Filipendula ulmaria,Dryas octopetala, Rubus saxatilis, Galium boreale and Thalictrum alpinum. Bryophytes cling to the rockcrevices and spill in hanging mats over the sheer, wet faces. Species include the calcicoles Tortella tortuosa,Ctenidium molluscum, Orthothecium rufescens, Neckera crispa, Ditrichum flexicaule and Anoectangiumaestivum as well as more widespread plants such as the mosses Hylocomium splendens, Rhytidiadelphustriquetrus, R. loreus, Hypnum jutlandicum and Breutelia chrysocoma and the liverworts Frullania tamarisci,Diplophyllum albicans, Pellia epiphylla and Scapania undulata.

Vegetation describable as willow scrub occupies barely 1% of the area in which the willows occur, andmany patches of willows could equally well be described as tall-herb ledge vegetation U17 or Dryas-Sileneheath CG14 with a few plants of Salix myrsinites growing in the sward.

W20 scrub is one of the least common types of vegetation in the British uplands. Most of the existingexamples are in the Breadalbane hills between Ben Lui in the west and Caenlochan in the east. There area few records in the central and north-west Highlands. Only on the limestone at Inchnadamph in Sutherlandand at Rassal in Wester Ross is there willow scrub dominated by Salix myrsinites and with no other speciesof willow. Elsewhere, S. myrsinites is a member of more mixed willow scrub where the canopy includes othermontane species such as S. lapponum, S. arbuscula and the rare S. lanata and S. reticulata.

W24 Rubus fruticosus agg.-Holcus lanatus underscrub

This vegetation consists of a dense tangle of brambles. There are patches of it among grassland and brackenin a small area near Achnacon in the lower part of the main glen. It appears to be at most only lightlygrazed. We did not examine it in detail: it appears to be species-poor and of little botanical interest. W24is widespread and common in lowland Britain, mainly on lightly grazed or ungrazed ground in places suchas roadsides and field margins.

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W25 Pteridium aquil inum-Rubus fruticosus agg. underscrub

This is tall bracken-dominated vegetation with much bramble. There are patches of it in mosaic withgrassland and bramble underscrub in a small area near Achnacon in the lower part of the main glen. Aswith the nearby W24 we did not examine the flora of this vegetaton in detail: it appears to be species-poorand of little botanical interest. W25 is widespread and common in lowland Britain.

W25D Pteridium aquil inum-Rubus fruticosus agg. underscrub, damp sub-community

This is a type of vegetation not yet described in the NVC. Quite common on suitable soils in the westernHighlands and Inner Hebrides, it also occurs in north Wales.

It is a community of damp, neutral to basic soils at low to moderate altitudes where grazing is only light,and is probably a relict of damp woodland. It occurs in the study area on the lower slopes of Sgorr namFiannaidh, mixed with the less herb-rich bracken community U20. From a distance it looks exactly like U20,and it can be appreciated only at close quarters. Under the cover of the fern there is an array of grasses andmesotrophic herbs: these include Festuca vivipara, Agrostis canina, Anthoxanthum odoratum, Deschampsiacespitosa, Filipendula ulmaria, Ranunculus acris, Cirsium heterophyllum, Valeriana officinalis, Geum rivale,Alchemilla glabra and smaller species such as Carex flacca, C. panicea, C. pulicaris and Linum catharticum.There are few bryophytes other than mats and wefts of species such as Hylocomium splendens, Scleropodiumpurum, Hypnum lacunosum, Calliergonella cuspidata, Breutelia chrysocoma and Frullania tamarisci.

H10a Calluna vulgaris-Erica cinerea heath, Typical sub-community

Most of the heather-dominated moorland on well-drained soils on steep slopes exposed to the sun is Calluna-Erica heath. The Typical sub-community is probably the least widespread form and occurs mostly in shelteredstream-gullies. It consists of dense, dark swards of Calluna vulgaris and Erica cinerea, speckled with Carexbinervis, Luzula multiflora, Deschampsia flexuosa, Galium saxatile and Potentilla erecta and with an underlayof bryophytes such as Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphus loreus, Hylocomiumsplendens and Frullania tamarisci.

H10a is the most common type of Calluna heath at low to moderate altitudes on well-drained soils in thewestern Highlands and the Inner Hebrides.

H10b Calluna vulgaris-Erica cinerea heath, Racomitrium lanuginosum sub-community

This is the most common type of dry Calluna heath in the study area. It is extensive on the east-facing sidesof Aonach Dubh, Gearr Aonach, Beinn Fhada, Buachaille Etive Beag and Buachaille Etive Mór, and on thesouth-facing slopes of Sgorr nam Fiannaidh and Aonach Eagach, giving way to the more upland, cold-tolerant Calluna-Vaccinium heath H12 (see below) with increasing altitude. The sward of dwarf shrubs isdense, but shorter or more open than it is in the Typical H10a. The mixtures of Calluna vulgaris and Ericacinerea grow through dense silvery carpets of the moss Racomitrium lanuginosum. There is a sprinkling ofsmall plants such as Antennaria dioica, Huperzia selago, Polygala serpyllifolia, Potentilla erecta, Solidagovirgaurea and Succisa pratensis, and shoots of Deschampsia flexuosa, Agrostis canina and Carex binervis.The carpet of Racomitrium is patched with many other species such as Breutelia chrysocoma, Hypnumjutlandicum, Rhytidiadelphus loreus, Hylocomium splendens, Pleurozium schreberi, Frullania tamarisci andDiplophyllum albicans.

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H10b usually occurs on steep rocky slopes, stream-sides and cliff ledges where the soils are coarse-texturedand free-draining. It is common in the west Highlands and in the Hebrides.

H10c Calluna vulgaris-Erica cinerea heath, Festuca ovina-Anthoxanthum odoratum sub-community

H10c is a more grassy form of the community: commonly the result of local heavy grazing. The sward ismore mixed and variegated, with the dwarf shrubs Calluna vulgaris and Erica cinerea intermingled withgraminoids such as Agrostis capillaris, A. canina, Anthoxanthum odoratum, Festuca ovina, F. vivipara,Deschampsia flexuosa, Nardus stricta, Carex panicea, C. binervis and Luzula multiflora. The sward is dottedwith species such as Potentilla erecta, Galium saxatile and Blechnum spicant. Bryophytes are not as easy tosee among the grasses as they are under the shrubs of the other sub-communities, but a search will usuallyreveal a reasonable number of species including Hylocomium splendens, Hypnum jutlandicum, Thuidiumtamariscinum, Pleurozium schreberi, Rhytidiadelphus loreus and R. squarrosus.

There are small patches of H10c on Meall Mór, in the main glen and in Glen Etive, generally in mosaicsof grassland. H10c is common in the west Highlands, particularly where grazing is heavy.

H10d Calluna vulgaris-Erica cinerea heath, Thymus polytrichus-Carex pulicaris sub-community

This is the herb-rich form of Calluna-Erica heath. It occurs on well-drained soils over neutral or basic rocks,and was recorded here only in one place: on stabilized shingle by the River Etive. At this place there is asmall patch of heath some 15cm tall, with about half of the sward made up of Calluna vulgaris and Ericacinerea. The grasses Agrostis capillaris, Anthoxanthum odoratum, Deschampsia flexuosa, Festuca ovina, F. rubra and Nardus stricta are common among these dwarf shrubs. H10d resembles the H10c heathdescribed above, but species such as Thymus polytrichus, Lotus corniculatus and Hypochoeris radicata showthat the soils are less acidic than those with the less herb-rich H10c. Prunella vulgaris, Plantago lanceolataand Rhinanthus minor occur in small quantity and also indicate richer soils. Other species here includeAntennaria dioica – here evidently taking advantage of the rather open ground conditions with littlecompetition from taller plants – and the moss Racomitrium ericoides which forms conspicuous patches as itcommonly does on river shingle.

H12b Calluna vulgaris-Vaccinium myr ti l lus heath, Vaccinium vitis-idaea-Cladonia impexasub-community

Much less common in the study area than the Calluna-Erica heath H10, H12b occurs on some steep, rockyslopes facing north-west to east and south-east. The soils are coarse-grained and free-draining. There arepatches of H12b on the eastern side of Beinn Fhada and on the northern side of Buachaille Etive Mór. It has a dense, dark-coloured sward of Calluna vulgaris, interleaved with a little Vaccinium myrtillus and V. vitis-idaea, with Carex binervis, Potentilla erecta and Galium saxatile. There is a dense and colourfulunderlay of bryophytes made up of species such as Pleurozium schreberi, Hylocomium splendens,Rhytidiadelphus loreus, Hypnum jutlandicum and Racomitrium lanuginosum.

H12b is widespread throughout the northern British uplands, especially in the central and eastern Highlands.

H12c Calluna vulgaris-Vaccinium myrtillus heath, Galium saxatile-Festuca ovina sub-community

This is a short, grassy form of H12 heath, in which Calluna vulgaris is very common but as low-grown, well-

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browsed shrubs among grasses, mosses and scattered Vaccinium myrtillus. The grasses are mainly Agrostiscanina, Festuca vivipara and Nardus stricta, and the commonest mosses are Hylocomium splendens,Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphus loreus and Racomitrium lanuginosum. Varioussmall vascular species can grow here, especially Alchemilla alpina, Potentilla erecta, Euphrasia agg.,Campanula rotundifolia and Polygala serpyllifolia.

There are small patches of H12c heath among the very extensive grasslands on the steep and rather heavilygrazed west-facing slopes on the eastern side of Fionn Ghleann, west of Bidean nam Bian.

H12c is widespread in upland Britain.

H13a Calluna vulgaris-Cladonia arbuscula heath, Cladonia arbuscula-Cladonia rangiferina

sub-community

On the summit ridge of Meall Mór – a level top reaching out to the north-west – there are a few small patchesof this lichen-rich, prostrate, montane Calluna heath. The heather grows in a grey-brown matted carpet,pressed tightly to the surface of the ground. Around it there is a dense white layer of the lichens Cladoniaportentosa, C. arbuscula and C. uncialis, blotched with the darker tones of Cladonia rangiferina, C. gracilis, Coelocaulon aculeatum and Cetraria islandica, the silvery shoots of Racomitrium lanuginosumand the green rosettes of Polytrichum alpinum. A few small plants prick up through the tight layer of heatherand lichens: species such as Empetrum nigrum ssp. hermaphroditum, Vaccinium myrtillus, Nardus stricta,Huperzia selago and Diphasiastrum alpinum.

This vegetation has more in common with the lichen-rich montane heaths of the eastern and central Highlandsthan with the more usual western communities with Racomitrium, so it has been classed as H13. It isinteresting that there are also lichen-rich patches of montane Nardus-Galium grassland U5e (see below) onthe same hill.

H14 Calluna vulgaris-Racomitrium lanuginosum heath

There is very little prostrate heath in the study area, but most of what occurs here is of this type. H14 has ashort sward of Calluna vulgaris, creeping in grey-brown strips and patches over gravel and rock outcropsand through soft, silvery swards of the moss Racomitrium lanuginosum. The shoot-tips of the heather and themoss all point away from the prevailing south-westerly winds. As in H13, a few small plants are able togrow in the meagre shelter. They include Carex pilulifera, C. bigelowii, Empetrum nigrum ssp.hermaphroditum, Nardus stricta, Vaccinium myrtillus, Trichophorum cespitosum and Potentilla erecta. Theremay be conspicuous patches of lichens such as Cladonia portentosa, C. uncialis and Cetraria islandica,but they are not as common as they are in the lichen-rich heath H13. The H14 in the study area is all ratherspecies-poor and the patches are small. It does not fit well into any of the three sub-communities, thoughperhaps has most in common with the Festuca ovina sub-community H14a.

We found H14 on Meall Mór and on Aonach Dubh and Gearr Aonach, on high exposed rocky ridgeswhich are exposed to the full force of the wind and generally blown clear of snow in winter. H14, thewestern, oceanic counterpart of the boreal lichen-rich heaths, is common on the higher hills of the northernand western Highlands. There are fragments in the Lake District and in North Wales.

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H18a Vaccinium myrtillus-Deschampsia flexuosa heath, Hylocomium splendens-Rhytidiadelphusloreus sub-community

These are rich green heathlands, forming small patches in mosaics with other heaths and grasslands on thelong steep slopes in the corries. They have a dense leafy sward of Vaccinium myrtillus, V. vitis-idaea andlocally Empetrum nigrum ssp. nigrum, spiked through by shoots of Deschampsia flexuosa, Festuca vivipara,Agrostis canina, Carex binervis and Luzula multiflora and woven with the trailing stems of Potentilla erectaand Galium saxatile. A few other species are dotted here and there, including Nardus stricta, Alchemillaalpina, Juncus squarrosus, Blechnum spicant and Luzula sylvatica. Under the vascular plants there is a deep,springy layer of bryophytes made up of large common mosses such as Pleurozium schreberi, Hylocomiumsplendens, Hypnum jutlandicum, Rhytidiadelphus loreus, R. squarrosus, Plagiothecium undulatum, Polytrichumcommune and Dicranum scoparium, with some clumps of Sphagnum capillifolium.

Most of the H18a in the study area lies within the former altitudinal range of woodland, and is probablythe remnant of heathy woodland vegetation of the Quercus-Betula-Dicranum type W17. Other patches lieat higher altitudes and have in them a few montane species such as Polytrichum alpinum or Empetrum nigrumssp. hermaphroditum. These may be more natural heaths, marking out places where snow lies long enoughto suppress the growth of Calluna vulgaris.

H18a heaths are common throughout the British uplands, though less so in the western Highlands. Someappear near-natural; some are evidently entirely anthropogenic. The flora does not vary very much whetherthe heaths are derived by burning and grazing from more natural woodlands or heathery heaths, or arenear-natural snow-bed and montane communities.

H18b Vaccinium myr ti l lus-Deschampsia flexuosa heath, Alchemilla alpina-Carex piluliferasub-community

H18b is more common than H18a in the study area, probably because of the moderately base-rich rocksand the considerable amounts of base-rich flushing in the steep-sided glens and corries where this type ofheath grows in great spreads of brilliant green vegetation. H18b is a Vaccinium myrtillus heath like H18a,but the blaeberry bushes are more sparse. The sward is more grassy and is sprinkled all over with thedelicate silvery leaves and small yellow-green flowers of Alchemilla alpina. There are small sprigs ofVaccinium vitis-idaea and clumps of Empetrum nigrum, growing with grasses such as Festuca vivipara,Anthoxanthum odoratum, Agrostis capillaris, Nardus stricta, Deschampsia flexuosa and Danthonia decumbens.Many other plants play a small part in the sward: species such as Polygala serpyllifolia, Potentilla erecta,Galium saxatile, Carex pilulifera, C. binervis, Campanula rotundifolia, Anemone nemorosa and Veronicaofficinalis. Lycopodium clavatum forms straggling patches, and more locally there are plants of the scarcerL. annotinum (Photograph 41). The bryophyte layer is much like that of H18a, with thick mats of speciessuch as Hylocomium splendens, Rhytidiadelphus loreus, R. squarrosus, Hypnum jutlandicum, Pleurozium schreberiand Racomitrium lanuginosum.

H18b is common in the study area on the steep slopes of the glens and straths. It occurs quite widely in theHighlands, but is most extensive on the Breadalbane hills running from Ben Lui north-east to Caenlochan.

H20a Vaccinium myrtillus-Racomitrium lanuginosum heath, Viola riviniana-Thymus polytrichussub-community

This is one of the montane Vaccinium myrtillus heaths: vegetation of steep, rocky ground at high altitudes

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and of rock-strewn plateaux and ridges. It has a mixed, rich-green sward of Vaccinium myrtillus, V. vitis-idaeaand V. uliginosum, variegated with clumps of Empetrum nigrum ssp. hermaphroditum. The shrubs areinterleaved with montane plants such as Carex bigelowii, Diphasiastrum alpinum and Deschampsiacespitosa ssp. alpina as well as the more widespread Carex pilulifera, Deschampsia flexuosa, Festucavivipara and Huperzia selago. Alchemilla alpina grows thickly in the sward, and there can be some Thymuspolytrichus as well. There is a deep and conspicuous layer of bryophytes. Racomitrium lanuginosum is themost common species, but there is also much Hylocomium splendens, Pleurozium schreberi, Rhytidiadelphusloreus, Barbilophozia floerkei and Diplophyllum albicans as well as a sprinkling of more choice montanespecies such as Conostomum tetragonum, Oedopodium griffithianum, Polytrichum alpinum and Kiaeria starkei.

Of all the sub-communities of Vaccinium-Racomitrium heath, H20a is the one most strongly associated with the richer rocks. It is common in the study area and there are patches of it on most of the high hills. It is especially extensive and species-rich on the precipitous upper slopes of Aonach Eagach (see Photograph 14). H20a occurs throughout the northern and western Highlands.

H20b Vaccinium myr til lus-Racomitrium lanuginosum heath, Cetraria islandica sub-community

This is the standard or ordinary montane Vaccinium-Racomitrium heath with neither the Alchemilla alpina,Thymus and grasses of H20a nor the oceanic liverworts of H20c. It has a dense deep-green sward ofVaccinium myrtillus, with much V. vitis-idaea and Empetrum nigrum ssp. hermaphroditum. Vacciniumuliginosum may grow here too. There are tufts of Deschampsia flexuosa, Nardus stricta, Festuca vivipara,Carex pilulifera and C. binervis, winding shoots of Potentilla erecta and Galium saxatile and a range ofsmall upland or montane species such as Carex bigelowii, Diphasiastrum alpinum and Huperzia selago.Around these plants there is a thick silver-grey felt of Racomitrium lanuginosum, variegated with otherbryophytes such as Hylocomium splendens, Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphusloreus, Diplophyllum albicans and Barbilophozia floerkei and sprinkled with lichens such as Cetrariaislandica, Cladonia portentosa, C. uncialis and Coelocaulon aculeatum.

H20b is common in the study area, especially on the more acid rocks of Buachaille Etive Beag andBuachaille Etive Mór. There are also patches on the main ridge and its spurs. H20b is the characteristicvegetation of high, stony slopes in the Scottish Highlands. It occurs in north Wales and probably also in theLake District.

H20c Vaccinium myr tillus-Racomitrium lanuginosum heath, Bazzania tricrenata-Mylia taylorisub-community

H20c is a Vaccinium-Racomitrium heath with a short green tufted sward of Vaccinium myrtillus, V. vitis-idaea,V. uliginosum and Empetrum nigrum ssp. hermaphroditum. This shelters species such as Carex bigelowii, C. pilulifera, Deschampsia flexuosa, Diphasiastrum alpinum, Huperzia selago, Blechnum spicant, Potentillaerecta and Galium saxatile. The sward of vascular plants is underlaid by dense wefts of Racomitriumlanuginosum. In this sub-community the bryophyte layer is more varied than it is in the other forms of H20.It includes a rich array of uncommon oceanic liverworts such as Bazzania tricrenata, B. pearsonii,Anastrepta orcadensis, Mylia taylorii, Lepidozia pearsonii, Pleurozia purpurea, Plagiochila carringtonii,Scapania ornithopodioides, S. nimbosa (Photograph 43) and Anastrophyllum donnianum (Photograph 40).There are also red hummocks of Sphagnum capillifolium and golden patches of Hylocomium splendens,Rhytidiadelphus loreus and Pleurozium schreberi.

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H20c is less common than the other sub-communities here, and is confined to slopes facing between north-west and east. We found it among boulders deep in the high corries of Bidean and in Coire na Tulaich onBuachaille Etive Mór (Photograph 39). The distribution of this type of vegetation is defined by the distributionsof the characteristic oceanic liverworts. H20c occurs in the north-west Highlands from Ben Hopesouthwards, and extends across the central Highlands to the Cairngorms. There are outlying examples withfewer liverwort species in the highest hills of north Wales.

H21a Calluna vulgaris-Vaccinium myr ti l lus-Sphagnum capil l i folium heath, Calluna vulgarisPteridium aquil inum sub-community

H21a is a damp heath with a tall shaggy canopy of Calluna vulgaris interleaved with Vaccinium myrtillus,V. vitis-idaea and in some places Empetrum nigrum or Erica cinerea or both. These shrubs grow over a deep,rich-red quilt of Spghagnum capillifolium, patched with other bryophytes such as Racomitrium lanuginosum,Breutelia chrysocoma, Hypnum jutlandicum, Rhytidiadelphus loreus, Hylocomium splendens, Pleuroziumschreberi, Plagiothecium undulatum, Dicranum scoparium, Polytrichum commune and Diplophyllum albicans.There are clumps of Blechnum spicant, and tufts and patches of species such as Potentilla erecta, Galiumsaxatile, Melampyrum pratense, Huperzia selago, Deschampsia flexuosa, Agrostis canina, Succisa pratensisand Luzula sylvatica. The heaths may be home to the tiny and uncommon orchid Listera cordata. Theclubmosses Lycopodium clavatum and L. annotinum can grow here too. The oceanic bryophytes whichdefine the Mastigophora-Herbertus sub-community H21b are scarce here, although there can be odd clumpsand patches of the more widespread species such as Scapania gracilis, Anastrepta orcadensis, Myliataylorii and Lepidozia pearsonii.

H21a is widespread in the study area. It is especially extensive on the craggy precipitous northern faces ofAonach Dubh, Gearr Aonach, Buachaille Etive Beag and Buachaille Etive Mór. It occurs through the Britishuplands from south-west England north to Orkney and Shetland, generally on shaded slopes facing betweennorth-west and east. In the western Highlands and the Hebrides it can also be found on level ground withdamp and peaty soils.

H21b Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath, Mastigophora woodsiiHerber tus aduncus hutchinsiae sub-community

Like H21a, this is a damp heath. It is less common than H21a in the study area, and all but tiny fragmentsare confined to the steep crags where they are nearly inaccessible. The tall untidy sward of Calluna vulgarisand Vaccinium myrtillus, speckled with V. vitis-idaea, Empetrum nigrum and Erica cinerea, covers anassemblage of smaller plants such as Blechnum spicant, Deschampsia flexuosa, Succisa pratensis, Huperziaselago, Galium saxatile and Potentilla erecta. The bryophyte carpet is as deep and continuous as it is inH21a, but is more richly varied. The red cushions of Sphagnum capillifolium, the silver mats of Racomitriumlanuginosum and the golden wefts of large mosses such as Hylocomium splendens, Pleurozium schreberi,Rhytidiadelphus loreus and Hypnum jutlandicum are mixed with clumps and hummocks of large oceanicliverworts. These include the bright orange Herbertus aduncus (Photograph 48), the purple Pleuroziapurpurea (Photograph 49), and Bazzania tricrenata, B. pearsonii, Scapania gracilis, S. nimbosa, S. ornithopodioides, Mastigophora woodsii, Mylia taylorii, Plagiochila carringonii, P. spinulosa andLepidozia pearsonii. Low down in the glen the more lowland species Saccogyna viticulosa and Bazzaniatrilobata are also common in these mats of liverworts, and the oceanic fern Hymenophyllum wilsonii growsamong the bryophytes and over rocks and tree-roots.

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H21b occurs in mosaics with H21a and other heaths, grasslands and woodlands on the cliffs of AonachDubh, Gearr Aonach, Beinn Fhada, Buachaille Etive Beag and Buachaille Etive Mór. There are fragmentson steep rocky ground below the cliffs in one or two places. Like the liverwort-rich Vaccinium-Racomitriumheath H20c, the distribution of H21b is defined by that of the liverwort species. It does not extend as far into the eastern Highlands as H20c, but is reasonably common down the western seaboard of theHighlands and in the Hebrides on suitable steep, damp, shaded, rocky slopes. It also occurs in the Lake District and in north Wales.

H22a Vaccinium myr ti l lus-Rubus chamaemorus heath, Polytrichum commune-Galium saxatilesub-community

This is a montane heath of steep and high but sheltered slopes which accumulate a reasonable amount of snow in winter and where the soils are deep and moist. It has a thick and rather uneven sward ofVaccinium myrtillus and Empetrum nigrum ssp. hermaphroditum, intermingled with Vaccinium uliginosum(Photograph 42), V. vitis-idaea and even a little Calluna vulgaris. These shrubs are mixed with grasses suchas Deschampsia flexuosa, Nardus stricta, Agrostis canina and Anthoxanthum odoratum, and with smallheathland species such as Potentilla erecta, Galium saxatile, Polygala serpyllifolia and Blechnum spicant.There is an assemblage of montane or upland species such as Carex bigelowii, Diphasiastrum alpinum,Huperzia selago and Alchemilla alpina and, in many patches, Cornus suecica, a plant which is quitecharacteristic of this type of vegetation. We also found the uncommon Pyrola minor in H22a on the northside of Buachaille Etive Beag, and Lycopodium annotinum in H22a on Buachaille Etive Mór (Photograph 41).Under the vascular plants there is a thick and colourful mat of bryophytes, made up of many speciesincluding Sphagnum capillifolium, Hylocomium splendens, Rhytidadelphus loreus, Hypnum jutlandicum,Pleurozium schreberi, Plagiothecium undulatum, Dicranum majus, Polytrichum commune, Barbilophoziafloerkei, Diplophyllum albicans, Anastrepta orcadensis and Tritomaria quinquedentata.

H22a occurs on the upper slopes of Aonach Dubh, Gearr Aonach, Beinn Fhada, Buachaille Etive Beag andBuachaille Etive Mór as well as on the main ridge of Bidean. Mostly it is in the upper corries or on shelteredconcave slopes. This type of vegetation is widespread in the higher hills of the Scottish Highlands and thereare similar heaths in a few high corries in north Wales.

HX Vaccinium myr ti l lus-Sphagnum capil l i folium heath

This type of vegetation is not described in the NVC, although we are finding it increasingly often in placeswhere there is or has been heavy grazing in Wales, northern England and parts of Scotland. It is a damp heathwhich seems to have been derived from Calluna-Vaccinium-Sphagnum heath H21 where the Calluna vulgarishas been eliminated by heavy grazing. This could well have happened in the study area, as many patchesare on steep north-facing or east-facing slopes where there are H21 heaths dominated by heather on ungrazedcliffs above. HX has a short and dense sward of Vaccinium myrtillus, tufted with species such as Deschampsiaflexuosa, Nardus stricta, Agrostis canina, Blechnum spicant, Potentilla erecta, Galium saxatile and Luzulasylvatica. These plants grow through a spongy layer of bryophytes consisting of mixtures of Sphagnumcapillifolium, Hypnum jutlandicum, Hylocomium splendens, Rhytidiadelphus loreus, Racomitrium lanuginosum,Polytrichum commune, Pleurozium schreberi, Barbilophozia floerkei and Diplophyllum albicans. Locally thereare some western liverwort species such as Scapania gracilis, Anastrepta orcadensis or Mylia taylorii.

HX is quite common on the north-west-facing sides of Coire nan Lochan and Coire Gabhail, and in Coire nan Cuilc on Meall Mór.

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M1 Sphagnum denticulatum bog pool community

This is the vegetation of the shallow peaty depressions and tiny pools which form a part of the wetter blanketmires (see Photograph 35). These depressions and pools may be widely scattered over the bog surface, oroccur close together in patterns looking rather like nets or ladders. They may be less than a metre across,or cover many square metres. They usually contain half-submerged, half-floating mats of Sphagnumdenticulatum, and may also have much S. cuspidatum, S. papillosum and S. fallax. These spongy mats arepierced through by plants such as Eriophorum angustifolium, Rhynchospora alba, Juncus bulbosus,Trichophorum cespitosum and Molinia caerulea, and dotted with smaller plants such as Drosera longifolia,D. rotundifolia, Narthecium ossifragum and Pleurozia purpurea. The scarce sedge Carex pauciflora(Photograph 47) grows in this type of vegetation in pools north of Buachaille Etive Beag. In the larger anddeeper examples there can be spreads of Menyanthes trifoliata growing through the carpets of Sphagnumand extending out into open water. Utricularia minor grows in some of these pools too.

Small patches of M1 vegetation are common within the Trichophorum-Eriophorum blanket bogs M17 in thestudy area, especially in the north-eastern corner of the site where the low ground around Buachaille Etive Beagand Buachaille Etive Mór is level and peat-covered. M1 mires occur throughout the western British uplands,following the distribution of the wetter Trichophorum-Eriophorum mires M17 with which they are associated.

M3 Eriophorum angustifolium bog pool community

In contrast to M1, patches of the M3 community can occur in terrestrial habitats with no standing water, althoughin spells of wet weather the ground may be submerged under a few centimetres of water. M3 is a vegetationtype of very shallow peaty pools, spreads of redistributed peat around hags, and channels and runnels in blanketbogs. It consists of dense swards of Eriophorum angustifolium, the red tips of its dark green leaves making thevegetation conspicuous from afar. If there are ever any other species in these pools they are few and the plantsare scattered. The most common are Trichophorum cespitosum, Molinia caerulea, Erica tetralix, Callunavulgaris, Narthecium ossifragum and the mosses Sphagnum papillosum, S. denticulatum and S. cuspidatum.

Although the NVC account of M3 states that this is the typical pool component of the drier, more northernand boreal Calluna-Eriophorum mires M19, in fact it is also common in the wetter, oceanic Trichophorum-Eriophorum mire M17, which is where it occurs in the study area. There are some large M3 pools in thebogs on the level low ground in the north-east of the study area. M3 mires are common in bogs throughoutthe British uplands.

M4 Carex rostrata-Sphagnum fallax mire

These acid Carex rostrata mires have a thin grey-green sward of the sedge, dotted with its heavy yellow-green heads in summer, growing through a bright green spongy carpet of Sphagnum palustre, S. fallaxand Polytrichum commune. This is speckled with a few small species such as Viola palustris, Carex nigra,C. echinata, Potentilla erecta and Ranunculus flammula. There can be a few shoots of Molinia caerulea,Eriophorum angustifolium, Juncus effusus or J. acutiflorus.

The typical sites for M4 are wet hollows and channels in blanket bogs or flushed ground at the edges oflochs. M4 occurs in the study area in the bogs in the north-eastern corner and on the shores of Loch Achtriochtan. It is a common type of mire vegetation in the uplands of Wales, northern England andScotland, though the individual patches are rarely large.

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M6a Carex echinata-Sphagnum fallax/denticulatum mire, Carex echinata sub-community

These are small, inconspicuous mires, hard to distinguish at a distance from grasslands. They have a sparse,short sward of Carex echinata, mixed with a little C. nigra, C. panicea, Agrostis canina, Holcus lanatusand Anthoxanthum odoratum. These plants grow through a soft and wet layer of bryophytes such asSphagnum fallax, S. palustre, S. denticulatum, Polytrichum commune, Rhytidiadelphus loreus andLophocolea bidentata. The mires are rarely herb-rich, and there is never more than a thin speckling amongthe sedges of small species such as Viola palustris, Potentilla erecta and Rumex acetosa.

M6a mires occur on flushed slopes in mosaics of grasslands, wet heaths and other mires and are mostcommon on Meall Mór and the southern slopes of the Aonach Eagach. M6a is one of the most commontypes of acid flush vegetation in the British uplands.

M6b Carex echinata-Sphagnum fal lax/dent iculatum mire, Carex nigra-Nardus s tr ic tasub-community

M6b mires are also small and generally dominated by sedges, but can be somewhat more variable thanM6a. The most common plant in these mires is usually Carex nigra, growing either as pure swards or invarying mixtures with C. echinata, C. panicea, Nardus stricta and Eriophorum angustifolium over the usualgreen carpet of acidophilous bryophytes Sphagnum fallax, S. palustre, S. denticulatum and Polytrichumcommune. Other bryophytes here include Rhytidiadelphus loreus and Lophocolea bidentata. There areusually a few other vascular species such as Potentilla erecta, Viola palustris, Rumex acetosa, Holcus lanatus,Agrostis canina, Anthoxanthum odoratum and Juncus squarrosus.

M6b is scarcer than M6a in the study area. It occurs in wet hollows and flushed channels among wetmontane heaths and grasslands in Coire nam Beitheach. This type of mire is common in the British uplands,especially within blanket bogs and wet heaths.

M6c Carex echinata-Sphagnum fallax/denticulatum mire, Juncus ef fusus sub-community

There are small patches of this type of rush-mire in the main glen; most of it around the shores of Loch Achtriochtan. It has a tall, dark dull-green sward of Juncus effusus, often flattened by heavy rain orflooding streams. This conceals the spongy underlayer of Sphagnum fallax, S. denticulatum, S. palustreand Polytrichum commune. There are generally few other species among the rushes apart from a little Carex echinata, C. nigra, Holcus lanatus, Agrostis canina, Potentilla erecta and Viola palustris.

M6c is widespread throughout the uplands of England, Wales, southern Scotland and the Inner Hebrides.It is less common in the north of Scotland.

M6d Carex echinata-Sphagnum fallax/denticulatum mire, Juncus acutiflorus sub-community

Most of the rush mires in the study area are of this type. They have a tall, dense and rich-green sward ofJuncus acutiflorus, interleaved with Molinia caerulea and Carex panicea and sprinkled with species such asPotentilla erecta, Carex echinata, Succisa pratensis, Viola palustris, Epilobium palustre and Eriophorumangustifolium. These plants grow through a thick wet underlay of bryophytes such as Sphagnum fallax, S. palustre, S. denticulatum, Polytrichum commune, Hylocomium splendens, Rhytidiadelphus squarrosus,Aulacomnium palustre and Lophocolea bidentata.

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Large and distinctive patches of M6d occur on the lower slopes of Meall Mór, all along the main glen andin Glen Etive. M6d is common in the mild climate of south and west Britain, but becomes more scarce inthe far north and in the east.

M7 Carex cur ta-Sphagnum russowii mire

The NVC description of this montane mire is rather vague and misleading, as Carex curta and the montaneSphagna can be absent. Acid soligenous mires at high altitudes can be quite variable, just as their sub-montane equivalents can, but they generally have a reasonably distinct flora including a few montanespecies. The quadrats show two examples. The sward of vascular plants is almost always sparse and open:certainly less dense than in typical examples of the sub-montane acid mires M6a and M6b. It consists of small sedges and related species such as Eriophorum angustifolium, Carex bigelowii, C. nigra and Juncus bulbosus. Carex curta does occur in some examples of these mires but is not ubiquitous. Under thesegraminoids is a patchy layer of bryophytes such as Sphagnum papillosum, S. denticulatum, Polytrichumalpinum, P. commune, Racomitrium lanuginosum and R. heterostichum. There is a speckling of small plantssuch as Potentilla erecta, Viola palustris, Narthecium ossifragum and the more montane Saxifraga stellaris,Alchemilla alpina and Deschampsia cespitosa ssp. alpina.

M7 mires occur in flushed hollows and channels at high altitudes, and along the edges of lochs or streams.We found them among the wet grasslands in Coire nan Lochan and on the plateau of Sgorr nam Fiannaidh.The community is widespread through the higher hills of the Highlands. It also occurs in the Lake District andthere is similar vegetation in north Wales.

M10a Carex dioica-Pinguicula vulgaris mire, Carex viridula ssp. oedocarpa-Juncus bulbosus/

kochii sub-community

These are small mires which mark out places of moderate base-rich flushing. They are usually small, dark-coloured, rather open and stony patches which follow lines of seepage on the hillsides. They have a shortand open sward of sedges such as Carex panicea, C. viridula ssp. oedocarpa, C. dioica, C. hostiana andC. pulicaris, interspersed with other small and base-tolerant species such as Prunella vulgaris, Selaginellaselaginoides, Ranunculus acris, Parnassia palustris and Thalictrum alpinum and dotted with the greasy lime-green rosettes of Pinguicula vulgaris. There are also generally a few plants of more acid mires, such asNarthecium ossifragum, Drosera rotundifolia, Molinia caerulea, Trichophorum cespitosum and Eriophorumangustifolium. Among the vascular plants are wisps and tufts of bryophytes such as Drepanocladus revolvens,Scorpidium scorpioides, Campylium stellatum, Ctenidium molluscum, Breutelia chrysocoma, Calliergonellacuspidata and Calliergon sarmentosum.

M10a mires are common on the hill slopes where base-rich water seeps out of the ground. They are themost widespread type of small mesotrophic to basic mire in the British uplands.

M11a Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Thalictrum alpinum-Juncus

triglumis sub-community

These are stony flushes, generally more open and sparsely-vegetated than the M10 mires described above.They are, however, usually more conspicuous. They have a distinctive sward of Carex panicea, C. viridulassp. oedocarpa, C. hostiana and C. pulicaris growing through mats of Saxifraga aizoides with its succulent

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shining green leaves and its tumbling cascades of starry yellow flowers. Set in this sward are many small species such as Selaginella selaginoides, Eleocharis quinqueflora, Thalictrum alpinum, Saxifragaoppositifolia, Ranunculus acris, Alchemilla alpina, A. glabra, Juncus triglumis and Linum catharticum. Some of the M11 mires on Meall Mór are home to the scarcer Tofieldia pusilla and Carex capillaris. Among the stones are cushions and patches of bryophytes such as Campylium stellatum, Blindia acuta,Drepanocladus revolvens, Scorpidium scorpioides, Calliergonella cuspidata, Calliergon trifarium, C. sarmentosum, Ctenidium molluscum, Hylocomium splendens and Breutelia chrysocoma. Photograph 6shows an example of an M11 mire on Meall Mór.

M11a mires are common in the west of the study area, on flushed slopes from the lower hillsides up intothe high corries. This type of mire is widespread in the Scottish Highlands and northern England. There aresimilar mires in the Southern Uplands and north Wales, where the characteristic species Saxifraga aizoidesdoes not occur.

M11b Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Palustriel la commutata-

Eleocharis quinqueflora sub-community

This sub-community of M11 occurs on the flushed northern slopes of Meall Mór at low to moderate altitudes.It resembles the more montane sub-community M11a, with its rich-green sward of Carex viridula ssp.oedocarpa, C. panicea, C. pulicaris, C. dioica and C. hostiana and attractive clumps of Saxifragaaizoides, but plants such as Eriophorum latifolium, Molinia caerulea, Narthecium ossifragum, Succisapratensis and Potentilla erecta are more common here, and the bright orange-gold moss Palustriellacommutata plays a large part in the array of bryophytes, growing with plants such as Ctenidium molluscum,Blindia acuta, Breutelia chrysocoma and Orthothecium rufescens. The more montane species such asThalictrum alpinum and Juncus triglumis are scarce or absent.

M11b accounts for some of the M11 mires at lower elevations here. Some M11b flushes are downstreamfrom Palustriella-Carex springs M37. The two vegetation types share many species.

M12 Carex saxati l is mire

There are a few patches of M12 in the upper part of Coire nam Beitheach, where the soils are flushed withcold base-rich water draining from the basic rocks above. These mires have dense deep-green swards ofCarex saxatilis interleaved with Deschampsia cespitosa ssp. alpina, Anthoxathum odoratum and Agrostiscanina and with a great array of small base-tolerant species such as Carex viridula ssp. oedocarpa, C. pulicaris, Ranunculus acris, Leontodon autumnalis, Geum rivale, Trollius europaeus, Persicaria vivipara,Pinguicula vulgaris, Thalictrum alpinum, Alchemilla glabra, A. alpina, Selaginella selaginoides, Saxifragahypnoides, S. aizoides and Crepis paludosa. The wet clumps and mats of bryophytes are half-hidden bythe taller plants, but there are many species including Blindia acuta, Hylocomium splendens, Drepanocladusrevolvens, Campylium stellatum, Ctenidium molluscum, Fissidens adianthoides, Dicranella palustris,Scapania undulata and Aneura pinguis.

M12 mires are common only in the Breadalbane hills, but they occur more sparingly throughout theHighlands, at high altitudes where they are covered by snow in winter and where they are flushed with coldbase-rich water.

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M15a Trichophorum cespitosum-Erica tetralix wet heath, Carex panicea sub-community

This type of vegetation is an acid flush or soligenous mire rather than a wet heath, although it has much incommon with the other sub-communities of M15. The sward is a sparse and uneven mixture of Erica tetralix,Calluna vulgaris, Molinia caerulea, Trichophorum cespitosum, Juncus bulbosus, J. squarrosus, Carex paniceaand Narthecium ossifragum. The soil underneath is wet, peaty and dotted with stones. It is dotted over withspecies including Succisa pratensis, Drosera rotundifolia, D. anglica, Pinguicula vulgaris and Pedicularissylvatica and patched with bryophytes such as Sphagnum denticulatum, S. papillosum, Racomitriumlanuginosum, Campylopus atrovirens and the large red-purple liverwort Pleurozia purpurea (Photograph 48).

M15a is common in the study area. It occurs as flushes among acid heaths and grasslands on the hillsides,and fills hollows, wet channels and seepage lines in blanket bogs. It is a common type of acid flushthroughout the British uplands.

M15b Trichophorum cespitosum-Erica tetralix wet heath, Typical sub-community

This type of wet heath vegetation is widespread on the gentle to moderate slopes on the lower flanks of thehills, covering wet but shallow peat. It is thick tussocky vegetation (see Photograph 44) with a dense green-ochre sward of Calluna vulgaris, Molinia caerulea, Trichophorum cespitosum, Erica tetralix, Nartheciumossifragum and Potentilla erecta. Species such as Succisa pratensis, Polygala serpyllifolia, Pedicularissylvatica, Dactylorhiza maculata and Drosera longifolia are sprinkled through the sward. In some patchesthere is much Myrica gale. There are big mats and cushions of bryophytes too, and the most common andcharacteristic species are Sphagnum denticulatum, S. capillifolium, Campylopus atrovirens, Hylocomiumsplendens, Pleurozium schreberi, Racomitrium lanuginosum, Hypnum jutlandicum and the liverworts Pleuroziapurpurea and Diplophyllum albicans. Carex panicea, Drosera species, Narthecium ossifranum, Pinguiculavulgaris, Sphagnum denticulatum, Campylopus atrovirens and Pleurozia purpurea are not as common hereas they are in the wetter M15a sub-community described above.

M15b is common in the study area, as it is through the whole of the west Highlands and Hebrides. Furthersouth and east in the British uplands it is widespread but not as extensive as in the western Highlands.

M15c Trichophorum cespitosum-Erica tetralix wet heath, Cladonia sub-community

M15c tends to take over from M15b on steeper slopes, on thinner peats with more bare rock and stone,and at higher altitudes. There may be an altitudinal sequence from one to the other, or they may occur inmosaics on the same slope. Most of the M15 in the study area belongs to this sub-community. M15c canlook quite different from M15b. The sward is generally thinner, shorter and more open. Calluna vulgaris andTrichophorum cespitosum are common, but Erica tetralix is usually joined by E. cinerea, and although theremay be much Molinia caerulea it usually occurs as well-spaced stems and clumps rather than as densetussocks. The vascular plants are matted round with the silver-grey shoots of Racomitrium lanuginosum.Succisa pratensis, Potentilla erecta, Dactylorhiza maculata, Polygala serpyllifolia, Pedicularis sylvatica,Narthecium ossifragum, Nardus stricta and Huperzia selago are common here. Clumps of bryophytes suchas Pleurozia purpurea, Campylopus atrovirens, Sphagnum denticulatum and Hypnum jutlandicum break thecarpets of Racomitrium lanuginosum, and there can be a light frosting of lichens including Cladoniaportentosa, C. uncialis, C. subcervicornis, C. furcata, C. rangiferina and Coelocaulon aculeatum. A fewuncommon plants find a home in this type of vegtation: the club-moss Lycopodium annotinum and the orchid

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Pseudorchis albida were recorded in M15c in the study area. On the north-facing slopes in the main glen,below the wooded crags, the oceanic liverworts Herbertus aduncus, Pleurozia purpurea, Mylia taylorii,Scapania gracilis, Plagiochila spinulosa, Bazzania tricrenata, Anastrepta orcadensis and Lepidoziapearsonii grow in patches in the sward of M15c. This is an unusual element in this type of vegetation, exceptin the far western Highlands. At higher altitudes the sward is generally open and thin, with much Nardusstricta and species such as Diphasiastrum alpinum and Carex pilulifera. On top of Beinn Fhada there is anunusual montane form of this type of vegetation, in which the usual species of M15c are joined by Carexbigelowii, Vaccinium uliginosum and Empetrum nigrum ssp. hermaphroditum.

M15c is common in the Highlands and is one of the most widespread types of vegetation in the far westand in the Hebrides. Though described as the Cladonia sub-community, it is only in the northern Highlandsthat lichens really thicken up in the sward, and further south and west it is the Racomitrium lanuginosum andErica cinerea which define M15c. Much M15c occurs above the tree-line, where it may be a near-naturalform of vegetation on wet soils.

M15d Trichophorum cespitosum-Erica tetralix wet heath, Vaccinium myr ti l lus sub-community

This is a mixed and rather grassy damp heath with a short, thick sward of Calluna vulgaris, Erica tetralix,Vaccinium myrtillus, Trichophorum cespitosum, Molinia caerulea, Agrostis capillaris, A. canina,Anthoxanthum odoratum, Deschampsia flexuosa, Nardus stricta and Juncus squarrosus. Shoots of Potentillaerecta and Galium saxatile wind among the other species, and there are scattered individuals of plants suchas Luzula sylvatica, Vaccinium vitis-idaea, Alchemilla alpina and Narthecium ossifragum. There is a densemat of bryophytes including Hypnum jutlandicum, Racomitrium lanuginosum, Pleurozium schreberi,Hylocomium splendens, Rhytidiadelphus loreus, Dicranum scoparium, Polytrichum commune, Campylopusflexuosus and the liverworts Diplophyllum albicans and Mylia taylorii. On the upper slopes of BuachailleEtive Mór there is a montane form of M15d in which the flora is augmented by a few montane species suchas Carex bigelowii, Vaccinium uliginosum, Empetrum nigrum ssp. hermaphroditum and the oceanicliverworts Bazzania pearsonii and Anastrophyllum donnianum. This is similar to the montane M15c on BeinnFhada, but the Vaccinium species are more common here.

We found M15d on steep slopes and in the corries in all parts of the study area. M15d is widespread in the British uplands. It seems to cover drier, firmer peats than those with the other sub-communities, and inmany places appears to be produced and maintained by grazing.

M17a Trichophorum cespitosum-Eriophorum vaginatum blanket mire, Drosera rotundifolia-Sphagnum sub-community

Most of the blanket bog in the study area is of this type. It is vegetation of flat ground or at most gentleslopes (see Photographs 35 and 45). Looked at from a little distance it is an even, ochre-gold mixture ofTrichophorum cespitosum, Molinia caerulea, Eriophorum vaginatum, E. angustifolium and Sphagnumpapillosum, speckled with darker clumps of Calluna vulgaris and Erica tetralix and with the fragrant greenbushes of Myrica gale. Many smaller plants grow in the shelter of the sward: species such as Nartheciumossifragum, Succisa pratensis, Pedicularis sylvatica, Dactylorhiza maculata, Drosera rotundifolia, Potentillaerecta and Polygala serpyllifolia. In wetter parts there can be lawns of Rhynchospora alba, conspicuous insummer with its sharp cream-white flowers. Carex pauciflora (Photograph 47) grows around some of thepools and wet hollows. Although Sphagnum papillosum (Photograph 51) is generally abundant, there are

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usually also hummocks of S. capillifolium (Photograph 52) and soft shoots of S. tenellum, dark patches ofCampylopus atrovirens and purple clumps of Pleurozia purpurea (Photograph 48) as well as Hypnumjutlandicum, Campylopus flexuosus, Racomitrium lanuginosum and liverworts such as Mylia taylorii, M. anomala and Odontoschisma sphagni.

There are large areas of M17a on the low ground around Buachaille Etive Beag and Buachaille Etive Mórand in Glen Etive, and smaller patches elsewhere throughout the study area. M17a is the typical low-altitudeblanket bog in the oceanic climate along the western seaboard of Britain from Cornwall northwards, thoughin south-west England it is a community of hill-tops and ridges rather than of the low ground.

M17b Trichophorum cespitosum-Eriophorum vaginatum blanket mire, Cladonia sub-community

M17b forms mosaics with M17a on the level flats to the north of Buachaille Etive Beag. Compared withM17a, the peat is drier and may be more hagged. In many instances this type of bog has been dried outas a result of frequent burning. The main components of the vegetation are the same as those in M17a:Eriophorum vaginatum, E. angustifolium, Trichophorum cespitosum, Molinia caerulea, Erica tetralix andCalluna vulgaris, but here the carpets of Sphagna are not extensive and are largely replaced by Racomitriumlanuginosum, growing either as silvery sheets over the bog surface or as large grey-green hummocks whichare visible from a distance. Hypnum jutlandicum is common too. There are clumps of Mylia taylorii andpatches of Campylopus atrovirens, and the introduced C. introflexus can colonise dry banks of peat. Thevascular flora is less varied than it is in M17a, and in many patches there is little more than Potentilla erecta,Succisa pratensis and Polygala serpyllifolia.

M17b is widespread in the wetter western half of Britain.

M18a Erica tetralix-Sphagnum papillosum raised and blanket mire, Sphagnum magellanicum-

Andromeda polifolia sub-community

Within the larger patches of blanket mire around Buachaille Etive Beag, Buachaille Etive Mór and in Glen Etive, there are areas of this really wet ombrogenous mire (see Photograph 35). Even more than in theTrichophorum-Eriophorum mire M17a, Sphagna play a large part in the vegetation. Here the rich-redSphagnum magllanicum (Photograph 53) grows in gloriously-coloured red, golden and green patchworkswith S. papillosum, S. denticulatum, S. capillifolium, S. subnitens, S. tenellum and S. fallax in a spongy layerover almost liquid peat. The bog liverwort Odontoschisma sphagni straggles through the hummocks of bogmosses, and there are cushions of Racomitrium lanuginosum and Hypnum cupressiforme as well as clumpsof Pleurozia purpurea. Plants such as Erica tetralix, Molinia caerulea, Eriophorum vaginatum, E. angustifolium,Empetrum nigrum ssp. nigrum, Calluna vulgaris, Myrica gale and Trichophorum cespitosum are rooted in thebryophyte carpet, and there is a speckling of smaller species such as Drosera rotundifolia, Nartheciumossifragum and Carex pauciflora (Photograph 47). The main difference between M17a and M18a is thatthe layer of Sphagna in M18a is more species-rich and more continuous and the sward of vascular plantsless dense and less varied. Otherwise the two have much in common.

M18a occurs through the British uplands from Wales northwards. It is an important component of the ‘flows’ of Caithness and Sutherland and south-west Scotland, and is the typical vegetation of lowland raisedmires.

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M18b Erica tetralix-Sphagnum papillosum raised and blanket mire, Empetrum nigrum nigrum-

Cladonia sub-community

M18b is the drier and impoverished counterpart of M18a, just as M17b is of M17a. Racomitrium lanuginosumreplaces much of the Sphagnum carpet, growing in great lawns with Sphagnum magellanicum, S. papillosumand S. capillifolium. These are pierced through by the shoots of Molinia caerulea, Trichophorum cespitosum,Eriophorum vaginatum, E. angustifolium, Calluna vulgaris and Erica tetralix. There is much Nartheciumossifragum, and Empetrum nigrum ssp. nigrum may grow in dense green clumps. Bare peat can be quiteextensive.

M18b is widespread in northern and western Britain.

M19a Calluna vulgaris-Eriophorum vaginatum blanket mire, Erica tetralix sub-community

Small areas in the bogs around the northern foot of the Buachaille Etive Mór are of this type, and the bogon the summit of Meall Mór is intermediate between this and the Trichophorum-Eriophorum mire M17a. M19 mires look quite different from M17, and the two are easy to distinguish even at a distance. Insteadof the flat-looking, pale sward of M17, M19a has a taller, dense, tufted, dark-green and dark reddish-browncanopy of Calluna vulgaris and Eriophorum vaginatum. E. angustifolium is common too, and is easy to seewith its glossy red-tipped leaves overtopping the other species. There are shoots of Vaccinium myrtillus andV. vitis-idaea too, and even a little Erica cinerea. There are a few of the more characteristic M17 speciessuch as Erica tetralix, Molinia caerulea and Trichophorum cespitosum, together with common upland plantssuch as Potentilla erecta, Juncus squarrosus and Deschampsia flexuosa. The bryophyte flora is also verydifferent from that of M17. Here there is a deep quilt of Sphagnum capillifolium and large mosses such asPleurozium schreberi, Rhytidiadelphus loreus, Plagiothecium undulatum, Hypnum jutlandicum, Racomitriumlanuginosum and Hylocomium splendens, sprinkled with a few white shoots of Cladonia portentosa or C. uncialis. M19a is one of the main habitats for the diminutive upland orchid Listera cordata.

M19 is the characteristic blanket bog of the cooler, drier, more continental parts of Britain, especially thecentral and eastern Highlands, the Southern Uplands and the English Pennines. M19a is the mostwidespread sub-community in the wetter climate of the west.

M23a Juncus effusus/acutiflorus-Galium palustre rush-pasture, Juncus acutiflorus sub-community

There are some fine examples of these flowery herb-rich mires on the lower northern slopes of Meall Mór(Photograph 10), where the soils are flushed with base-rich water from the limestone rocks above. M23aalso occurs elsewhere on Meall Mór and around the shores of Loch Achtriochtan. M23 mires are amongthe most attractive types of upland vegetation. The tall, dark-green sward of Juncus acutiflorus is entwinedwith species such as Holcus lanatus, Agrostis canina, Anthoxanthum odoratum and Molinia caerulea andwith a great array of tall mesotrophic herbs such as Crepis paludosa, Angelica sylvestris, Geum rivale,Ranunculus acris, R. flammula, Filipendula ulmaria, Galium palustre, Senecio aquaticus, Parnassia palustris,Cardamine pratensis, Valeriana officinalis, Epilobium palustre, Lysimachia nemorum, Cirsium palustre,Lychnis flos-cuculi and Geranium sylvaticum. Under these are thin wefts of bryophytes such as Calliergonellacuspidata, Hylocomium splendens, Rhizomnium punctatum, Plagiomnium undulatum, Brachythecium rivulareand Trichocolea tomentella.

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In one place on the north side of Meall Mór there is M23a mire in which the dominant species is Juncusarticulatus rather than J. acutiflorus. Otherwise the flora is much the same as that just described.

M23a mires are common at low altitudes in the south-west Highlands and in the Hebrides, but become raretowards the north and east where the climate is cooler. They are one of the more thermophilous of our uplandvegetation types.

M25a Molinia caerulea-Potenti l la erecta mire, Erica tetralix sub-community

This type of wet grassland is common on the lower slopes of the hills, in the main glen and in Glen Etive. It covers shallow peats or wet peaty gleyed soils, generally on gentle to moderate concave slopes or onlevel ground. However, on the sides of Meall Mór it ascends onto ground which is remarkably steep andwell-drained for this type of vegetation. M25a has a tall, dense and tussocky sward and can be one of themost difficult types of upland vegetation to walk through – an especially frustrating matter in the Lairig Eildeand the Lairig Gartain where there is a difficult choice between tall Molinia grassland or a wet erodedfootpath through peat. The tussocks of Molinia caerulea are separated by deep runnels: damp channels half-filled with decomposing leaf-litter. The vegetation is species-poor. Potentilla erecta trails among the tussocks.There are clumps of Calluna vulgaris and Erica tetralix, and in some places green spreads of Myrica gale.Otherwise there is just a thin sprinkling of species such as Narthecium ossifragum, Carex panicea, Agrostiscanina, Festuca vivipara, Viola palustris and Succisa pratensis. On the bases of the tussocks there are matsof bryophytes such as Hypnum jutlandicum, Hylocomium splendens and Pleurozium schreberi, but in the wetrunnels there is little more than etiolated shoots of Rhytidiadelphus squarrosus or cushions of Sphagnumdenticulatum, S. palustre or S. fallax.

M25a is common and widespread in western Britain, from Cornwall and Devon northwards. Much of it isapparently derived from wet heaths by grazing in a mild wet climate.

M25b Molinia caerulea-Potenti l la erecta mire, Anthoxanthum odoratum sub-community

The sward of M25b is more of a mixture of grasses than those of the other sub-communities. Here the Moliniacaerulea grows in a tall, thick turf with Agrostis capillaris, A. canina, Festuca vivipara, F. rubra,Anthoxanthum odoratum, Danthonia decumbens, Nardus stricta and Deschampsia flexuosa. Hidden amongthe grasses are herbs such as Potentilla erecta, Polygala serpyllifolia, Campanula rotundifolia, Violariviniana, V. palustris, Succisa pratensis, Rumex acetosa and Narthecium ossifragum. On the ground thereare patches of bryophytes including Rhytidadelphus squarrosus, R. loreus, Hypnum jutlandicum, Hylocomiumsplendens, Pleurozium schreberi, Racomitrium lanuginosum and Thuidium tamariscinum.

M25b is common in the study area. It clothes the steep slopes of Meall Mór and the lower slopes of thecorries and glens. It usually occurs on better-drained soils than the Erica tetralix sub-community M25a. It iswidespread in the uplands of western Britain.

M25c Molinia caerulea-Potenti l la erecta mire, Angelica sylvestris sub-community

This is the herb-rich form of Molinia-Potentilla grassland. It is the least common of the three sub-communitiesin the study area, though there are some splendid patches of it on the northern slopes of Meall Mór(Photograph 9). Here a tall sward of Molinia caerulea, Agrostis canina, Anthoxanthum odoratum, Holcus

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lanatus, Helictotrichon pratense and Festuca vivipara is enriched with many large mesotrophic herbsincluding Ranunculus acris, Geum rivale, Filipendula ulmaria, Angelica sylvestris, Crepis paludosa, Trolliuseuropaeus, Valeriana officinalis and Parnassia palustris, as well as smaller species such as Carex viridulassp. oedocarpa, C. hostiana, C. pallescens, C. pulicaris, Pedicularis palustris, Selaginella selaginoides,Linum catharticum and Saxifraga aizoides. The uncommon montane species Silene acaulis and Carexcapillaris and the uncommon moss Orthothecium rufescens grow in M25c on Meall Mór. The mats ofbryophytes under the taller plants are made up of species such as the mosses Hylocomium splendens,Calliergonella cuspidata, Rhizomnium punctatum, Plagiomnium undulatum, Campylium stellatum andCtenidium molluscum.

M25c occurs throughout the western parts of the British uplands, and is locally common in the south-westand mid-west Highlands and in the Inner Hebrides.

M28a Iris pseudacorus-Fil ipendula ulmaria mire, Juncus spp sub-community

There are small patches of M28 on the lower slopes of Meall Mór and at the western end of the main glen.It is a community of neutral flushed soils at low altitudes. It is made up of tall green swards of Iris pseudacorus;its sword-like leaves scattered in summer with its untidy yellow flowers. Among the iris an array of grassesand mesotrophic herbs includes Agrostis canina, Festuca ovina, Anthoxanthum odoratum, Holcus lanatus,Rumex acetosa, Ranunculus acris, Stachys palustris, Lathyrus pratensis, Stellaria uliginosa, Galium palustreand Myosotis secunda. There is barely space for bryophytes amid the thick growth of vascular species, butthere are generally a few straggling wisps and wefts of Rhytidiadelphus squarrosus and Rhizomniumpunctatum. M28 is common in the western Highlands and Hebrides, mostly near the sea.

M31 Anthelia julacea-Sphagnum denticulatum spring

The silvery-green patches of this type of vegetation are made up of the tightly-overlapping tiny leaves of theliverwort Anthelia julacea. The mass of shoots forms a firm, swelling cushion of vegetation which is studdedwith small species such as Saxifraga stellaris, Selaginella selaginoides, Gnaphalium supinum, Narrtheciumossifragum, Pinguicula vulgaris, Nardus stricta and Huperzia selago. There are patches of a few otherbryophytes such as Sphagnum denticulatum, Oligotrichum hercynicum, Marsupella emarginata andScapania undulata.

Although these springs are widely distributed in the study area, most are at high altitudes and are especiallyassociated with snow-bed heaths and grasslands in the upper corries. Here they form part of the greatspreads of mixed spring and snow-bed vegetation in flushed hollows and on wet slopes. The M31 springsat lower elevations lack the more montane species and the patches are generally smaller. M31 occursthroughout the British uplands from north Wales northwards.

M32a Philonotis fontana-Saxifraga stel laris spring, Sphagnum denticulatum sub-community

This is the less diverse and species-rich form of M32 and is most common around springs where the wateremerges slowly through peat or peaty soils. It consists of distinctive red-gold patches of Sphagnumdenticulatum, pierced by shoots of species such as Agrostis canina, Juncus bulbosus, J. articulatus andAnthoxanthum odoratum and speckled with small plants including Montia fontana and Viola palustris. Thereare generally a few other bryophytes such as Scapania undulata, Philonotis fontana and Polytrichum commune.

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M32a mires are common throughout the study area, especially on the lower slopes and among the bogsand wet heaths. This type of spring occurs throughout the British uplands.

M32b Phi lonot i s fon tana-Saxi f raga s te l lar i s spr ing, Mont ia fontana-Chr ysosp len ium

opposi t i fo l ium sub-community

These springs are like little jewels of bright vegetation; sparkling with water and set among the drab coloursof the surrounding heaths and grasslands. They are almost always a bright and conspicuous shade of green.The cushions or mats are composed of a number of species which may grow in pure patches or in intricatemixtures. The most characteristic species are the mosses Philonotis fontana, Dicranella palustris, Bryumpseudotriquetrum, Calliergonella cuspidata, Drepanocladus fluitans, Sphagnum denticulatum and theliverwort Scapania undulata. Over these cushions are sprinkled small plants such as Saxifraga stellaris,Montia fontana, Chrysosplenium oppositifolium, Rumex acetosa, Agrostis canina, Ranunculus acris, R. flammula and the dwarf montane form of marsh marigold Caltha palustris var. radicans.

In the upper parts of Coire nam Beitheach, at over 900m, there are more montane examples with muchPhilonotis seriata and with the rare Saxifraga rivularis as well as the more widespread species.

There are M32b springs throughout the study area, but they are especially common in the high corries,where they form large spreads of vegetation along spring-lines and following the edges of small rills andstreams. M32b occurs throughout the British uplands from Wales northwards, but is rare below about 450mexcept in the far north and west.

M33 Pohlia wahlenbergii var glacialis spring

The dominant plant in these springs is the montane moss Pohlia wahlenbergii var. glacialis (Photograph 19).It has beautiful pale mint-green leaves with a waxy surface and a crisp but pliant texture. They are alwaysbeaded with drops of icy water and are the very essence of coolness. P. w. glacialis grows with smalleramounts of other bryophytes such as Philonotis fontana, Sphagnum denticulatum and Scapania undulata,and the montane snow-bed species Pohlia ludwigii. The cushions of bryophytes are dotted with Deschampsiacespitosa ssp. alpina, Saxifraga stellaris, Epilobium anagallidifolium, Chrysosplenium oppositifolium and, onthe high back wall of Coire nam Beitheach, the rare Saxifraga rivularis (Photographs 18–19).

There are M33 springs in Coire nan Lochan and in the upper part of Coire nan Gabhail as well as in Coirenam Beitheach. This is a high montane community. It occurs only in the higher hills of the Scottish Highlands,in corries and on high slopes where snow lies late in spring. M33 is never common except on the largerhills of the central and eastern Highlands, where in spring and early summer there is a great deal of late-lying snow.

M37 Palustriel la commutata-Festuca rubra spring

This type of spring marks out the places of emergence of base-rich water. There are many examples on MeallMór, and a few in the corries of Bidean and on the slopes of the Aonach Eagach. M37 mires are distinctiveand easy to see. They consist of dripping mats or rounded cushions of Palustriella commutata: a large mosswith orange-gold shoots and tightly-curling leaves. The similar but slightly smaller Cratoneuron filicinum cangrow here too, and some springs consist entirely of this species. There are generally patches of Bryum

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pseudotriquetrum and Scapania undulata. Few vascular plants grow in these springs apart from Festucarubra, which forms a thin sward over the layer of mosses. There are odd shoots of Carex panicea, C. flacca,Viola palustris, Selaginella selaginoides and Chrysosplenium oppositifolium.

M37 mires occur throughout the British uplands from Wales northwards, wherever there are outcrops ofbase-rich rock.

MX Small sedge mire

Although this type of mire is not yet described by the NVC, it is widespread in the uplands of Wales andScoland and almost certainly occurs in northern England too. It has the sedges of the Carex-Sphagnummires M6 with the tall herbs and grasses of the Juncus-Galium rush pastures M23. The sward is made up of Carex echinata, C. nigra, C. panicea, C. viridula ssp. oedocarpa and C. hostiana, interleaved withgrasses such as Agrostis canina, Anthoxanthum odoratum and Holcus lanatus and with mesotrophic herbssuch as Filipendula ulmaria, Angelica sylvestris, Geum rivale, Crepis paludosa, Parnassia palustris,Pedicularis palustris and Trollius europaeus. Over the ground are wet mats of bryophytes made up ofCalliergonella cuspidata, Campylium stellatum, Plagiomnium undulatum and Rhizomnium punctatum.

There are some fine MX mires on the slopes of Meall Mór, where the slopes are flushed with base-rich waterfrom the limestone.

MG9 Holcus lanatus-Deschampsia cespitosa grassland

This distinctive type of vegetation consists of a tall, dense growth of the massed tussocks of Deschampsiacespitosa; their leaves reaching up to about 60–80cm tall and overtopped by conspicuous flowering stemsabout 80–100cm in height. Other plants are quite sparse in what little room is left among the D. cespitosatussocks. They typically include Agrostis capillaris, Anthoxanthum odoratum, Holcus lanatus and scattered plantsof Potentilla erecta, Ranunculus acris, Achillea millefolium, A. ptarmica, Viola palustris, Carex binervis andJuncus effusus. Bryophytes are sparse and inconspicuous, the commonest species being the moss Rhytidiadelphussquarrosus. The flora appears to indicate more or less neutral soils. Grazing is evidently rather light.

MG9 is scarce at this site. There are patches of it among Molinia grassland on level ground in one part ofthe valley floor of Glen Etive, at the south-eastern edge of the site, and among bracken, bramble and shortergrassland near Achnacon in the lower part of the main glen. MG9 is widespread at low altitudes in Britain.

CG10a Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland, Trifolium repens-Luzula

campestris sub-community

These are herb-rich grasslands on well-drained neutral to basic soils. The short, grey-green sward is madeup of Festuca ovina, F. vivipara, Agrostis capillaris, A. canina and Anthoxanthum odoratum and entangledwith the long creeping shoots of Thymus polytrichus. It is enriched with small base-tolerant species includingBellis perennis, Ranunculus acris, Lotus corniculatus, Prunella vulgaris, Carex caryophyllea, Trifolium repens,Achillea millefolium and Plantago lanceolata, as well as the less exacting Potentilla erecta and Galiumsaxatile. There are thin mats of bryophytes through the turf, consisting of species such as Hylocomiumsplendens, Ctenidium molluscum, Rhytidiadelphus squarrosus, Racomitrium lanuginosum, Breutelia chrysocoma,Diplophyllum albicans and Frullania tamarisci.

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CG10a occurs on the slopes of the Fionn Gleann. It is less common in the study area than the damper sub-community CG10b. CG10a occurs widely through the British uplands.

CG10b Festuca ovina-Agrostis capil laris-Thymus polytrichus grassland, Carex pulicaris-Carex

panicea sub-community

This is the most widespread form of Festuca-Agrostis-Thymus grassland in the study area. Like CG10a, it hasa short, tight sward of grasses such as Festuca ovina, F. vivipara, Agrostis capillaris, A. canina,Anthoxanthum odoratum and a little Nardus stricta, matted together with Thymus polytrichus. There are alsomore species of damp, flushed soils such as Carex panicea, C. pulicaris, Selaginella selaginoides, Violapalustris, Cirsium palustre and Succisa pratensis. These grow together with Prunella vulgaris, Ranunculusacris, Campanula rotundifolia, Alchemilla alpina, A. glabra and Linum catharticum. Wefts of bryophytes arewoven through the turf. They consist of species such as Hylocomium splendens, Rhytidiadelphus squarrosus,R. loreus, Dicranum scoparium, Ctenidium molluscum and Scleropodium purum.

CG10b is most extensive over the limestone on Meall Mór (see Photographs 7, 9 and 10), but there areexamples on flushed slopes with free-draining mineral soils throughout the study area. CG10b occursthroughout the British uplands.

CG10c Festuca ovina-Agrostis capil laris-Thymus polytrichus grassland, Saxifraga aizoides-

Ditrichum flexicaule sub-community

This is the most montane form of Festuca-Agrostis-Thymus grassland. Like CG10b, it is a grassland of damp,flushed soils, and the short silvery-green turf of Festuca vivipara, Agrostis canina, A. capillaris, Anthoxanthumodoratum and Nardus stricta is interleaved with Carex panicea, C. pulicaris, C. viridula ssp. oedocarpa,C. hostiana and species such as Selaginella selaginoides, Linum catharticum, Saxifraga aizoides, Geum rivale,Trollius europaeus, Alchemilla alpina, A. glabra, Ranunculus acris, Parnassia palustris, Thalictrum alpinum,Leontodon autumnalis and Persicaria vivipara. There is much Thymus polytrichus, Prunella vulgaris andPlantago lanceolata, and in some examples there is a little Carex bigelowii, Deschampsia cespitosa ssp.alpina or Helictotrichon pratense. There is a rich array of bryophytes including Breutelia chrysocoma, Hylocomiumsplendens, Hypnum lacunosum, Ctenidium molluscum, Calliergonella cuspidata, Ditrichum flexicaule,Plagiobryum zieri, Rhytidiadelphus squarrosus, R. triquetrus, Thuidium tamariscinum and Tortella tortuosa.

We found CG10c on Meall Mór (see Photographs 7 and 9) and on the steep south-facing slopes of AonachEagach. CG10c is widespread though rarely common throughout the Scottish uplands.

CG10 Festuca ovina-Agrostis capil laris-Thymus polytrichus grassland on river shingle

There are some interesting CG10 grasslands on the spreads of river shingle in the Lost Valley of CoireGabhail (Photograph 27), in the Lairig Eilde and in the main glen. Grasses are sparse, and the swardsconsist of assemblages of Thymus polytrichus, Lotus corniculatus, Festuca vivipara and Anthoxanthumodoratum, growing in dense pale-golden carpets of Racomitrium ericoides. There is a fine speckling of afew other species such as Deschampsia flexuosa, Anthoxanthum odoratum, Galium saxatile, Ranunculusacris, Viola riviniana, Plantago lanceolata, Prunella vulgaris and Polygala serpyllifolia, and large untidyrosettes of Digitalis purpurea. Most of these swards are likely to be flooded in winter when the rivers are inspate, and are obviously subject to much disturbance and redistribution of the shingle. This vegetation is

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quite different from the three sub-communities of CG10 mainly because of the very extensive carpets of R. ericoides, but also because grasses are more sparse than Thymus and dicotyledonous herbs.

CG11a Festuca ovina-Agrostis capil laris-Alchemilla alpina grassland, Typical sub-community

This is the drier sub-community of CG11. The bright green silver-edged leaves of Alchemilla alpina growthickly in a short and open sward of Festuca vivipara, Agrostis capillaris, A. canina, Anthoxanthumodoratum, Deschampsia flexuosa and Nardus stricta. There are trailing mats of Thymus polytrichus, and anarray of other species such as Potentilla erecta, Campanula rotundifolia, Viola riviniana, Carex pilulifera, C. flacca, C. bigelowii, Linum catharticum, Lotus corniculatus and Huperzia selago. In most examples thereare a few montane species such as Diphasiastrum alpinum, Sibbaldia procumbens or Gnaphalium supinum.This is a grassland of stony and usually steep ground and there is plenty of space for bryophytes in the thinsward. Many species grow here, including Hylocomium splendens, Racomitrium lanuginosum, R. ericoides,R. fasciculare, Breutelia chrysocoma, Rhytidiadelphus squarrosus, R. loreus, R. triquetrus, Polytrichum alpinumand Scleropodium purum.

CG11a clothes considerable areas of the upper slopes and corrie walls, and also occurs as small patcheswithin mosaics of other grasslands. It is widespread in the Scottish Highlands and also occurs in the Lake District.

CG12 Festuca ovina-Alchemilla alpina-Silene acaulis community

These velvety emerald green swards of Silene acaulis bear vivid pink flowers in summer and are dotted witha great array of small species such as Persicaria vivipara, Alchemilla alpina, A. glabra, Trollius europaeus,Thymus polytrichus, Ranunculus acris, Carex pulicaris, C. capillaris, C. pallescens, Festuca ovina, F. vivipara,Helictotrichon pratense, Plantago lanceolata, Saxifraga aizoides, Viola riviniana, Linum catharticum,Filipendula ulmaria, Geum rivale and Parnassia palustris. The turf is threaded with bryophytes such asHylocomium splendens, Ctenidium molluscum, Hypnum lacunosum, Plagiobryum zieri, Tortella tortuosa,Ditrichum flexicaule, Scleropodium purum, Anoectangium aestivum and Breutelia chrysocoma.

CG12 is an attractive herb-rich community of damp basic soils at moderate to high altitudes. There are manysmall patches of it on the steep limestone slopes of Meall Mór. CG12 is widespread though rarely extensivein the Scottish Highlands.

CG14 Dr yas octopetala-Silene acaulis community

This herb-rich heath forms flowery swards on the steep rock faces and ungrazed ledges of Meall Mór. Thedistinctive and very decorative Dryas octopetala has dark green leaves with silvery scalloped edges, massesof creamy flowers and delicate feathery seed-heads. It grows here with species such as Alchemilla alpina,A. glabra, Thymus polytrichus, Silene acaulis, Trollius europaeus, Carex capillaris, C. pulicaris, Luzulasylvatica, Geum rivale, Geranium sylvaticum, Angelica sylvestris, Filipendula ulmaria, Saxifraga aizoides,S. oppositifolia, Rubus saxatilis, Oxyria digyna, Selaginella selaginoides, Sedum rosea, Deschampsiacespitosa, Festuca vivipara, Helictrotrichon pratense and Linum catharticum. In some patches of CG14 thereare plants of Salix myrsinites. There are cushions and mats of bryophytes on the ledges and hanging overthe rock faces. These are made up of species such as Neckera crispa, Ctenidium molluscum, Tortellatortuosa, Anoectangium aestivum, Orthothecium rufescens, Ditrichum flexicaule, Hylocomium splendens,

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Racomitrium lanuginosum, Scleropodium purum, Breutelia chrysocoma, Hypnum lacunosum and Frullaniatamarisci. CG14 and the Salix myrsinites scrub W20 have similar floras and grade from one to the other.

There are small patches of CG14 on the very steep upper north-eastern slope of Meall Mór. CG14 is oneof the less common types of upland vegetation in Britain. It is confined to ungrazed ledges or lightly-grazedslopes where the rocks are base-rich. It occurs on the Breadalbane hills and, more rarely, on basic outcropsthrough the central and north-west Highlands.

U4a Festuca ovina-Agrostis capil laris-Galium saxati le grassland, Typical sub-community

This is the common acid upland grassland. Almost always grazed short and dense, it has a green sward ofFestuca ovina, F. vivipara, F. rubra, Agrostis capillaris and Anthoxanthum odoratum. Potentilla erecta andGalium saxatile grow among the grasses and their small bright flowers enliven the green herbage in summer.The turf is dotted with a few other species such as Carex binervis, Viola riviniana, V. palustris, Polygalaserpyllifolia, Luzula multiflora, Blechnum spicant and Conopodium majus. There are dense mats ofbryophytes, made up of species including Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphussquarrosus, R. loreus, Hylocomium splendens, Scleropodium purum and Dicranum scoparium.

U4a is common in the study area, on well-drained mineral soils on moderate to steep slopes. It is ubiquitousin the British uplands, though there is more of it on hills with a long history of sheep grazing.

U4b Festuca ovina-Agrostis capil laris-Galium saxati le grassland, Holcus lanatus-Trifolium

repens sub-community

This type of Festuca-Agrostis-Galium grassland usually covers richer soils than those with the Typical sub-community U4a. Whether they are derived from more nutrient-rich rocks, or have had farmyard manure orlime added, the effect on the flora is similar. The sward of Festuca ovina, F. rubra, Agrostis capillaris,Anthoxanthum odoratum and Carex binervis is enriched by species such as Cynosurus cristatus, Holcuslanatus, Achillea millefolium, Prunella vulgaris, Trifolium repens, Plantago lanceolata and Ranunculus acris.The common species of acid grassland Potentilla erecta, Galium saxatile, Polygala serpyllifolia andConopodium majus grow here too but can be scarcer than in U4a. There are many bryophytes scattered inthe turf, including Rhytidiadelphus squarrosus, R. loreus, Hypnum jutlandicum, Thuidium tamariscinum,Hylocomium splendens, Pleurozium schreberi and Scleropodium purum.

U4b grassland covers the fields in the floor of the main glen betwen Achnambeitheach and Achtriochtan. It is common throughout the British uplands, especially over richer rocks such as basalt and where there hasbeen some agricultural improvement in the past.

U4d Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Luzula multiflora-Rhytidiadelphus

loreus sub-community

This is a grassland of cool, damp slopes with well-drained but moist soils. It covers many of the north-facingand east-facing lower slopes in the study area. From a distance it looks just like the Typical sub-communityU4a, but Agrostis canina is more common than A. capillaris in the sward of Festuca vivipara, F. rubra,Anthoxanthum odoratum and Deschampsia cespitosa, and there is much Carex binervis and Luzulamultiflora. The usual small grassland herbs grow here: Potentilla erecta, Galium saxatile, Polygala

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serpyllifolia, Viola riviniana and Euphrasia officinalis. There may be a sprinkling of Alchemilla alpina andthe turf may be sprigged with Vaccinium myrtillus. There can be a few other heathland species such asCornus suecica, Blechnum spicant, Vaccinium vitis-idaea, Lycopodium clavatum and L. annotinum(Photograph 41). U4d has a more mossy turf than either U4a or U4b. There are red, golden and greenconspicuous mats and patches of species such as Rhytidiadelphus loreus, R. squarrosus, Hylocomiumsplendens, Hypnum jutlandicum, Pleurozium schreberi, Polytrichum commune, Dicranum scoparium, Thuidiumtamariscinum, Barbilophozia floerkei and Tritomaria quinquedentata, and there can be hummocks ofSphagnum capillifolium.

As the flora suggests, U4d is usually derived by grazing from some kind of damp heath. Indeed it is commonto see spreads of U4d on slopes which are open to grazing and Calluna-Vaccinium-Sphagnum heath H21or Vaccinium-Rubus heath H22 on ungrazed ledges and cliffs above. On the north-facing slopes of the mainglen there are clumps and patches of the oceanic liverworts Herbertus aduncus, Pleurozia purpurea, Myliataylorii, Scapania gracilis, Plagiochila spinulosa, Bazzania tricrenata, Anastrepta orcadensis and Lepidoziapearsonii in the swards of U4d. These are almost certainly relicts from the days when these slopes wereclothed with dwarf-shrub heaths or woodland.

U4d is widespread in the uplands of northern Britain.

U4e Festuca ovina-Agrost is capi l laris-Galium saxati le grassland, Vaccinium myr t i l lus-Deschampsia flexuosa sub-community

This is the most montane form of Festuca-Agrostis-Galium grassland. It is a community of thin, well-drained,stony soils such as the ribs of raised ground between the incised gullies on the upper slopes. U4e has athin, short, open, silver-green turf of Festuca vivipara, Agrostis capillaris, A. canina, Anthoxanthum odoratumand Nardus stricta growing through a dense grey underlay of Racomitrium lanuginosum. There are speciessuch as Galium saxatile, Potentilla erecta, Viola riviniana and Carex panicea, and short, grazed shoots ofCalluna vulgaris or Vaccinium myrtillus or both. There are also clumps and patches of other bryophyes suchas Hylocomium splendens, Hypnum jutlandicum and Frullania tamarisci.

U4e is common on the upper slopes of the study area, especially on the exposed slopes facing south andwest: the opposite to the Luzula-Rhytidiadelphus sub-community U4d. The two can form intimate mosaics onthe steep slopes corrugated by gullies, with U4d on the sheltered sides of the gullies and U4e on theexposed ribs or ridges in between. U4e is common in the British uplands from Wales northwards.

U4eA Festuca ovina-Agrost is capi l lar is-Gal ium saxat i le grass land, Vaccinium myr t i l lus-Deschampsia flexuosa sub-community, Alchemilla alpina variant

This type of vegetation, not yet described by the NVC, has a grassy sward with much Alchemilla alpina. It resembles CG11 but lacks the calcicole species and Thymus polytrichus which occur there. It is probablybest thought of as a Racomitrium-Alchemilla form of U4. U4eA consists of an open turf of Festuca vivipara,Agrostis capillaris, A. canina, Anthoxanthum odoratum, Deschampsia flexuosa and Nardus stricta, thickly-grown with Alchemilla alpina and set in silvery carpets of Racomitrium lanuginosum. There is a fine sprinklingof species such as Potentilla erecta, Viola riviniana, Galium saatile, Vaccinium myrtillus, Euphrasia officinalisagg., Luzula sylvatica and Solidago virgaurea, and tufts of bryophytes including Hypnum jutlandicum,Rhytidiadelphus loreus, Hylocomium splendens, Pleurozium schreberi and Polytrichum formosum.

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U4eA is common in the glens and corries of the study area, on steep slopes with thin, well-drained rockysoils. It occurs throughout the western Highlands and in the Inner Hebrides.

U4F Festuca ovina-Agrostis capil laris-Galium saxati le grassland, provisional Fi l ipendulaulmaria sub-community

Another community not yet described in the NVC, this is a herb-rich Festuca-Agrostis-Galium grassland. As inthe other sub-communities, the sward is made up of Festuca ovina, F. vivipara, F. rubra, Anthoxanthumodoratum, Agrostis capillaris, A. canina, Holcus lanatus, Helictotrichon pratense, Luzula multiflora and Carex binervis. Instead of the Thymus polytrichus and small herbs of CG10 and CG11, U4F is enrichedwith larger, mesotrophic species such as Ranunculus acris, Geum rivale, Filipendula ulmaria, Trollius europaeus,Angelica sylvestris, Cardamine pratensis, Alchemilla glabra, Centaurea nigra, Galium verum, Heracleumsphondylium, Carex pallescens and Rhinanthus minor. These grow here with smaller plants such as Carexpulicaris, Linum catharticum, Saxifraga aizoides, Trifolium repens, Parnassia palustris, Prunella vulgaris,Carex hostiana and C. flacca, as well as more typical U4 species such as Potentilla erecta, Galium saxatile,Viola riviniana, Polygala serpyllifolia, Conopodium majus and Plantago lanceolata. The ground under thevascular plants is moist and shaded, and there are mats of bryophytes such as Hylocomium splendens,Scleropodium purum, Rhytidiadelphus loreus, R. squarrosus, Thuidium tamariscinum and Breutelia chrysocoma.

U4F is widespread on the steep northern and north-eastern slopes of Meall Mór, in mosaics with other typesof species-rich grassland (see Photographs 7, 9 and 10). This is its typical habitat. U4F occurs on steep N-NE-facing slopes with deep, flushed mineral soils and moderate to light grazing in several places in thewestern Highlands and the Inner Hebrides. It is not described in the NVC but we have generally referred to it as U4F where we have found it in surveys: the letter ‘F’ (in upper case to avoid confusion with formalsub-communities) continues on from ‘e’ in the series of NVC sub-communities, and also stands for both‘Flushed’ and ‘Filipendula’.

U5a Nardus stricta-Galium saxati le grassland, Species-poor sub-community

Many hectares of the hillsides are clothed with the pale white-blond swards of this type of acid grassland.The thick turf of Nardus stricta is threaded through with a little Festuca ovina, F. vivipara, Agrostis capillaris,A. canina, Anthoxanthum odoratum and Carex binervis. It is starred in summer with the small bright flowersof Potentilla erecta, Galium saxatile, Viola riviniaina, V. palustris and Polygala serpyllifolia. The sward isdotted with the dark rosettes of Juncus squarrosus and the green sprigs of Vaccinium myrtillus. There is adense mass of bryophytes made up of large common species such as Pleurozium schreberi, Rhytidiadelphusloreus, R. squarrosus, Hylocomium splendens, Hypnum jutlandicum, Dicranum scoparium, Polytrichum commune,P. strictum and Thuidium tamariscinum.

U5a is common throughout the British uplands, especially in the hills of Wales, northern England andsouthern Scotland where there has been a long history of heavy grazing by ewes and lambs. These animalsavoid the tough leaves of Nardus, enabling it to spread at the expense of more palatable species.

U5b Nardus s tr ic ta-Gal ium saxat i le grass land, Agrost is canina-Polytr ichum communesub-community

This is a damp form of Nardus grassland. It can be really wet underfoot; even boggy, with big swellingcushions of Sphagnum capillifolium, S. papillosum, Polytrichum commune and P. strictum interrupting the pale

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swards of Nardus stricta. There is much Juncus squarrosus, Agrostis canina and Carex panicea in the sward,together with Agrostis capillaris, Anthoxanthum odoratum, Festuca ovina and F. vivipara. The turf is speckledwith small plants such as Galium saxatile, Viola palustris, Potentilla erecta, Polygala serpyllifolia, Pedicularissylvatica, Narthecium ossifragum, Eriophorum angustifolium, Trichophorum cespitosum and Blechnumspicant. Among these are mats and wefts of bryophytes such as Hypnum jutlandicum, Pleurozium schreberi,Rhytidiadelphus loreus, Plagiothecium undulatum, Hylocomium splendens, Thuidium tamariscinum andliverworts such as Lophocolea bidentata, Mylia taylorii, Scapania undulata and S. gracilis.

U5b is common on concave slopes and in hollows where ill-drained mineral soils are kept constantly moist.It is the characteristic vegetation of grassy channels among blanket bogs and wet heaths. It also occurs upin the high corries in wet hollows and on steps and shelves among montane heaths and snow-beds,wherever deep irrigated soil has been able to accumulate. U5b is common throughout the British uplandsfrom Wales northwards.

U5c Nardus stricta-Galium saxatile grassland, Carex panicea-Viola riviniana sub-community

U5c is a herb-rich Nardus grassland. The dense tussocky straw-coloured sward of N. stricta, Agrostiscapillaris, A. canina, Anthoxanthum odoratum, Festuca ovina, F. vivipara, F. rubra and Deschampsiaflexuosa conceals a rich assemblage of small plants such as Ranunculus acris, Carex pulicaris, C. panicea,C. viridula ssp. oedocarpa, Thalictrum alpinum, Linum catharticum, Alchemilla alpina, A. glabra, Thymuspolytrichus, Geum rivale, Filipendula ulmaria, Trollius europaeus, Selaginella selaginoides and Geum rivale,as well as less demanding species such as Viola riviniana, V. palustris, Campanula rotundifolia, Potentillaerecta, Galium saxatile, Rumex acetosa, Luzula multiflora, Narthecium ossifragum, Carex binervis and C. nigra. Among these plants are creeping shoots and clumps of bryophytes such as Calliergonellacuspidata, Hylocomium splendens, Rhytidiadelphus loreus, R. squarrosus, R. triquetrus, Hypnum lacunosum,Hylocomium splendens, Racomitrium fasciculare and Isothecium myosuroides var. brachythecioides.

U5c is not common in the study area. There are scattered patches on Meall Mór and in Coire namBeitheach. In the British uplands it occurs widely from Wales northwards, but is local and confined to placeswith base-rich rock or base-rich flushing. Only in the Breadalbanes is it at all extensive.

U5e Nardus stricta-Galium saxati le grassland, Racomitrium lanuginosum sub-community

This is the most montane form of Nardus grassland. Unlike the other sub-communities which are producedand maintained by heavy grazing, some U5e might be near-natural. The sward is open and sparse. Thewhite clumps of Nardus stricta are interspersed with bushes of Calluna vulgaris or Empetrum nigrum ssp.hermahroditum, and with sedges including C. panicea and C. pilulifera. They are set in a silvery carpet ofRacomitrium lanuginosum, which may be dotted with montane species such as Carex bigelowii, Salixherbacea, Diphasiastrum alpinum and Polytrichum alpinum and with a ‘frosting’ of lichens such as Cladoniaportentosa, C. uncialis, C. furcata, C. rangiferina, Coelocaulon aculeatum, Cetraria islandica and Alectorianigricans. Indeed, some of the U5e on the summit ridge of Meall Mór is so lichen-rich that the turf lookswhite from a distance and is distinctively crisp underfoot (see Photograph 4). In some of the patches of U5ethere are a few wet heath species such as Narthecium ossifragum or Trichophorum cespitosum. There aresparse wisps or tufts of other bryophytes such as Hypnum jutlandicum, Pleurozium schreberi, Hylocomiumsplendens, Rhytidiadelphus loreus, Frullania tamarisci, Anastrepta orcadensis and Pleurozia purpurea.

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We found U5e on the summit ridge of Meall Mór, in the high corries of Bidean and on the upper slopesand summit ridges. It occurs throughout the British uplands from Wales northwards, but is most extensive inthe western Highlands and in the Hebrides.

U6a Juncus squarrosus-Festuca ovina grassland, Sphagnum sub-community

U6a is a Juncus squarrosus bog of level ground on hill tops and plateaux, and the summits of broad ridgeswhere peat has been able to accumulate. It has a dense, dark green sward of Juncus squarrosus; its glossyrosettes set in a matrix of Festuca vivipara, Agrostis canina, Eriophorum angustifolium, Deschampsiaflexuosa, Carex binervis and C. panicea. These plants grow through rich red and gold patches ofSphagnum capillifolium, S. papillosum, S. fallax, Polytrichum commune, Hypnum jutlandicum, Hylocomiumsplendens, Plagiothecium undulatum and Pleurozium schreberi. The turf is sprinkled with species such asGalium saxatile, Potentilla erecta, Narthecium ossifragum, Polygala serpyllifolia, Pedicularis sylvatica, Violapalustris and Leontodon autumnalis.

U6a occurs on the summit of Meall Mór. It is common throughout the British uplands, but is most extensivein Wales, northern England and southern Scotland, where there has been heavy grazing over many years.

U6c Juncus squarrosus-Festuca ovina grassland, Vaccinium myr ti l lus sub-community

This is a rush heath in which the firm tussocky sward of Juncus squarrosus is densely sprigged with smallshoots of Vaccinium myrtillus. There is also a little Festuca ovina, Agrostis capillaris, A. canina, Anthoxanthumodoratum and Nardus stricta set in the spaces around the rosettes of the Juncus squarrosus. There is afreckling of small plants such as Galium saxatile, Potentilla erecta, Viola riviniana and Vaccinium vitis-idaea,and mats of bryophytes including Rhytidiadelphus squarrosus, R. loreus, Hypnum jutlandicum, Pleuroziumschreberi, Hylocomium splendens, Polytrichum commune.

There are patches of U6c among grasslands on the slopes of Meall Mór. It is common throughout the British uplands except in the north-west Highlands.

U6d Juncus squarrosus-Festuca ovina grass land, Agrost is capi l lar is-Luzula mul t i f lora

sub-community

In this type of Juncus rush heath, the sward of grasses, sedges and heath rush is very mixed. It is made upof Juncus squarrosus, Festuca ovina, Agrostis capillaris, A. canina, Anthoxanthum odoratum, Nardus stricta,Deschampsia flexuosa and Carex binervis. The plants may grow together in even mixtures or as discreteclumps and patches, but always forming a patterned, variegated turf. In this are rooted small plants such asGalium saxatile, Potentilla erecta, Viola riviniana, Luzula multiflora, Vaccinium myrtillus and Polygalaserpyllifolia. There is a sparse array of bryophytes made up of large common mosses such as Hylocomiumsplendens, Hypnum jutlandicum, Pleurozium schreberi, Rhytidiadelphus squarrosus, R. loreus and Polytrichumcommune.

We found U6d among mosaics of grasslands on the slopes of Meall Mór. U6d is widespread throughoutthe British uplands, and is especially common on the grazed, grassy hills of Wales, northern England andsouthern Scotland.

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U6Cv Juncus squarrosus-Festuca ovina grassland, provisional Calluna vulgaris sub-community

This is yet another type of vegetation which is not described by the NVC. It is a heathy mixture of the tightgreen rosettes of Juncus squarrosus and short dark brown bushes of Calluna vulgaris, together with shoots ofVaccinium myrtillus and clumps of Empetrum nigrum ssp. nigrum. Among these are grasses such as Nardusstricta, Agrostis capillaris and Festuca ovina. U6Cv is generally, though not always, wet and boggy withmuch Sphagnum capillifolium as well as mosses such as Hylocomium splendens, Pleurozium schreberi,Hypnum jutlandicum, Rhytidiadelphus loreus and Plagiothecium undulatum.

There are small patches of U6Cv in Coire na Cuilce on Meall Mór. We have seen this type of vegetationin many parts of the British uplands. It appears to be a stage in recovery from grazing, with the rush heathgradually being colonised by dwarf shrubs.

U6R Juncus squarrosus-Festuca ovina grassland, provisional herb-rich sub-community

There is no place for this type of upland vegetation in the NVC, although it was recognised in the 1950sin the Breadalbane hills by McVean & Ratcliffe (1962). A rush heath of damp, flushed soils, it has a darkgreen, close-set sward of Juncus squarrosus. The stiff rosettes of the rush shelter many small base-tolerantspecies such as Thalictrum alpinum, Ranunculus acris, R. flammula, Geum rivale, Carex pulicaris, C. panicea,C. viridula ssp. oedocarpa, Crepis paludosa, Leontodon autumnalis and Saxifraga stellaris. There are alsograsses such as Deschampsia cespitosa, Agostis canina and Anthoxanthum odoratum. The bryophytes, too,show that the soils are moist and flushed with nutrient-rich water. They include Calliergonella cuspidata,Calliergon sarmentosum, Rhizomnium punctatum, Plagiomnium undulatum, Hylocomium splendens,Drepanocladus revolvens, Riccardia multifida and Scapania undulata.

There are small patches of U6R on the limestone slopes of Meall Mór. This type of vegetation occurs quitewidely in the British uplands from Wales northwards, but seems to be common only across the Breadalbanehills.

U7a Nardus str ic ta-Carex bigelowii grass heath, Empetrum nigrum hermaphroditum-

Cetraria islandica sub-community

These are pale golden-blond patches of vegetation which occur in hollows on the summit ridges and in the high corries, where snow lies late in spring and drainage is held up by heavy soils or impervious rocks.The short, tufted, even sward of Nardus stricta is interleaved with Trichophorum cespitosum and Carexbigelowii and patched with Empetrum nigrum ssp. hermaphroditum, Vaccinium myrtillus and V. uliginosum.It is sprinkled over with species such as Diphasiastrum alpinum, Huperzia selago, Narthecium ossifragum,Succisa pratensis, Carex pilulifera, Viola palustris and Deschampsia cespitosa ssp. alpina. There is a richarray of bryophytes: silvery patches of Racomitium lanuginosum, red hummocks of Sphagnum capillifoliumand golden-green wefts of Hypnum jutlandicum, Hylocomium splendens, Rhytidiadelphus loreus, R. squarrosus, Polytrichum commune, P. alpinum and Pleurozium schreberi. There are liverworts such asDiplophyllum albicans, Barbilophozia floerkei, Ptilidium ciliare and the western, oceanic Anastreptaorcadensis, Pleurozia purpurea, Scapania gracilis and Bazzania tricrenata. Over these is a thin dusting oflichens such as Cetraria islandica, Cladonia portentosa, C. uncialis, C. furcata, C. rangiferina andCoelocaulon aculeatum.

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U7a is common in the high corries and on the ridges of Bidean and Sgorr nam Fiannaidh. It is widespreadthrough the Scottish Highlands, picking out the places where snow lies late and where the soils are moistthroughout the year.

U7b Nardus stricta-Carex bigelowii grass heath, Typical sub-community

Less distinctive in its flora than the other two sub-communities of U7, this is a snow-bed grassland of well-drained slopes in the high corries. The patches typically form a crescent-shaped necklace of bedraggled,straw-pale vegetation around the upper slopes of a corrie or around the base of a cliff. U7a is a wettergrassland and U7c is more herb-rich. This, in contrast, is dry and heathy. The turf of Nardus stricta, Carexbigelowii and a few other grasses and sedges such as Anthoxanthum odoratum, Deschampsia flexuosa,Agrostis canina, Festuca vivipara and Carex pilulifera, is thickly grown with shoots of Vaccinium myrtillusand V. vitis-idaea and with clumps of V. uliginosum and Empetrum nigrum ssp. nigrum. Many other speciesgrow here, including Potentilla erecta, Galium saxatile, Huperzia selago, Diphasiastrum alpinum andBlechnum spicant. The mats of bryophytes are made up of Racomitrium lanuginosum, Hylocomiumsplendens, Pleurozium schreberi, Polytrichum alpinum, Plagiothecium undulatum, Rhytidiadelphus loreus andR. squarrosus. Cetraria islandica grows in most patches of U7b, together with Cladonia uncialis, C. portentosaand C. furcata.

There are patches of U7b in the high corries of Bidean and on the ridge of Sgorr nam Fiannaidh. U7b iswidespread throughout the Scottish Highlands, occupying fairly free-draining soils where snow lies late in spring.

U7c Nardus stricta-Carex bigelowii grass heath, Alchemilla alpina-Festuca ovina sub-community

This is the usual type of Nardus snow-bed where the soils are flushed with base-rich water. It is also the mostcommon at moderate altitudes and in the far west Highlands and in the Hebrides. Though Nardus stricta isstill the dominant plant, giving its characteristic bleached-blond colour to the vegetation, the sward is moremixed than it is in U7a and U7b. There can be Anthoxanthum odoratum, Festuca vivipara, Agrostis canina,Deschampsia flexuosa and D. cespitosa as well as Carex bigelowii and C. pilulifera. There are usuallyflowery clumps of Alchemilla alpina growing with Galium saxatile, Potentilla erecta, Vaccinium myrtillus,Empetrum nigrum ssp. hermaphroditum, Diphasiastrum alpinum, Huperzia selago and Viola riviniana.Among these are wefts of bryophytes such as Racomitrium lanuginosum, R. ericoides, Hypnum jutlandicum,Hylocomium splendens, Rhytidiadelphus loreus, Dicranum scoparium and Polytrichum alpinum, and patchesof Cetraria islandica, Cladonia portentosa, C. uncialis and C. furcata.

U7c is the least common of the three types of U7 in the study area. We found it on Sgorr nam Fiannaidhand in Coire nam Beitheach, in hollows and on high slopes where the underlying rocks and the irrigatingwater are base-rich. U7c occurs throughout the western and central Highlands and in the Hebrides.

U8 Carex bigelowii-Polytrichum alpinum heath

This is the least extensive of the montane heaths in the study area. There is a tiny patch on the summit ofBidean, and fragments on Sgorr nam Fiannaidh, Stob Coire Leith and Meall Dearg. The pale grey-greeneven sward of Carex bigelowii, dotted in summer with its tight black flowers, grows through a thick carpetof Polytrichum alpinum, Dicranum fuscescens, Kiaeria starkei, Racomitrium lanuginosum, R. heterostichumand R. fasciculare. Very little else grows here beyond a sprinkling of Alchemilla alpina, grasses such as

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Deschampsia flexuosa, Agrostis capillaris or Festuca vivipara and scattered tufts of other bryophytes such asAnthelia juratzkana, Oligotrichum hercynicum and Conostomum tetragonum.

U8 is a community of high hilltops where snow lies far into spring. Many but not all patches are in placeswhere drainage is slow, or where water is channelled from melting snow for weeks at a time and whichmay well have standing water well into summer. In these situations U8 is almost a Carex bigelowii mire. The vegetation is equally common on convex summits and on steep slopes where the soils are well-drained.U8 occurs throughout the eastern and central Highlands. There are outlying patches as far south and westas Ben Lui, but most of it is on the high plateaux of hills such as Creag Meagaidh, Ben Alder, the hills aroundGlen Clova, and the Cairngorms and Lochnagar.

U10a Carex bigelowii-Racomitrium lanuginosum moss heath, Galium saxatile sub-community

We found this form of Carex-Racomitrium heath only around the summit of Bidean and on rocky ground onthe steep slopes to the south-east of Stob Coire Sgreamhach. The thick silver-grey mat of Racomitriumlanuginosum which carpets the ground is pricked through by Carex bigelowii, Deschampsia flexuosa,Festuca vivipara and Nardus stricta. There are trailing masses of Galium saxatile, short shoots of Vacciniummyrtillus, rosettes of Saxifraga stellaris and tufts of other bryophytes such as Polytrichum alpinum, Dicranumfuscescens, Rhytidiadelphus loreus, Racomitrium sudeticum, R. fasciculare and Pleurozium schreberi.

U10a is the form of Carex-Racomitrium heath most commonly encountered on hills in the parts of the Britishuplands where the climate is slightly less bleak and inhospitable than is usual for this type of vegetation. Thisis certainly not the case for Bidean nam Bian: a high stony summit more often than not engulfed in cloudand subject to bitter freezing wind, driving rain and intense frosts as well as to scorching sunshine anddrought on the rare clear summer days. U10a is the most common sub-community of U10 on the hills ofnorth Wales, the Lake District and south and west Scotland. This form of U10, in which vascular plants covermuch of the ground and the moss Racomitrium lanuginosum is rather patchy, may be the result of the mosscarpet being damaged by grazing and trampling or by eutrophication from animal droppings or fromatmospheric pollution. It may occur on the well-trodden summit and ridge of Bidean because of tramplingby people and animals, because of the nutrient enrichment from sheep dung and urine or because of nitratedeposition from mist as well as rain and snow.

U10b Carex bigelowii-Racomitrium lanuginosum moss heath, Typical sub-community

Most of the Carex-Racomitrium heath in the study area is of this type. It is a community of the most severelyexposed hill summits and ridges which are blown clear of snow in winter (see Photographs 13, 26 and 34).The growing-season is not dramatically shortened as it is in the snow-beds, but the plants which grow inthese heaths have to be able to endure huge variations in temperature from the bitterest winter cold to theburning heat of summer, repeated thawing and freezing in the winter, drenching cold rain, mist and blowinglow cloud and raging winds. All of these make the soils unstable and prevent any but the smallest plantsgaining a foothold. One of the species which can tolerate these inhospitable conditions is the mossRacomitrium lanuginosum. It spreads over the ground in a thick, felted, silver-green blanket (Photograph 24)in may places with its curling shoot-tips pointed away from the prevailing wind. Though it must provide onlymeagre shelter, a number of other species take root in the Racomitrium carpet. Many of them are low,creeping plants with long rhizomes or prostrate stems giving them a firm hold on the ground: species suchas Carex bigelowii, Salix herbacea, Vaccinium myrtillus and Diphasiastrum alpinum. Then there are other

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small plants such as Juncus trifidus, Huperzia selago, Alchemilla alpina, Empetrum nigrum ssp.hermaphroditum, Carex pilulifera, Nardus stricta and Deschampsia flexuosa. Among the Racomitriumlanuginosum there can be a few tufts or clumps of other bryophytes such as Polytrichum alpinum,Rhytidiadelphus loreus, Hypnum jutlandicum, Pleurozium schreberi, Dicranum fuscescens and Racomitriumsudeticum. There may be quite a scattering of lichens too, including species such as Cetraria islandica,Ochrolechia frigida, Cladonia uncialis, C. furcata, C. subcervicornis and C. rangiferina.

On the summit of Stob Coire nan Lochan there is Racomitrium heath in which the dominant species is R. ericoides rather than R. lanuginosum.

U10b occurs on Meall Mór, on Sgorr nam Fiannaidh, along the Aonach Eagach to Meall Dearg, on themain ridge of Bidean from An t-Sron to Stob Coire Sgreamheach and on the ridges of Aonach Dubh, Gearr Aonach, Beinn Fhada, Buachaille Etive Beag and Buachaille Etive Mór. U10b occurs throughout theScottish Highlands and in Galloway and the Lake District. It is also in north Wales, but is very rare there.

U10c Carex bigelowii-Racomitrium lanuginosum moss heath, Silene acaulis sub-community

This is a herb-rich form of Carex-Racomitrium heath. The patches on the summit of Bidean nam Bian and theridge leading out to Stob Coire nan Lochan – the only part of the study area where we found U10c – arenot sensationally species-rich, but are sufficiently so to be classed as this sub-community. The carpet ofRacomitrium lanuginosum is quite open, with patches of the moss scattered over rocky ground. There arevivid green cushions of Silene acaulis, mats of Armeria maritima and shoots of Persicaria vivipara andRanunculus acris, together with a sprinkling of Festuca vivipara, tufts of Polytrichum alpinum and lichens suchas Ochrolechia frigida, Cladonia subcervicornis and C. portentosa.

U10c is most common on the high summits of the north-west Highlands, where it can cover many hectaresin green herb-rich carpets. There are outlying examples over the base-rich rocks of the Breadalbanes.

Snow-bed vegetation

Almost all the vegetation on the high slopes of Coire nan Lochain, Coire Gabhail and, especially, Coirenam Beitheach, owes its existence and special characteristics to late-lying snow (Photograph 23). Theseupper slopes are composed largely of loose gravel and spreads of boulders, constantly shifting andchanging and irrigated by melt-water and by small rills as well as by rain. Many of the hollows in the corriefloors fill up with water after heavy rain and become deep but very short-lived pools. Because of this thevegetation is extraordinarily complicated, with much variation on a small scale. Although the NVCcategories can be recognised, only some of the vegetation actually fits neatly into one or another. Most ofit is intermediate vegetation with characteristics of more than one type, and there are extensive mosaicswhere the individual patches of distinct types of vegetation are less than a metre square. The NVC accountsof the snow-beds are themselves inadequate to describe the actual variation in these types of vegetation; afact which is freely admitted by the authors (Rodwell et al. 2000). This should be borne in mind whenreading the following accounts of the snow-bed communities U11, U12, U13b and U14. The accounts arean attempt to describe the range of variation which exists in the high corries by reference to the NVC noda.

U11a Polytrichum sexangulare-Kiaeria starkei snow-bed, Typical sub-community

Most of the moss-dominated snow-bed vegetation in the upper corries belongs to this type or is clearly

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related to it. The dark green mossy carpets, over ground which is generally steep and rocky, are made upof any of quite a range of bryophytes. They grow in fairly pure swards or as intricate mixtures. Some patchesconsist of Kiaeria starkei, K. falcata, K. blyttii, Polytrichum alpinum, Racomitrium lanuginosum, Oligotrichumhercynicum, Conostomum tetragonum, Diplophyllum albicans and Barbilophozia floerkei, growing invariegated sheets of many shades of green. These are studded with plants such as Gnaphalium supinum,Galium saxatile, Saxifraga stellaris, Carex bigelowii, Deschampsia cespitosa ssp. alpina, D. flexuosa,Luzula spicata, Sibbaldia procumbens, Huperzia selago, Vaccinium vitis-idaea, Cryptogramma crispa andAlchemilla alpina. In the layer of bryophytes there are small tufts and wisps of species such as Andreaeaalpina, Oedopodium griffithianum, Anthelia juratzkana, Marsupella emarginata, M. sphacelata, M. alpinaand Pleurocladula albescens and a few oceanic or western plants such as Bazzania tricrenata, Mylia tayloriiand Anastrepta orcadensis. In upper Coire nam Beitheach the rare moss Brachythecium glaciale grows inthis type of vegetation. These patches of vegetation are a reasonable fit for the NVC type U11a.

There is just as much vegetation, especially among boulders or in earthy gullies, in which the Kiaeriaspecies, Polytrichum alpinum, Oligotrichum hercynicum or Conostomum tetragonum occur in pure patchestogether with the characteristic vascular plants, or where the dominant bryophytes are Racomitriumheterostichum or R. fasciculare, or Barbilophozia floerkei, Diplophyllum albicans or Andreaea alpina.According to Rodwell (1992), some of these could be acommodated in the Species-Poor sub-communityU11b. However, they are not species-poor at all, and many patches are in fact more species-rich than theTypical sub-community as described in the NVC. There are also patches of vegetation on eroded or gravellyground where the typical vascular plants of U11a occur in a thin and open sward with very few of themosses and liverworts. One of the most common of these species is Luzula spicata, which can grow inalmost pure lawns covering several square metres. All of these are clearly mossy snow-beds, but the rangeof variation is very great.

Rothero (1991), in an extensive study of Scottish snow-beds, suggested that U11 could be more sensiblydivided into three sub-communities: a Typical one, a hepatic-rich one with much Nardia scalaris,Barbilophozia floerkei, Cephalozia bicuspidata and Pleurocladula abescens, and one with Racomitriumheterostichum. These categories would get closer to accommodating the range of variation which exists inthe high corries of Bidean.

U11 snow-beds have been recorded throughout the Scottish Highlands, but are most extensive in the coldereastern and central hills. The community is rare in the north-west where winters are milder, and has not beenrecorded south of the Highlands.

Pohlia ludwigii snow-bed

This is another type of vegetation which Rodwell (1992) mentions briefly, saying that this, too, could beconsidered a species-poor form of U11. The Pohlia ludwigii snow-beds on Bidean, however, are not reallythat similar to U11. Indeed Rothero (1991), in his much more comprehensive study, considered that thePohlia ludwigii snow-beds should be classed as a new community. There are big patches of this type ofvegetation on the upper slopes of the eastern arm of Coire nam Beitheach, on almost vertical gravel. Thesegravels are saturated with water, which makes them desperately slippery and unstable. The vegetationconsists of bright green lawns of P. ludwigii, dotted with Deschampsia cespitosa ssp. alpina, Saxifragastellaris, Chrysosplenium oppositifolium, Cochlearia pyrenaica ssp. alpina and bryophytes such as Kiaeriastarkei, K. falcata, Racomitrium lanuginosum, Pohlia wahlenbergii var. glacialis and Oedopodium

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griffithianum. The rare Saxifraga rivularis and S. cernua also occur in this type of vegetation on the upperslopes of the corrie.

Pohlia ludwigii snow-beds have not been comprehensively recorded. They certainly occur in the Cairngormsand on Ben Nevis (Rothero 1991) and they are probably thinly-scattered among the higher hills of the centraland eastern Highlands.

U12a Salix herbacea-Racomitrium heterostichum snow-bed, Silene acaulis-Luzula spicata sub-community

This attractive assemblage of plants is very much like the Festuca-Alchemilla-Silene dwarf-herb communityCG12 but with more montane species. It may be an altitudinal replacement for it. U12a is characteristic ofhollows, banks and gully-sides at high altitudes where snow lies very late, and where the underlying rocksor irrigating waters appear to be at least moderately base-rich. It consists of a patchy but colourful anddecorative sward of Silene acaulis, Carex bigelowii, Luzula spicata, Thymus polytrichus, Alchemilla alpina,Nardus stricta, Selaginella selaginoides, Huperzia selago, Anthoxanthum odoratum, Deschampsia cespitosassp. alpina, D. flexuosa, Thalictrum alpinum, Gnaphalium supinum, Persicaria vivipara, Juncus trifidus, J. triglumis, Sibbaldia procumbens, Cerastium arcticum, Saxifraga hypnoides and S. stellaris. Among thesespecies grows a great array of bryophytes such as Andreaea alpina, Anthelia juratzkana, Conostomumtetragonum, Pleurocladula albescens, Oligotrichum hercynicum, Racomitrium lanuginosum, R. heterostichum,R. ericoides, R. fasciculare, Kiaeria falcata, K. starkei, K. blyttii, Barbilophozia floerkei, Diplophyllumalbicans, Scapania undulata, Gymnomitrion obtusum and Ctenidium molluscum. There are obvioussimilarities with the Polytrichum-Kiaeria snow-bed U11 and with the Alchemilla-Sibbaldia snow-bed U14.Indeed, there is much vegetation intermediate between these three types.

We found U12a in both branches of Coire nam Beitheach, where the emerald green cushions of Sileneacaulis stand out vividly against the dull tones of the other montane grasslands, heaths and snow-beds.U12a occurs on the Cairngorms and on the higher hills of the central and western Highlands.

U12b Salix herbacea-Racomitrium heterostichum snow-bed, Gymnomitrion concinnatumsub-community

This type of vegetation is generally so utterly different from U12a that it is hard to imagine how they havecome to be classified as forms of one community. U12b looks from a distance like a sprinkling of smallvascular plants on dark-coloured soil. Close inspection, however, reveals that what looks like bare soil is infact a dense tight crust of diminutive liverworts. Species such as Gymnomitrion concinnatum, Nardiacompressa, N. scalaris, Anthelia juratzkana, A. julacea, Marsupella emarginata, M. sphacelata and M. alpina form a greyish mat which is dotted with plants such as Salix herbacea, Gnaphalium supinum,Deschampsia cespitosa ssp. alpina, Huperzia selago, Diphasiastrum alpinum, Carex bigelowii, Galiumsaxatile and Sibbaldia procumbens. There are also tufts of mosses such as Andreaea alpina, Racomitriumheterostichum, R. lanuginosum, R. ericoides and Polytrichum alpinum.

These liverwort crusts are among the more distinctive and easily-recognisable types of snow-bed vegetation,as they do not look like any other plant community. The dark patches stand out well among the green andgolden tones of the surrounding heaths and grasslands. U12b is not as common in the study area as U11and the related types. It is extensive only on the steep back walls of Coire Gabhail, where it forms mosaics

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with other types of snow-beds, moss heaths and montane grasslands. U12b snow-beds are the mostwidespread of this type of vegetation in Scotland. Like U11 they are most common on the high cold hills in the central and eastern Highlands, but they also occur well to the north and west and in the Hebrides.There are fragments of this or very closely-related vegetation in the Southern Uplands.

U13a Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemillaalpina sub-community

U13a is one of the most extensive types of vegetation in the high corries of the study area. The dark greentussocky sward of Deschampsia cespitosa, with its distinctive tough, grooved, sharp-edged stiff leaves,clothes the corrie floors and steep walls, especially under cliffs or boulder-fields where the soils are kept moistby cold and acid water. The D. cespitosa is interleaved with Anthoxanthum odoratum, Festuca vivipara,Agrostis canina, Carex bigelowii and Luzula multiflora. The turf is speckled with small plants such as Violapalustris, Rumex acetosa, Alchemilla alpina, Galium saxatile, Potentilla erecta, Cerastium fontanum, Euphrasiaofficinalis agg., Oxalis acetosella and Blechnum spicant. All are interwoven with deep spongy carpets ofbryophytes such as Rhytidiadelphus loreus, R. squarrosus, Pleurozium schreberi, Hylocomium splendens,Hypnum jutlandicum, Sphagnum denticulatum and Plagiothecium undulatum. In the upper part of Coire namBeitheach there are patches of U13a with much Salix herbacea: a rather unusual plant in this type of vegetation.

U13a is common in Coire nam Beitheach (see Photographs 15 and 21), Coire nan Lochan and upper Coire Gabhail (Photograph 22), and occurs on high slopes and cliffs throughout the study area. This typeof upland grassland is generally near-natural, covering ground above the altitudinal limit of woodland andeven of dwarf-shrub heaths on cool and shaded slopes. U13a is covered with snow through most of thewinter and irrigated with icy water from the melting snow-patches in spring and early summer. U13a iscommon throughout the Highlands. It is especially extensive on the high hills which form the west Highlandwatershed. There are fragmentary outliers in southern Scotland, the Lake District and north Wales.

U13ahr Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemillaalpina sub-community, herb-rich form

This type of herb-rich Deschampsia cespitosa grassland is not described in the NVC. McVean & Ratcliffe(1962) found herb-rich D. cespitosa grasslands quite widely in Scotland, between the Breadalbanes andthe north-west Highlands. Most of their data for this vegetation seem to have been incorporated into theAgrostis-Rhytidiadelphus sub-community of the Luzula-Geum tall-herb community U17c. This seems fairenough for the swards where there are dwarfed basal rosettes of the characteristic tall herbs of U17, andwhere it is clear that the vegetation would revert to a tall-herb community in the absence of grazing. Thereare good examples of this on Ben Lui (Averis & Averis 1999a). But the swards on Bidean are different. Theyare flushed Deschampsia grasslands, enriched with a few small base-tolerant species. They do not look likegrazed swards of tall-herb vegetation, and indeed the set of species which grow in them are quite differentfrom those which grow in the ungrazed tall-herb vegetation on nearby cliffs. They are the equivalent of theherb-rich Nardus grassland U5c and the herb-rich Juncus squarrosus rush heath U6R, with a sward ofDeschampsia cespitosa. As always, the swards are not pure, and there is much Anthoxanthum odoratum,Festuca vivipara, Agrostis capillaris and A. canina as well as Carex bigelowii. Thymus polytrichus iscommon in some patches, growing with an array of species including Ranunculus acris, Selaginellaselaginoides, Saxifraga hypnoides, Thalictrum alpinum, Viola palustris, Alchemilla alpina, A. glabra,Persicaria vivipara, Cerastium fontanum, Campanula rotundifolia, Galium saxatile and Potentilla erecta.

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There are wet mats and cushions of bryophytes including Rhytidiadelphus loreus, R. squarrosus, Hylocomiumsplendens, Calliergonella cuspidata, Rhizomnium punctatum, Plagiomnium undulatum, Hypnum lacunosum,Pleurozia purpurea and even the odd patch of Sphagnum capillifolium.

We found this type of vegetation on wet ledges and in hollows in the upper parts of Coire nam Beitheach.

U13b Deschampsia cespitosa-Galium saxatile grassland, Rhytidiadelphus loreus sub-community

This is the typical vegetation of the steep upper walls of corries in the west Highlands where snow builds upinto a cornice over the winter, and where great masses of snow can linger far into spring. It consists of red-golden patches of Rhytidiadelphus loreus: the dense carpets of its curled and springy shoots are sparselystudded with Deschampsia cespitosa ssp. alpina, Nardus stricta, Festuca vivipara, Anthoxanthum odoratum,Galium saxatile, Vaccinium myrrtillus, V. vitis-idaea, Saxifraga stellaris, Alchemilla alpina and Rumexacetosa. Hidden among the carpets of Rhytidiadelphus loreus are tufts, clumps and mats of many otherbryophytes such as Hylocomium splendens, Racomitrium lanuginosum, Hypnum jutlandicum, Polytrichumalpinum, Kiaeria starkei, Pleurozium schreberi, Anastrepta orcadensis, Barbilophozia floerkei, Scapaniagracilis, Ptilidium ciliare and Nardia compressa.

U13b is very common on the high slopes of Coire nam Beitheach, Coire nan Lochan and Coire Gabhail.It is one of the most extensive types of snow-bed vegetation in the hills of the north and west Highlands,where it can cover vast areas of ground around the upper rims of the corries.

U14 Alchemilla alpina-Sibbaldia procumbens dwarf-herb community

This, like U12a, is one of the snow-beds with plenty of vascular plants as well as mosses and liverworts. It has an attractive delicate green flowery sward of Alchemilla alpina and Sibbaldia procumbens, growingwith an array of small plants such as Carex bigelowii, Deschampsia cespitosa ssp. alpina, Huperzia selago,Diphasiastrum alpinum, Saxifraga stellaris, Gnaphalium supinum, Campanula rotundifolia and Veronicaserpyllifolia ssp. humifusa. Under these is a carpet of bryophytes. One of the most common is Racomitriumfasciculare, which can form a distinctive golden-brown underlay to some patches of U14. There can alsobe a more mixed and variegated patchwork of species such as Racomitrium ericoides, R. lanuginosum,Polytrichum alpinum, Andreaea alpina, Conostomum tetragonum, Oligotrichum hercynicum, Kiaeria falcata,K. starkei, Dicranum fuscescens, Hypnum jutlandicum, Nardia compressa, Barbilophozia floerkei, Marsupellaemarginata and M. sphacelata.

U14 is especially characteristic of the unstable ice-plucked ground around the upper rim of a corrie, but alsooccurs in earthy gullies, on steep crumbling slopes and on spreads of wet gravel lower down in the corries.We found it in all the high corries in the study area. The Alchemilla-Potentilla community mentioned byMcVean & Ratcliffe (1962), in which Sibbaldia procumbens is replaced by Potentilla erecta, also occurs inthese high corries. U14 is widespread throughout the Highlands, on the higher hills where there is much late-lying snow. The Alchemilla-Potentilla form is common in the west Highlands, and there is related vegetationin the Lake District.

U15 Saxifraga aizoides-Alchemilla glabra banks

This herb-rich community of flushed rocks and banks is intermediate in flora and in habitat between theCarex-Saxifraga mires M11 and the more terrestrial tall-herb community U17. Common on the limestone

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rocks of Meall Mór, U15 is made up of thick hanging masses of Saxifraga aizoides and S. oppositifolia,their succulent leaves streaming with water. The branching sprays of golden-yellow flowers of the S. aizoidesmake these mats conspicuous from a distance in summer. Growing among the saxifrages in rock crevicesand in the saturated carpets of bryophytes is an assemblage of small base-tolerant plants such as Selaginellaselaginoides, Alchemilla alpina, A. glabra, Carex pulicaris, C. capillaris, C. viridula ssp. oedocarpa, C. flacca, Thymus polytrichus, Silene acaulis, Linum catharticum, Sagina procumbens, Saussurea alpina,Thalictrum alpinum, Plantago maritima, Pinguicula vulgaris, Asplenium viride, Galium boreale and Primula vulgaris. Bryophytes are plentiful among the vascular plants and hanging down over the rock faces.Species include Ctenidium molluscum, Breutelia chrysocoma, Anoectangium aestivum, Blindia acuta,Calliergonella cuspidata, Hymenostylium insigne, Bryum pseudotriquetrum, Orthothecium rufescens,Hylocomium splendens, Fissidens adianthoides, Scorpidium scorpioides, Scapania undulata, Frullaniatamarisci, Plagiochila porelloides, Aneura pinguis and Preissia quadrata.

U15 occurs on base-rich rocks and banks throughout the Highlands.

U16a Luzula sylvatica-Vaccinium myr til lus tall-herb community, Dryopteris dilatata-Dicranum

majus sub-community

There are tiny patches of this type of vegtation on the ledges of the larger cliffs. Entirely out of reach, theyconsist of lush swards of ferns such as Dryopteris dilatata, D. filix-mas and Oreopteris limbosperma growingwith much Luzula sylvatica, Vaccinium myrtillus and Deschampsia cespitosa. The bryophytes were notidentifiable through binoculars, but in this kind of vegetation generally include clumps of Dicranum majusand D. scoparium, and mats of large common species such as Pleurozium schreberi, Hypnum jutlandicum,Hylocomium splendens, Rhytidiadelphus loreus and Racomitrium lanuginosum. There can be an array of smallervascular species such as Oxalis acetosella, Potentilla erecta, Phegopteris connectilis and Blechnum spicant.

U16a is the characteristic vegetation of acid cliff ledges in the British uplands.

U16b Luzula sylvatica-Vaccinium myr ti l lus tall-herb community, Anthoxanthum odoratum-

Festuca ovina sub-community

This is another tall community of ungrazed cliffs. It is like U16a (see above) without the ferns. There are goodpatches on the cliffs on the western side of Coire nam Beitheach, with a thick, lush green sward of Luzulasylvatica and Vaccinium myrtillus overtopped by the flowering heads of Deschampsia cespitosa, D. flexuosa,Agrostis capillaris, Festuca ovina and Carex binervis. There are tufts of Blechnum spicant and Huperziaselago, and bushy masses of Potentilla erecta and Galium saxatile. Under these are dense layers ofbryophytes such as Rhytidiadelphus loreus, R. squarrosus, Hypnum jutlandicum, Pleurozium schreberi,Hylocomium splendens, Polytrichum commune, Dicranum majus and D. scoparium.

There are scattered patches of U16b on the cliffs in the study area. U16b is widespread on acid cliffs andsteep banks throughout the British uplands.

U16c Luzula sylvatica-Vaccinium myr ti l lus tall-herb community, Species-poor sub-community

This is the form of Luzula sylvatica vegetation which covers open accessible slopes and forms vast areas oftall, russet-green, rustling swards on the steep north-facing slopes of the study area. Though the vegetation

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looks lush it is, in fact, excessively species-poor. Luzula sylvatica is usually completely dominant. Very littlecan push up between its tight rosettes except for sparsely scattered clumps of Deschampsia flexuosa, Agrostis capillaris or Festuca vivipara, sprigs of Vaccinium myrtillus, or trailing stems of Galium saxatile orPotentilla erecta. There is hardly any room at all for bryophytes. All that grow here are sparse shoots ofRhytidiadelphus squarrosus, Pleurozium schreberi, Dicranum scoparium or Polytrichum commune.

U16c is common on the northern slopes of the main glen and on the sides of Aonach Dubh, Gearr Aonachand Beinn Fhada. This type of vegetation is widespread in the British uplands from Wales northwards, butseems to form the largest patches in the western Highlands.

U17a Luzu la sy l va t i ca -Geum r i va le ta l l -he rb commun i ty, A l chemi l la g labra -Br yumpseudotr iquetrum sub-community

Of the four types of Luzula-Geum vegetation, this is the one of the wettest soils and with the highest numberof small spring and flush species. Most of the tall-herb vegetation on the base-rich andesite ledges at the topof Coire nam Beitheach is of this type (see Photgraph 17). The patches of short but lush vegetation clotherock ledges and steep gravelly slopes. The swards are made up of species such as Luzula sylvatica, L. spicata, Sedum rosea, Oxyria digyna, Ranunculus acris, Trollius europaeus, Cerastium arcticum,Selaginella selaginoides, Carex flacca, C. pulicaris, C. viridula ssp. oedocarpa, Saxifraga hypnoides, S. aizoides, Alchemilla alpina, A. glabra, Sibbaldia procumbens, Juncus triglumis, Thalictrum alpinum,Cochlearia pyrenaica ssp. alpina, Poa alpina, Trollius europaeus, Arabis petraea, Asplenium viride,Polystichum lonchitis and Silene acaulis. The lowland Silene dioica is an unexpected member of some ofthese swards. The rare species of the study area Saxifraga nivalis, S. rivularis, S. cernua, Poa glauca, Juncus biglumis, Cystopteris montana, Draba norvegica and Cerastium alpinum also occur in this type ofvegetation, but they are only locally plentiful. There are colourful clumps and patches of bryophytes on therocks, made up of species such as Ctenidium molluscum, Tortella tortuosa, Philonotis fontana, P. calcarea,P. seriata, Hylocomium splendens, Orthothecium rufescens, Amphidium mougeotii, Bryum pseudotriquetrum,Campylium stellatum, Blindia acuta, Herbertus stramineus and Scapania undulata.

U17a occurs at lower altitudes too, but these patches lack the more montane species. U17a is widespreadthrough the Scottish Highlands and also occurs in southern Scotland, the Lake District and Wales. In its floraand habitat it is quite similar to the Saxifraga-Alchemilla banks U15.

U17b Luzula sylvatica-Geum rivale tall-herb community, Geranium sylvaticum sub-community

These are the taller, more lush swards of Luzula-Geum vegetation. They can resemble the ground vegetationof ungrazed Fraxinus-Sorbus-Mercurialis woodland W9, and on Meall Mór the two types occur closetogether. There are patches of U17b on the larger cliff ledges right up to near the summit ridge, and thistype of vegetation is also common around waterfalls and in ravines where the rocks are base-rich. Theluxuriant swards are like miniature meadows of grasses and tall herbs. They are colourful attractive mixturesof Luzula sylvatica, Deschampsia cespitosa, Geum rivale, Alchemilla glabra, A. alpina, Angelica sylvestris,Filipendula ulmaria, Valeriana officinalis, Ranunculus acris, Oxyria digyna, Geranium sylvaticum, G. robertianum, Rubus saxatilis, Trollius europaeus, Cirsium palustre, C. heterophyllum, Solidago virgaurea,Hieracium spp., Saussurea alpina, Galium boreale, Rhinanthus minor, Silene dioica, Crepis paludosaand Parnassia palustris. There is also a great host of smaller plants such as Selaginella selaginoides,Thalictrum alpinum, Saxifraga aizoides, S. oppositifolia, Carex flacca, C. pulicaris, C. viridula ssp. oedocarpa,

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Pinguicula vulgaris, Persicaria vivipara, Linum catharticum, Lotus corniculatus, Cystopteris fragilis, Aspleniumtrichomanes and A. viride. These sensationally rich assemblages clothe the wet ledges and spill over therock faces. Fine streams of water pour down through the herbage and sustain an array of bryophytes suchas Ctenidium molluscum, Tortella tortuosa, Neckera crispa, Orthothecium rufescens, Ditrichum flexicaule,Plagiomnium undulatum, Breutelia chrysocoma, Anoectangium aestivum, Mnium stellare, Plagiobryum zieri,Hylocomium splendens, Hymenostylium insigne, Rhytidadelphus triquetrus, Thuidium tamariscinum, Fissidensadianthoides, Herbertus stramineus, Scapania gracilis, S. aspera, Preissia quadrata and Plagiochilaporelloides.

U17b is the sub-community where the tall herbs are most common and luxuriant. The many fine patches ofthe community on the Breadalbane hills are mostly of this type. It occurs throughout the Highlands, in southernScotland and in the Lake District and north Wales.

U17d Luzula sylvatica-Geum rivale tall-herb community, Primula vulgaris-Hyperichum pulchrumsub-community

Some of the patches of tall-herb vegetation on the ledges of the wooded cliffs and in ravines, at low tomoderate altitudes, are of this type. Although the tall herbs are common this is a heathy assemblage of plants, with big ungrazed bushes of Calluna vulgaris, Vaccinium myrtillus and Erica cinerea growing inlush swards of Luzula sylvatica, Deschampsia cespitosa, Anthoxanthum odoratum, Agrostis capillaris andFestuca vivipara. Many of the plants which grow here are woodland species such as Primula vulgaris,Oxalis acetosella, Anemone nemorosa, Hyacinthoides non-scripta, Dryopteris dilatata, Teucrium scorodoniaand Geranium robertianum. They are interleaved with species such as Filipendula ulmaria, Alchemillaglabra, Heracleum sphondylium, Valeriana officinalis, Rubus saxatilis, Angelica sylvestris and Sedum rosea.As in the other types of U17 there are spreading mats and patches of bryophytes among the vascular plants,including Breutelia chrysocoma, Neckera crispa, Rhytidiadelphus triquetrus, Racomitrium lanuginosum,Tortella tortuosa, Ctenidium molluscum, Thuidium tamariscinum, Hylocomium splendens, Eurhynchiumpraelongum, Isothecium myosuroides var. brachythecoides, Frullania tamarisci and Aneura pinguis.

U17d is the most common type of Luzula-Geum tall-herb vegetation at low to moderate altitudes and in thefar west Highlands and in the Hebrides. It also occurs in the Lake District and Wales.

U18 Cr yptogramma crispa-Athyrium distentifolium snow-bed

The brilliant green patches of ferns in the high corries mark out places where snow lies late and deep indamp hollows among boulders and on scree. Cryptogramma crispa is by far the most common species. Itsnew fronds unroll in a lacy froth of green over the dead red-brown mats of last year’s leaves. Only in a fewshaded places among boulders are there also clumps of Athyrium distentifolium, but other ferns such asPhegopteris connectilis, Blechnum spicant, Dryopteris dilatata and the scarce D. oreades are reasonablycommon. They grow in a thin sward of Deschampsia cespitosa and Festuca vivipara with loose tufts ofGalium saxatile and Potentilla erecta. A dark layer of bryophytes clothes the ground among the screes orboulders. This is made up of snow-tolerant species such as Anthelia juratzkana, Pleurocladula albescens,Kiaeria blyttii, Polytrichum alpinum, Racomitrium fasciculare, Barbilophozia floerkei, Diplophyllum albicans,Dicranum fuscescens, Conostomum tetragonum, Oligotrichum hercynicum and a number of oceanicliverworts including Anastrophyllum donnianum, Scapania nimbosa, S. ornithopodioides, Bazzania pearsonii,Anastrepta orcadensis and Mylia taylorii. These liverworts, like the ferns, are able to grow at high altitudes

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because the covering of snow in winter ameliorates the temperature and protects them from frost. Indeed,Anastrophyllum donnianum is extraordinarily common in U18 among the boulders in the upper part of Coire nam Beitheach, growing in pure, dark red-brown swards several metres across. It is anattractive plant with two rows of long narrow leaves which make the shoots look like miniature combs(Photograph 40). The mats of bryophytes are dotted with small plants such as Huperzia selago, Saxifragastellaris, Rumex acetosa and Sibbaldia procumbens.

There are large areas of U18 in the upper parts of Coire nam Beitheach, Coire nan Lochan and CoireGabhail. This type of vegetation occurs throughout the Highlands, extending eastwards to the Cairngorms,but is most widespread and extensive on the hills of the west Highland watershed from Ben More Assyntdown to Ben Lui.

U19 Oreopteris l imbosperma-Blechnum spicant community

All over the north-facing slopes of Meall Mór and Bidean nam Bian, at low to moderate altitudes, there are huge patches of this type of vegetation. The smooth steep slopes are clothed with the light-green, citrus-scented fronds of Oreopteris limbosperma. U19 is common too on stream banks and on the steep sidesof rocky knolls where the soils are deep and quite fertile but well-drained. The canopy of Orepteris concealsan array of other species including ferns such as Blechnum spicant and Phegopteris connectilis, woodlandherbs such as Hyacinthoides non-scripta, Anemone nemorosa, Oxalis acetosella and Lysimachia nemorumand grassland species such as Festuca ovina, F. rubra, Agrostis capillaris, A. canina, Anthoxanthumodoratum, Carex binervis, Galium saxatile, Potentilla erecta, Viola riviniana and V. palustris. Under these,on ground which can be stony, there are swathes of bryophytes such as Pleurozium schreberi, Hypnumjutlandicum, Hylocomium splendens, Thuidium tamariscinum, Scleropodium purum, Rhytidiadelphussquarrosus, R. loreus, Polytrichum commune, and Dicranum scoparium. Locally the soils are more moist andspecies such as Molinia caerulea, Succisa pratensis, Carex panicea and Sphagnum capillifolium growamong the ferns. In some patches of U19, generally those below cliffs with either Quercus-Betula-Dicranumwoodland W17 or with Calluna-Vaccinium-Sphagnum damp heath H21, there are clumps and cushions ofoceanic western liverworts including Herbertus aduncus, Pleurozia purpurea, Mylia taylorii, Scapaniagracilis, Plagiochila spinulosa, Bazzania tricrenata, Anastrepta orcadensis and Lepidozia pearsonii. This isan unusual and interesting element in this type of vegetation.

U19 is common in the study area. Though it occurs throughout the British uplands, it is most widespread andextensive in the hills of the south-west Highlands.

U20a Pteridium aquilinum-Galium saxatile community, Anthoxanthum odoratum sub-community

Of all the types of bracken-dominated vegetation in the study area, this is the most common and the mostwidespread. It forms distinctive dark green sharp-edged patches at low to moderate altitudes on the slopesall around the hills. It is especially common on the lower slopes of the Aonach Eagach (visible as orange-brown patches in Photographs 32–33). Under the dense spreading canopy of the fern there is vegetationvery much like a Festuca-Agrostis-Galium grassland, with a thick green turf of Festuca ovina, F. rubra, Agrostis capillaris, A. canina, Danthonia decumbens, Deschampsia flexuosa and Anthoxanthum odoratum.It is speckled with small plants such as Potentilla erecta, Galium saxatile, Viola riviniana, Carex binervis,Euphrasia officinalis agg. and Rumex acetosa, and with wispy patches of bryophytes such as Rhytidadelphussquarrosus, Thuidium tamariscinum, Scleropodium purum, Eurhynchium praelongum and Hypnum jutlandicum.

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On the lower, north-facing slopes in the main glen there is much Hyacinthoides non-scripta among thebracken, and the young fronds unroll through a cloud of sweet-scented blue-purple flowers. As in theOreopteris community U19, there are patches of bracken on north-facng slopes which shelter mats andcushions of oceanic liverworts such as Herbertus aduncus, Pleurozia purpurea, Mylia taylorii, Scapaniagracilis, Plagiochila spinulosa, Bazzania tricrenata, Anastrepta orcadensis and Lepidozia pearsonii. It iseven more unusual to find these species in U20 than it is to find them in U19.

U20a is common throughout the British uplands, especially in England, Wales, southern Scotland, the south-west Highlands and the Inner Hebrides. It is generally believed to have spread as a result of sheep grazingrather than cattle. Although cattle do not browse it much, they trample it hard, and this breaks up therhizomes and restricts its spread. Also at one time it was a common practice to cut bracken and dry it forwinter bedding. With the falling populations of both people and cattle and with the shortage of cheaplabour, this is no longer done. As a result the fern has spread on deep and fertile soils.

U20c Pteridium aquil inum-Galium saxati le community, Species-poor sub-community

There are a few patches of this type of vegetation in the floor of the main glen, with a dense andimpenetrable canopy of bracken standing over a deep underlayer of its own litter. Few species are able to grow here, and there is usually little more than a thin sprinkling of the characteristic species of U20:vascular plants such as Agrostis capillaris, Holcus lanatus and Potentilla erecta and bryophytes such asRhytidiadelphus squarrosus, Hypnum jutlandicum, Scleropodium purum, Hylocomium splendens andThuidium tamariscinum. Such intractable vegetation as this has little to offer either for agriculture or for natureconservation. Even when it is cleared by cutting or spraying, the resulting bare soil is easily eroded intounsightly and long-lasting scars on the landscape.

U20c occurs throughout the uplands but is rare in the northern Highlands.

U21 Cr yptogramma crispa-Deschampsia flexuosa community

This is essentially the more lowland or sub-montane counterpart of the Cryptogramma-Athyrium snow-bedU18. Nevertheless, since it is a community of dry acid screes, it can and does occur well up in the corrieswhere its range overlaps with that of U18. The two communities can look very similar, with their greenmounds of Cryptogramma crispa stippling the stony slopes. Other than that their floras are quite different.Instead of the snow-tolerant vascular plants and bryophytes of U18, U21 includes species such asDeschampsia flexuosa, Potentilla erecta, Galium saxatile, Agrostis capillaris, Festuca ovina, Rumex acetosa,Vaccinium myrtillus and Digitalis purpurea. The mats of bryophytes among the stones consist of species suchas Racomitrium lanuginosum, R. ericoides, R. fasciculare, R. sudeticum, Pleurozium schreberi, Campylopusflexuosus, Dicranum scoparium and Hypnum jutlandicum. There can be grey patches of lichens includingCladonia portentosa, C. subcervicornis, C. furcata and Peltigera canina. In some of the patches in the highcorries there are montane plants such as Carex bigelowii, Diphasiastrum alpinum and Kiaeria starkei, butgenerally it is easy to tell the U18 snow-beds from these U21 communities of loose scree.

U21 is common in north Wales, northern England and the Southern Uplands. It is also plentiful in the south-west Highlands and in the Inner Hebrides, but becomes scarce further north and east, as Cryptogrammacrispa is sensitive to frost and survives in cold districts only where there is a protective covering of snow inwinter.

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S9a Carex rostrata swamp, Carex rostrata sub-community

S9a consists of grey-green thin pure swards of Carex rostrata growing in shallow water up to about half ametre deep. It covers a large area in the southern part of Loch Achtriochtan, and part-fills Lochan nan Folaon the edge of Rannoch Moor in the north-east of the study area. There is also a patch of it in one of thelochans in the upper part of Coire nan Lochan.

S9a is common in pools and lochs in the British uplands. It can be found in slow-flowing streams and rivers,especially in the reaches of dead water where they pass through blanket bogs. It also occurs in the lowlands.

S9b Carex rostrata swamp, Menyanthes trifoliata-Potamogeton polygonifolius sub-community

Some of the Carex rostrata swamps consist of slightly more continuous vegetation. In shallow water with abottom of peaty mud, they are made up of swards of Carex rostrata growing through mats of emergentPotamogeton polygonifolius, Menyanthes trifoliata, Juncus bulbosus and Utricularia minor.

There are small patches of this vegetation around the edges of open water at Lochan na Fola in the north-east of the site. The surrounding vegetation here is mainly a mosaic of different types of blanket bog. S9b is widespread in Britain, in bog pools and shallow peaty channels through bogs and wet heaths.

S10a Equisetum fluviati le swamp, Equisetum fluviati le sub-community

The sward of S10a is even thinner, more open and more sparse than that of S9a. It is hard to see it as acommunity at all. It consists of the supple jointed stems of Equisetum fluviatile growing in water up to abouta metre deep.

We found it around the edge of Loch Achtriochtan. Like S9a, S10a is a swamp of pools, lochans and slow-moving streams and rivers, mostly in the uplands of Britain.

OW + Menyanthes = Menyanthes trifoliata vegetation in open water

This is a very sparse but rather conspicuous scatter of numerous shoots of Menyanthes trifoliata growing inopen water, usually accompanied by Equisetum fluviatile. This type of vegetation grows in lochans in theinteresting bog complex at the eastern end of the site, east of Buachaille Etive Mór (Photograph 45). Similarvegetation occurs in bog pools elsewhere in the Highlands, especially in the far north. This vegetation doesnot fit into any NVC type. It is shown in our vegetation maps using the code ‘OW + Menyanthes’.

Juncus ef fusus

Vegetation labelled ‘Je’ for Juncus effusus on the maps consists of dense patches of this species with nounderstorey that could be used to assign them to any of the rush mires or wet pastures. Other than the Juncusitself it is rare to find much in these patches apart from a few shoots of grasses such as Holcus lanatus,Festuca rubra, F. ovina, Anthoxanthum odoratum and Agrostis capillaris, small herbs such as Rumex acetosa,Ranunculus acris and Cerastium fontanum, and the odd shoot of bryophytes such as Rhytidiadelphussquarrosus.

There are small patches of this vegetation among grassland and bracken in two places in the bottom of themain glen: near Achnacon and near Achtriochan.

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Patches of this kind of vegetation are presumably the result of clonal growth over many years, and arecommon in the hills of Wales, northern England and southern Scotland where the vegetation has been muchchanged by heavy grazing over many years. They occur more sporadically in the Highlands, where theland-use is less intensive.

Racomitrium ericoides shingle vegetation

Racomitrium ericoides is a common and successful coloniser of bare scree and gravel and the sides of tracks.There are great spreads of it on the river gravels and shingles beside the River Coe in the floor of the mainglen. Presumably much of this is inundated in winter when the river is in spate, and the constant shifting,sorting and redistribution of the gravel must prevent most plants from becoming established and allow themoss to persist.

Fell-f ield

Fell-fields are the stony, level patches of gravel on the ridges and high summits. The result of ablation anderosion by wind, from a distance they look completely bare and indeed they are nearly so. However, thereis a characteristic assemblage of small plants which are able to colonise and persist on these surfaces in theface of continued churning by frost and trampling by people and grazing animals. It includes Saxifragastellaris, Campanula rotundifolia, Solidago virgaurea, Vaccinium myrtillus, V. vitis-idaea, Deschampsia flexuosa,Festuca vivipara, Carex panicea, Salix herbacea, Galium saxatile and bryophytes such as Racomitiumlanuginosum, R. ericoides, R. sudeticum and R. heterostichum.

Fell-fields are not described in the NVC, though they occur very widely in the uplands. Many people arefamiliar with the pink granite gravel of summits and ridges in the Cairngorms, and with the black basaltgravels of Mull and northern Skye. Both are fell-fields, and both granite and basalt seem particularly proneto developing fell-fields, partly because they form flat-topped hills and plateaux and partly because the rockweathers into small and uniform fragments. Fell-fields also occur on other types of rock and on hills with moresteep terrain, and can be found in north Wales, northern England and throughout the uplands of Scotland.

Garden

Though hardly gardens in either the decorative or horticultural senses, the enclosed grounds around the farmsteadings at Achnambeithach and Achtriochtan have been given this label to distinguish them from thesurrounding, less heavily-modified land.

Acid Rock

Almost all of the rock in the study area is hard and acidic. It is also one of the most extensive habitats. Evento the most casual observer, the vast precipices of Aonach Dubh and Gearr Aonach and the Aonach Eagachgive this glen much of its scenic grandeur, and the cones of Buachaille Etive Beag and Buachaille Etive Mórare almost pure rock. Needless to say, though, little rock is absolutely bare. Small plants cling to the cracksand crevices, commonly in attenuated forms of the plant communities of the surrounding more vegetatedground. Even the sheer faces have a sparse growth of plants such as the mosses Andreaea rupestris,Racomitrium sudeticum, R. lanuginosum, R. heterostichum, R. fasciculare, Hedwigia stellata, Hypnumcupressiforme, Campylopus atrovirens and Dicranum scoparium, the liverworts Diplophyllum albicans,

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Marsupella emarginata, Gymnomitrion obtusum, G. crenulatum, Barbilophozia floerkei and Scapaniagracilis, and the lichens Cladonia subcervicornis and Parmelia saxatilis.

Basic Rock

The largest expanse of basic rock in the study area is the outcrop of limestone which forms the eastern partof Meall Mór. Base-rich volcanic rocks outcrop in many parts of the study area, from the lower ground rightup to the col under the west top of Bidean at over 1100m. Outcrops of both types of rock have a generallysimilar flora: a scatter of many small plants such as the mosses Tortella tortuosa, Neckera crispa, Blindia acuta, Anoectangium aestivum, Ditrichum flexicaule, Schistidium apocarpum, Ctenidium molluscum,Hypnum lacunosum, Homalothecium sericeum, Dryptodon patens, Orthothecium rufescens, Grimmia funalisand Plagiobryum zieri and the liverworts Frullania tamarisci, Plagiochila porelloides, Scapania aspera andCololejeunea calcarea. The oceanic liverwort Herbertus stramineus occurs on some volcanic outcrops. The rare oceanic moss Hymenostylium insigne grows on some damp limestone outcrops on Meall Mór. Small vascular species such as Thymus polytrichus, Asplenium trichomanes and Hypericum pulchrum growhere and there on limestone or igneous outcrops, where their roots can get a foothold on a small ledge orin a soily crevice.

Even on the predominantly acid cliffs, there are innumerable more basic dykes and igneous intrusionsbringing lines of herb-rich vegetation down through expanses of species-poor acid heaths and grasslands.This is well seen on the Aonach Eagach and on the eastern side of the Lairig Eilde. The most notable of thebase-rich outcrops, however, are those in upper Coire nam Beithach, where the rare Saxifraga cernua, S. rivularis, S. nivalis, Draba norvegica, Juncus biglumis, Cerastium alpinum, Cystopteris montana and Poa glauca grow in crevices, on tiny ledges and beneath overhangs together with an array of calcicolebryophytes including Orthothecium rufescens and Herbertus stramineus.

Acid Scree

Most of the great scree-fans and scree-choked gullies in the study area are of hard acidic rock which hasweathered into medium-sized angular fragments. Many are recent or subject to continuous new rockfall andmovement and are almost unvegetated. Others are coated with a greasy layer of mosses such asRacomitrium lanuginosum, R. ericoides, R. heterostichum, R. sudeticum, Andreaea rupestris, Campylopusatrovirens, Polytrichum formosum and a great range of crustose lichens which were investigated morethoroughly by Fryday (1993). Vascular plants are rare except at the lower edges of the scree-runs, and evenhere there is just a sparse assembly of plants such as Digitalis purpurea, Succisa pratensis, Rumex acetosa,Solidago virgaurea, Viola riviniana, Agrostis canina, A. capillaris, Festuca vivipara, Teucrium scorodonia,Blechnum spicant and Vaccinium myrtillus.

Basic Scree

Very little of the scree is basic, even where it has tumbled down through areas of base-rich soils on the lowerslopes. There are, however, a few scree-fans in the Lairig Eilde and the Lairig Gartain where the rocks areovergrown with species such as Thymus polytrichus, Lotus corniculatus, Ajuga reptans, Prunella vulgaris,Cirsium heterophyllum and even the odd plant of Oxyria digyna. These are obviously fragments of morebase-rich rock.

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Boulder-field

In the upper corries and at the north-western entrance to the Lost Valley, there are fields of large boulders.These are the result of cliffs collapsing in late glacial or early post-glacial times. The boulder-field in the LostValley is partly wooded and there is an impressive array of bryophytes and ferns among the boulders andon the tops and sides of the larger blocks: see the description of W17a woodland. The boulder-fields in thehigh corries are more sparsely vegetated. Many of the patches of Cryptogramma-Athyrium snow-bed U18and of liverwort-rich Vaccinium-Racomitrium heath H20c lie in the shelter of these boulders. Generally,though, the vegetation consists mainly of patches of brophytes which are able to tolerate prolonged snow-lie and high humidity. The most notable of these are the oceanic liverworts Anastrophyllum donnianum,Scapania nimbosa, S. ornithopodioides and Bazzania pearsonii, and montane rarities such as Pleurocladulaalbescens, Diplophyllum taxifolium, Brachythecium glaciale, Polytrichum sexangulare, Moerkia blyttii, Lescuraeapatens, Mnium thomsonii, Marsupella boeckii and Isopterigium muellerianum. The most notable vascularplants of these high boulder-fields are Athyrium distentifolium, Dryopteris oreades, Saxifraga rivularis, S. nivalis and Cerastium arcticum.

Shingle

This label was used for bare spreads of shingle beside the rivers. These have evidently been brought downby the most recent spates, and at the time of survey had only a sparse array of plants such as Potentillaerecta, Digitalis purpurea and Racomitrium species.

Open Water

Apart from small pools in the bogs and the lochans in the high corries, the only substantial areas of openwater in the study area are Loch Achtriochtan and Lochan nam Fola. Both are shallow with much emergentvegetation.

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Table 1 List of vegetation types recorded in this sur vey at Glen Coe in 2002


W4 Betula pubescens-Molinia caerulea woodland

W7b Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland, Carex remota-Cirsium palustresub-community

W7c Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland, Deschampsia cespitosa sub-community

W9b Fraxinus excelsior-Sorbus aucuparia-Mercurialis perennis woodland, Crepis paludosa sub-community

W11a Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Dryopteris dilatata sub-community

W11b Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Blechnum spicant sub-community


W17b Quercus petraea-Betula pubescens-Dicranum majus woodland, Typical sub-community

W17c Quercus petraea-Betula pubescens-Dicranum majus woodland, Anthoxanthum odoratum-Agrostis capillarissub-community

W18d Pinus sylvestris-Hylocomium splendens woodland, Sphagnum capillifolium/quinquefarium sub-community


W24 Rubus fruticosus agg.-Holcus lanatus underscrub

W25 Pteridium aquilinum-Rubus fruticosus agg. underscrub

W25D Pteridium aquilinum-Rubus fruticosus agg underscrub, damp sub-community *

H10a Calluna vulgaris-Erica cinerea heath, Typical sub-community

H10b Calluna vulgaris-Erica cinerea heath, Racomitrium lanuginosum sub-community

H10c Calluna vulgaris-Erica cinerea heath, Festuca ovina-Anthoxanthum odoratum sub-community

H10d Calluna vulgaris-Erica cinerea heath, Thymus polytrichus-Carex pulicaris sub-community

H12b Calluna vulgaris-Vaccinium myrtillus heath, Vaccinium vitis-idaea-Cladonia impexa sub-community

H12c Calluna vulgaris-Vaccinium myrtillus heath, Galium saxatile-Festuca ovina sub-community

H13a Calluna vulgaris-Cladonia arbuscula heath, Cladonia arbuscula-Cladonia rangiferina sub-community


H18a Vaccinium myrtillus-Deschampsia flexuosa heath, Hylocomium splendens-Rhytidiadelphus loreussub-community

H18b Vaccinium myrtillus-Deschampsia flexuosa heath, Alchemilla alpina-Carex pilulifera sub-community

H20a Vaccinium myrtillus-Racomitrium lanuginosum heath, Viola riviniana-Thymus polytrichus sub-community

H20b Vaccinium myrtillus-Racomitrium lanuginosum heath, Cetraria islandica sub-community

H20c Vaccinium myrtillus-Racomitrium lanuginosum heath, Bazzania tricrenata-Mylia taylori sub-community

H21a Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath, Calluna vulgaris-Pteridium aquilinumsub-community

H21b Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath, Mastigophora woodsii-Herbertus aduncus hutchinsiae sub-community

H22a Vaccinium myrtillus-Rubus chamaemorus heath, Polytrichum commune-Galium saxatile sub-community

HX Vaccinium myrtillus-Sphagnum capillifolium heath *

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Table 1 (continued)




M6a Carex echinata-Sphagnum fallax/denticulatum mire, Carex echinata sub-community

M6b Carex echinata-Sphagnum fallax/denticulatum mire, Carex nigra-Nardus stricta sub-community

M6c Carex echinata-Sphagnum fallax/denticulatum mire, Juncus effusus sub-community

M6d Carex echinata-Sphagnum fallax/denticulatum mire, Juncus acutiflorus sub-community

M7 Carex curta-Sphagnum russowii mire

M10a Carex dioica-Pinguicula vulgaris mire, Carex viridula ssp. oedocarpa-Juncus bulbosus/kochii sub-community

M11a Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Thalictrum alpinum-Juncus triglumis sub-community

M11b Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Palustriella commutata-Eleocharis quinqueflorasub-community

M12 Carex saxatilis mire

M15a Trichophorum cespitosum-Erica tetralix wet heath, Carex panicea sub-community

M15b Trichophorum cespitosum-Erica tetralix wet heath, Typical sub-community

M15c Trichophorum cespitosum-Erica tetralix wet heath, Cladonia sub-community

M15d Trichophorum cespitosum-Erica tetralix wet heath, Vaccinium myrtillus sub-community

M17a Trichophorum cespitosum-Eriophorum vaginatum blanket mire, Drosera rotundifolia-Sphagnum sub-community

M17b Trichophorum cespitosum-Eriophorum vaginatum blanket mire, Cladonia sub-community

M18a Erica tetralix-Sphagnum papillosum raised and blanket mire, Sphagnum magellanicum-Andromeda polifolia sub-community

M18b Erica tetralix-Sphagnum papillosum raised and blanket mire, Empetrum nigrum nigrum-Cladonia sub-community


M23a Juncus effusus/acutiflorus-Galium palustre rush-pasture, Juncus acutiflorus sub-community

M25a Molinia caerulea-Potentilla erecta mire, Erica tetralix sub-community

M25b Molinia caerulea-Potentilla erecta mire, Anthoxanthum odoratum sub-community

M25c Molinia caerulea-Potentilla erecta mire, Angelica sylvestris sub-community

M28a Iris pseudacorus-Filipendula ulmaria mire, Juncus spp sub-community


M32a Philonotis fontana-Saxifraga stellaris spring, Sphagnum denticulatum sub-community

M32b Philonotis fontana-Saxifraga stellaris spring, Montia fontana-Chrysosplenium oppositifolium sub-community


M37 Palustriella commutata-Festuca rubra spring

MX Small sedge mire *

MG9 Holcus lanatus-Deschampsia cespitosa grassland

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Table 1 (continued)

CG10a Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland, Trifolium repens-Luzula campestrissub-community

CG10b Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland, Carex pulicaris-Carex panicea sub-community

CG10c Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland, Saxifraga aizoides-Ditrichum flexicaulesub-community

CG10 Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland on river shingle *

CG11a Festuca ovina-Agrostis capillaris-Alchemilla alpina grassland, Typical sub-community


CG14 Dryas octopetala-Silene acaulis community

U4a Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Typical sub-community

U4b Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Holcus lanatus-Trifolium repens sub-community

U4d Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Luzula multiflora-Rhytidiadelphus loreussub-community

U4e Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Vaccinium myrtillus-Deschampsia flexuosasub-community

U4eA Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Vaccinium myrtillus-Deschampsia flexuosasub-community, Alchemilla alpina variant *

U4F Festuca ovina-Agrostis capillaris-Galium saxatile grassland, provisional Filipendula ulmaria sub-community*

U5a Nardus stricta-Galium saxatile grassland, Species-poor sub-community

U5b Nardus stricta-Galium saxatile grassland, Agrostis canina-Polytrichum commune sub-community


U5e Nardus stricta-Galium saxatile grassland, Racomitrium lanuginosum sub-community

U6a Juncus squarrosus-Festuca ovina grassland, Sphagnum sub-community

U6c Juncus squarrosus-Festuca ovina grassland, Vaccinium myrtillus sub-community

U6d Juncus squarrosus-Festuca ovina grassland, Agrostis capillaris-Luzula multiflora sub-community

U6Cv Juncus squarrosus-Festuca ovina grassland, provisional Calluna vulgaris sub-community *

U6R Juncus squarrosus-Festuca ovina grassland, provisional herb-rich sub-community *

U7a Nardus stricta-Carex bigelowii grass heath, Empetrum nigrum hermaphroditum-Cetraria islandicasub-community




U10a Carex bigelowii-Racomitrium lanuginosum moss heath, Galium saxatile sub-community

U10b Carex bigelowii-Racomitrium lanuginosum moss heath, Typical sub-community




U12b Salix herbacea-Racomitrium heterostichum snow-bed, Gymnomitrion concinnatum sub-community

U13a Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemilla alpina sub-community

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Table 1 (continued)

U13ahr Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemilla alpinasub-community, herb-rich form *




U16a Luzula sylvatica-Vaccinium myrtillus tall-herb community, Dryopteris dilatata-Dicranum majus sub-community

U16b Luzula sylvatica-Vaccinium myrtillus tall-herb community, Anthoxanthum odoratum-Festuca ovinasub-community

U16c Luzula sylvatica-Vaccinium myrtillus tall-herb community, Species-poor sub-community

U17a Luzula sylvatica-Geum rivale tall-herb community, Alchemilla glabra-Bryum pseudotriquetrum sub-community


U17d Luzula sylvatica-Geum rivale tall-herb community, Primula vulgaris-Hyperichum pulchrum sub-community

U18 Cryptogramma crispa-Athyrium distentifolium snow-bed

U19 Oreopteris limbosperma-Blechnum spicant community

U20a Pteridium aquilinum-Galium saxatile community, Anthoxanthum odoratum sub-community

U20c Pteridium aquilinum-Galium saxatile community, Species-poor sub-community

U21 Cryptogramma crispa-Deschampsia flexuosa community

S9a Carex rostrata swamp, Carex rostrata sub-community

S9b Carex rostrata swamp, Menyanthes trifoliata-Potamogeton polygonifolius sub-community

S10a Equisetum fluviatile swamp, Equisetum fluviatile sub-community

OW + Menyanthes trifoliata vegetation in open water *Men-yanthes

Je Species-poor Juncus effusus-dominated vegetation *

Rac. eri Racomitrium ericoides shingle vegetation *

FF Fell-field *

G Garden *

AR Acid Rock *

BR Basic Rock *

AS Acid Scree *

BS Basic Scree *

BF Boulder-field *

Poh. lud. Pohlia ludwigii snow-bed *

Shingle Almost bare river shingle *

OW Open Water *

Note: these are all vegetation types of the National Vegetation Classification except those marked with an asterisk, which donot fit into any NVC type and are therefore classed separately.

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5 EVALUATION

5.1 General description of the site

Glen Coe is one of the wildest and most scenic glens in Scotland. The main glen is a great U-shape,deepened and oversteepened by ice and walled in by colossal precipices. The pinnacled battlements of theAonach Eagach rise on the north side, and the black near-vertical cliffs of the Three Sisters to the south.Between the jutting promontories of the Three Sisters, steep green glens run back into the hills and terminatein high rocky corries under the summit ridge. The study area falls into four distinct regions: Meall Mór; Sgorr nam Fiannaidh, Aonach Eagach and Meall Dearg; Bidean nam Bian and its subsidiary tops; and thedistinct peaks of Buachaille Etive Beag and Buachaille Etive Mór.

The tall green triangle of Meall Mór guards the western end of the glen. It rises in a long, steep sweep ofgrassland, dotted over with wooded crags, to a level summit.

Sgorr nam Fiannaidh and Stob Coire Leith are two summits at either end of a high-level ridge draped insilvery screes. To their east the black towers of Aonach Eagach lead to the blunt red cone of Meall Dearg.The south-facing slopes drop to the main glen in almost vertical tiers of craggy slopes cut through bynumerous stream gullies.

Bidean nam Bian, the peak of the bens, is well-named. It consists of a series of pointed tops linked by anarrow and airy ridge. The ridge runs from north-west to south-east and divides the north-facing corries wheresnow blows into deep drifts in winter and where montane snow-bed vegetation covers the ground, from thesteep slopes facing south-west where sub-montane grasslands reach nearly to the summit. The long narrowridges of the Three Sisters – Aonach Dubh, Gearr Aonach and Beinn Fhada – reach out to the north. Theirsides drop in dark, tree-bedecked precipices to the green glens.

To the east the pink granite cones of Buachaille Etive Beag and Buachaille Etive Mór rise in solitary grandeurfrom the wet heaths and bogs of Rannoch Moor. They are divided from each other by the long, smooth-sided, symmetrical green glen of the Lairig Gartain, and from the main group of hills by the Lairig Eilde.

5.2 Vegetation

We recorded 129 types of vegetation and habitat here – more than on any other site we have yet surveyed.

5.2.1 General patterns of vegetation

The glens and the lower slopes are green and grassy, and heaths persist only on the steepest ground. Thispattern is the result of a long history of grazing, but there is still much woodland on the cliffs, in the ravinesand in the lower glen of the Allt Coire Gabhail.

Around the two Buachailles there is much Trichophorum-Eriophorum blanket bog M17, with Erica-Sphagnumbog M18 in the wetter reaches as well as many fine examples of bog pools. In Glen Etive, the LairigGartain, the Lairig Eilde and in the main glen there are Trichophorum-Erica wet heaths M15 and Molinia-Potentilla wet grasslands M25, with smaller patches of Trichophorum-Eriophorum bog M17. In the main glenthere are semi-improved fields with Festuca-Agrostis-Galium grassland U4b.

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On the steep north-facing and east-facing slopes of the main ridge there is much Oreopteris limbospermafern-dominated vegetation U19, mossy Festuca-Agrostis-Galium grassland U4d and bracken U20. The greenvegetation on these slopes is also made up of Vaccinium-Deschampsia heaths H18, Vaccinium-Sphagnumheaths HX, Nardus-Galium grassland U5, Trichophorum-Erica wet heath M15 and small patches of Festuca-Agrostis-Thymus grassland CG10. Higher up these pass into more montane communities such as Vaccinium-Racomitrium heath H20, Vaccinium-Rubus heath H22, Luzula sylvatica swards U16c and Deschampsia-Galium grassland U13a. On the cliffs there are Quercus-Betula-Dicranum woodlands W17, dark patchesof Calluna-Vaccinium-Sphagnum damp heath H21 and bright green areas of Luzula-Vaccinium tall-herbvegetation U16. Where the rocks are more basic there are herb-rich swards of the Luzula-Geum community U17.

On the south-facing and west-facing slopes of the main ridge and the Aonach Eagach there are similarmixtures of vegetation. Here there is generally more Festuca-Agrostis-Thymus grassland CG10, Festuca-Agrostis-Alchemilla grassland CG11 and also more bracken, at least in the main glen. There is not so muchmossy grassland or Oreopteris limbosperma vegetation, and instead of the damp heaths there are Calluna-Erica H10 and Calluna-Vaccinium H12 dry heaths.

In the north-facing corries there are intriguing mixtures of snow-bed vegetation. The pale swards of Nardus-Carex grassland U7 and the green expanses of Deschampsia-Galium grassland U13a and Vaccinium-Racomitrium montane heaths H20, and the rich shades of the Polytrichum-Kiaeria snow-beds U11, Salix-Racomitrium snow-beds U12, Rhytidiadelphus loreus snow-beds U13b and Alchemilla-Sibbaldia snow-bedsU14 form a colourful patchwork over the ground. This is threaded with a brightly-coloured lacework ofPhilonotis-Saxifraga springs M32, Anthelia-Sphagnum springs M31 and Pohlia wahlenbergii var. glacialissprings M33.

On the summit ridges there are Calluna-Racomitrium heaths H14, mossy Nardus-Galium grasslands U5eand, on the highest ground, Carex-Racomitrium heaths U10. Some of these belong to the herb-rich formU10c. In hollows there are Nardus-Carex snow-beds U7, and there are also patches of Trichophorum-Ericaheath M15, locally with many montane species in the sward. On the top of Bidean nam Bian and on thehills on the north side of the glen there are small patches of Carex-Polytrichum heath U8.

The vegetation of Meall Mór is rather different, because of the limestone rocks. There are Molinia-Potentillagrasslands M25 on the steep slopes: some with a mixed grassy sward; some very herb-rich. There are largeexpanses of Festuca-Agrostis-Thymus grassland CG10 and herb-rich Festuca-Agrostis-Galium grassland U4F,small patches of Festuca-Alchemilla-Silene swards CG12 and, on crags, Dryas-Silene ledge vegetationCG14. There are also some fine examples of Luzula-Geum vegetation U17 and herb-rich woodland W7and W9. On the upper cliffs there are a few patches of Salix myrsinites scrub W20. The summit of the hillis acid and there are the same mixtures of montane heaths and grasslands as there are elsewhere in thestudy area. More notably, there are tiny patches of lichen-rich Calluna-Cladonia heath H13 and largerpatches of lichen-rich Nardus-Galium grassland U5e.

There are small Carex-Pinguicula M10, Carex-Saxifraga M11 and Carex-Sphagnum M4 and M6 miresthroughout the study area, on the slopes and in the glens. In the main glen there are big, dark-green patchesof Juncus acutiflorus-Sphagnum mire M6d, and on Meall Mór there are J. acutiflorus-Galium rush pasturesM23a.

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5.2.2 Wider Brit ish and European distributions of vegetation types

Some vegetation types found here are widespread and common in Britain, at least in the northern andwestern upland areas. Other types are scarce in Britain. Similarly, some types have counterparts which arewidespread or common in mainland Europe, while others are markedly oceanic and in Europe are foundonly the oceanic west, especially Britain and Ireland. Table 2 shows the British distribution of the differentNVC types recorded in this survey, and the wider European distribution of similar vegetation.

5.2.3 Uncommon vegetation types

Several vegetation types found at this site are especially interesting because they are uncommon in Great Britain. They are:


W9b Fraxinus excelsior-Sorbus aucuparia-Mercurialis perennis woodland, Crepis paludosa sub-community




W25D Pteridium aquilinum-Rubus fruticosus agg underscrub, damp sub-community



H18b Vaccinium myrtillus-Deschampsia flexuosa heath, Alchemilla alpina-Carex pilulifera sub-community

H20a Vaccinium myrtillus-Racomitrium lanuginosum heath, Viola riviniana-Thymus polytrichus sub-community

H20b Vaccinium myrtillus-Racomitrium lanuginosum heath, Cetraria islandica sub-community

H20c Vaccinium myrtillus-Racomitrium lanuginosum heath, Bazzania tricrenata-Mylia taylori sub-community

H21b Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath, Mastigophora woodsii-Herbertus aduncus hutchinsiae sub-community

H22a Vaccinium myrtillus-Rubus chamaemorus heath, Polytrichum commune-Galium saxatile sub-community

M7 Carex curta-Sphagnum russowii mire

M11a Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Thalictrum alpinum-Juncus triglumis sub-community

M11b Carex viridula ssp. oedocarpa-Saxifraga aizoides mire, Palustriella commutata-Eleocharis quinqueflorasub-community


M25c Molinia caerulea-Potentilla erecta mire, Angelica sylvestris sub-community




CG10c Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland, Saxifraga aizoides-Ditrichum flexicaulesub-community

CG11a Festuca ovina-Agrostis capillaris-Alchemilla alpina grassland, Typical sub-community


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CG14 Dryas octopetala-Silene acaulis community

U4eA Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Vaccinium myrtillus-Deschampsia flexuosasub-community, Alchemilla alpina variant

U4F Festuca ovina-Agrostis capillaris-Galium saxatile grassland, provisional Filipendula ulmaria sub-community


U6R Juncus squarrosus-Festuca ovina grassland, provisional herb-rich sub-community

U7a Nardus stricta-Carex bigelowii grass heath, Empetrum nigrum hermaphroditum-Cetraria islandicasub-community







U12b Salix herbacea-Racomitrium heterostichum snow-bed, Gymnomitrion concinnatum sub-community

U13a Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemilla alpinasub-community

U13ahr Deschampsia cespitosa-Galium saxatile grassland, Anthoxanthum odoratum-Alchemilla alpinasub-community, herb-rich form




U17a Luzula sylvatica-Geum rivale tall-herb community, Alchemilla glabra-Bryum pseudotriquetrum sub-community


U17d Luzula sylvatica-Geum rivale tall-herb community, Primula vulgaris-Hyperichum pulchrum sub-community


U21 Cryptogramma crispa-Deschampsia flexuosa community

Pohlia ludwigii snow-bed

Fell-field

Most of these are not very extensive at this site. However, M25c and U4F are quite extensive on Meall Mór,and there are large areas of U13a in the higher corries on the north side of the Bidean ridge.

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The vegetation types which are interesting because they have markedly western, oceanic distributions inEurope and are most common in Britain and Ireland are:

W11a Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Dryopteris dilatata sub-community

W11b Quercus petraea-Betula pubescens-Oxalis acetosella woodland, Blechnum spicant sub-community



W25 Pteridium aquilinum-Rubus fruticosus agg underscrub

H10 Calluna vulgaris-Erica cinerea heath (all sub-communities)



H20 Vaccinium myrtillus-Racomitrium lanuginosum heath (all sub-communities)

H21 Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath (all sub-communities)



M15 Trichophorum cespitosum-Erica tetralix wet heath (all sub-communities)

M17 Trichophorum cespitosum-Eriophorum vaginatum blanket mire (all sub-communities)


M25 Molinia caerulea-Potentilla erecta mire (all sub-communities)


U20 Pteridium aquilinum-Galium saxatile community (all sub-communities)

Among these, H10, M15, M17, M25 and U20 are extensive here.

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Table 2 Brit ish distribution of NVC types recorded in this sur vey at Glen Coe in 2002, and European distribution of similar vegetation

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15W E Or S N Wa SW SE Ir Fr Sc Fa CE Sp NHi Hi + Sc En le En En el Ge an er Eu + Af

NVC gh gh Sh ot gl s gl gl and Ho di oe ro Po ri

W3 + + . + + . . . . . * . . . .W4 +++ +++ . ++ ++ ++ ++ + c c * . . . .W7b +++ +++ . ++ +++ ++ ++ + ? c . . c . .W7c ++ ++ . ++ +++ +++ + + ? c . . c . .W9b +++ +++ . + ++ . . . c . c . . . .W11a + . . ++ +++ +++ ++ . ? c . . . . .W11b +++ . . + + + . . * c * . . . .W17a +++ + . + ++ ++ + . ++ ? * . . . .W17b +++ +++ . ++ ++ +++ + . * ? ? . . . .W17c +++ +++ . ++ ++ +++ + . * ? ? . . . .W18d + ++ . . . . . . . . * . . . .W20 + + . . . . . . . . * . . . .W24 ++ ++ ? ++ +++ +++ +++ +++ * * . . . . .W25 +++ +++ ? +++ +++ +++ +++ +++ * ? . . . . .H10a +++ ++ ++ +++ + + + . +++ . c * . . .H10b ++ + ? + + . . . ? . c ? . . .H10c +++ ++ . +++ + + . . ? . c ? . . .H10d ++ + + + . . . . ? . c ? . . .H12bc + +++ + +++ +++ +++ ++ . . . * . . . .H13a + +++ . + + + . . . . * . . . .H14 ++ + + . . . . . * . . . . . .H18ab ++ +++ . ++ ++ ++ + . . . * . . . .H20a + + . . . . . . . . . . . . .H20b ++ ++ . . . . . . . . ? . . . .H20c + . . . . . . . . . . . . . .H21a +++ ++ ++ ++ ++ + . . * . * . . . .H21b ++ . + . + + . . * . . * . . .H22a + ++ . + . . . . . . ? . . . .M1 +++ . . ++ ++ ++ ++ + * . * . . . .M3 +++ +++ . ++ +++ + . . . * * . . . .M4 ++ ++ . ++ ++ ++ + . . . . . . . .M6abcd +++ +++ +++ +++ +++ +++ +++ . * * * . . . .M7 + ++ . . + . . . . . * . . . .M10a +++ +++ +++ +++ +++ ++ . . . * * . * . .

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Table 2 (continued)



M11ab +++ +++ + + ++ + . . . * * . * . .M12 ++ ++ . . . . . . . . * * . . .M15a +++ ++ ++ ++ + + + . +++ . * * . . .M15b +++ ++ ++ ++ + + + . +++ . * * . . .M15c +++ + ++ ++ + + + . +++ . * * . . .M15d + + + ++ ++ ++ ++ . ? . ? ? . . .M17a +++ ++ ++ ++ + ++ + . +++ . . * . . .M17b +++ ++ ++ ++ . + . . ? . . ? . . .M18a + +++ . +++ ++ ++ . + . c . . . . .M18b + ++ . ++ ++ ? . . . c . . . . .M19a +++ ++ ++ +++ ++ ++ . . * . c . c . .M23a +++ ++ . +++ ++ ++ ++ + * . . . . . .M25a +++ ++ . +++ ++ +++ ++ + ? . . . . . .M25b + . . ++ + +++ ++ . ? . . . . . .M25c + . . . + ++ +++ . ? . . . . . .M28a +++ . +++ . . + + . * . * . . * *

M31 ++ ++ . + + + . . ? . ? ? . . .M32ab ++ +++ ++ ++ ++ + ? . . . * * * . .M33 + ++ . . . . . . . . * . . . .M37 ++ +++ . ++ ++ + ? ? ? ? ? . ? . .U4a +++ +++ +++ +++ +++ +++ +++ . * ? ? * . . .U4b +++ +++ +++ +++ +++ +++ +++ . ? ? ? ? . . .U4d ++ ++ ++ ++ + + . . ? ? ? ? . . .U4e + + ? ++ ++ ++ ++ . ? ? ? ? . . .U5a +++ +++ ? +++ +++ +++ ? . . c c . . . .U5b ++ ++ ? +++ +++ +++ ? . . c c . . . .U5c + ++ + + + . . . . c c . . . .U5e +++ ++ +++ ++ + . . . . c c * . . .U6a ++ ++ ? +++ +++ ++ ? . . c c c . c .U6c ++ ++ ? +++ +++ ++ ? . . c c c . c .U6d + ++ ? +++ +++ ++ ? . . c c c . c .U7a ++ ++ . . . . . . . . c c . . .U7b ++ ++ . + . . . . . . c c . . .U7c + + . + + . . . . . c c . . .

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Table 2 (continued)



U8 + ++ . . . . . . . . * . . . .U10a +++ +++ . + + + . . * . ? ? . . .U10b +++ +++ . + + . . . ? . * +++ . . .U10c ++ + . + . . . . . . * ++ . . .U11a + ++ . . . . . . . . * . * . .U12ab + ++ . + . . . . . . c . c . .U13a ++ ++ . + . . . . . . c . . . .U13b ++ ++ . + . . . . . . c . . . .U14 + ++ . . . . . . . . * . . . .U15 ++ ++ . . + . . . . . * . . . .U16a ++ ++ . ++ + . . . c . c c . . .U16b ++ ++ ++ ++ + . . . c . c c . . .U16c ++ ++ ++ ++ + . . . c . c c . . .U17a + + . + + + . . . . ? ? . . .U17b + + . . . . . . . . * ? . . .U17d ++ . ++ . . . . . . . ? ? . . .U18 + ++ . . . . . . . . * . . . .U19 ++ ++ ? ++ ++ ++ + . . . ? . . . .U20ac +++ +++ +++ +++ +++ +++ +++ ++ * . . . . . .U21 + + . + ++ ++ . . ? ? ? . ? ? .CG10a +++ +++ ++ ++ +++ ++ . . ? ? ? * ? . .CG10b +++ +++ ++ + + + . . ? ? ? * ? . .CG10c + + + . . . . . . ? ? ? ? . .CG11a ++ ++ . . + . . . . . ? * . . .CG12 + ++ . . . . . . . . * . . . .CG14 + ++ . . + + . . . . * . . . .MG9 + + ? + ++ + ++ +++ ? * . . ? . .S9ab ++ ++ ++ ++ ++ ++ . + ? ? ? . . . .S10a ++ ++ ++ ++ ++ ++ ++ ++ ? ? ? . . . .

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Note: vegetation types not described in the NVC are omitted from this table because there is insufficient information availableabout their British and European distributions.

Key to areas1 W Highlands 4 S Scotland 7 SW England 10 France/Germany/Belgium/Holl./Lux. 13 Central Eur.2 E Highlands 5 N England 8 SE England 11 Scandinavia 14 Spain/Port.3 Orkney/Shetl. 6 Wales 9 Ireland 12 Faroe Islands 15 N AfricaKey to symbols+ scarce ++ fairly common +++ common * present ? possibly presentc present, but information only meaningful at NVC community level

5.2.3.1 Qualifying EC Habitats Directive Annex I Habitats

Several vegetation types found here are so scarce in Europe that they are listed among the Annex I habitatsin the EC Habitats Directive. Some of these are considered by Scottish Natural Heritage to be importantenough here to count as ‘Qualifying Annex I Habitats’ for the proposed Glencoe Special Area for Conservation:

Alpine and boreal heaths

NVC types found here and assigned to this EC Annex I type by UK nature conservation agencies:

H13 Calluna vulgaris-Cladonia arbuscula heath


H20 Vaccinium myrtillus-Racomitrium lanuginosum heath

H21 Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath (where above tree-line)

H22 Vaccinium myrtillus-Rubus chamaemorus heath

The montane prostrate heaths H13 and H14 are not common, because there is so little high rocky plateau.H13 occurs only on Meall Mór and covers only a few square metres. It is interesting because H13 is morecommon in the more boreal climate of the north and east Highlands than it is in the south and west, thoughthere are also fragments on Ben Lui (Averis & Averis 1999a). H14 is most extensive on the exposed tops ofthe Three Sisters. H20 is very extensive on the upper slopes, especially on steep, rocky and broken ground.Examples of H21 above the presumed tree-line occur on some little-grazed or ungrazed middle slopes facingnorth, and H22 is quite common on the upper slopes. H20c and H21b heaths contain excellent, internationallyimportant assemblages of oceanic bryophytes.

European dr y heaths


H10 Calluna vulgaris-Erica cinerea heath

H12 Calluna vulgaris-Vaccinium myrtillus heath

H21 Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath (where below tree-line)

There are large patches of H10 and H21 and smaller areas of H12 on many steep slopes and cliffs: H10 mostly faces from W through S to SE, and H21 mostly from NW through NE to E. The vegetation is madeup mainly of nationally common species and the flora is generally not very species-rich, but the H21b sub-communitycontains excellent, internationally important assemblages of oceanic bryophytes. H10, H12 and H21a arecommon in the Highlands, but this does not reduce the importance of the vegetation on a European scale.

Sil iceous scree of the montane to snow levels



Also AS Acidic scree (a non-NVC type) which is associated with snowbed vegetation

These vegetation and habitat types are common in the upper parts of the high corries. They form part ofsome of the most impressive examples of snowbed vegetation in the west Highlands.

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Sil iceous alpine and boreal grasslands


U7 Nardus stricta-Carex bigelowii grass heath


U10 Carex bigelowii-Racomitrium lanuginosum moss heath

U11 Polytrichum sexangulare-Kiaeria starkei snow-bed

U12 Salix herbacea-Racomitrium heterostichum snow-bed

U14 Alchemilla alpina-Sibbaldia procumbens snow-bed

U7 snow-bed grasslands are common in the upper corries; they also occur in small patches on summitridges. Their flora is generally not very notable in a British context but the comparative scarcity of similarvegetation in mainland Europe gives it special importance here. U11, U12 and U14 snowbeds andvegetation intermediate between these types cover large areas in the upper parts of the high corries. Theyconstitute some of the most impressive examples of snowbed vegetation in the west Highlands. They aremore extensive and varied than those on Ben Lui (Averis & Averis 1999a). U8 is scarce on damp ground inthese same areas. U10 occurs on many exposed summit ridges but is not extensive.

Sil iceous rocky slopes with chasmophytic vegetation

The habitat found here which appears to belong to this EC Annex I type is:

AR Acid rock outcrops and cliffs. Note: this habitat and vegetation is not described in the NVC

This habitat is very common throughout most of this site. The flora of acid rocks can be rather species-poor,but in many places the habitat is intimately associated with bryophyte-rich woodlands, heaths and snow-beds of obvious high value for nature conservation.

Alpine and subalpine calcareous grasslands



CG14 Dryas octopetala-Silene acaulis heath

CG12 and CG14 occur as small patches on the steep upper north-eastern slope of Meall Mór. They havea rich flora, and are undoubtedly of great value for nature conservation.

Species-rich Nardus grassland on si l iceous substrates in mountain areas


CG10 Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland

CG11 Festuca ovina-Agrostis capillaris-Alchemilla alpina grassland


U4F Festuca ovina-Agrostis capillaris-Galium saxatile grassland, Filipendula ulmaria type

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The herb-rich grasslands U5c, CG10, CG11 and U4F occur throughout the study area, except on the acidrocks of the two Buachailles. Only on Meall Mór are there large areas of CG10, CG11 and U4F, but theyare impressive and herb-rich. U5c, as usual, occurs in small patches on flushed slopes.

Calcareous rocky slopes with chasmophytic vegetation


BR Basic rock outcrops and cliffs. Note: this habitat and vegetation is not described in the NVC

This habitat occurs in several places on Meall Mór and on igneous outcrops on the north side of Bidean.The flora can be quite rich, and this habitat is typically accompanied by interesting species-rich, herb-richgrassland, woodland or tall herb vegetation.

Alpine pioneer formations of the Caricion bocoloris-atrofuscae


M11 Carex viridula ssp. oedocarpa-Saxifraga aizoides mire


M11 mires are common in the study area, on flushed slopes from the lower hillsides up into the high corries.M12 has been found only in the upper part of Coire nam Beitheach. The M11 and M12 mires are never large.

Alkaline fens


M10 Carex dioica-Pinguicula vulgaris mire

M10 mires are common on the hill slopes at this site. They are all small.

Hydrophilous tall herb fringe communities of plains and montane to alpine levels


(U15 Saxifraga aizoides-Alchemilla glabra banks)

U17 Luzula sylvatica-Geum rivale tall-herb community

Only U17 is included in the current definition of this Annex I type, but U15 is also mentioned here becauseit occurs within the same area as does U17 on the steep NE slope of Meall Mór and has a similar floraand habitat to U17. U17 also occurs widely on cliffs elsewhere at this site.

Sub-arctic Salix sp. scrub



W20 occurs on the east-facing upper slopes of Meall Mór. The patches are tiny and are confined to cragsand steep broken ground.

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5.2.3.2 Non-qualifying EC Annex I Habitats

Several other vegetation and habitat types found in this survey belong to EC Annex I habitats, but areconsidered by Scottish Natural Heritage as ‘Non-qualifying Annex I Habitats’ for the Glen Coe Special Areafor Conservation:

Nor thern Atlantic wet heaths with Erica tetralix


M15 Trichophorum cespitosum-Erica tetralix wet heath

M15 is widespread and extensive on the lower ground, but ascends to over 800m. There is a wide rangeof forms of oceanic wet heath in the study area, and it occurs over a particularly large altitudinal range here.The most montane forms of M15 which we have found here add to the range of wet heathland typesdescribed so far from Britain; previously, montane forms of British wet heath were hardly described.

Transit ion mires and quaking bogs



M4 is scarce at this site. There are small patches of it among bogs in the north-eastern corner and on theshores of Loch Achtriochtan.

Blanket bog (active)


M17a Trichophorum-Eriophorum vaginatum blanket mire, Drosera-Sphagnum sub-community

M18a Erica tetralix-Sphagnum papillosum mire, S. magellanicum-Andromeda sub-community

M19 Calluna vulgaris-Eriophorum vaginatum blanket mire

Bogs of these types are widespread, common and locally extensive on the lower ground at this site. Thereare excellent examples of blanket bog in two areas in the NE of the site, with good and varied Sphagnumcarpets, some uncommon plant species and much small-scale variation in wetness and microtopography.These bogs are evidently little-disturbed and are of obvious high value for nature conservation.

Depressions on peat substrates of the Rhynchosporion



There are many small patches of this habitat among the wetter parts of M17 and M18 bog in the north-eastof the study area (see under ‘Blanket bog (active)’ (above)).

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Calcareous and calcshist screes of the montane to alpine levels


BS Basic scree. Note: this habitat and vegetation is not described in the NVC

This is rare in the study area: there are small patches among a few scree-fans in the Lairig Eilde and theLairig Gartain.

Petrifying springs with tufa formation


M37 Palustriella commutata-Festuca rubra spring

There are many examples of M37 on Meall Mór, and a few in the corries of Bidean and on the slopes ofAonach Eagach.

Old sessile oak woods with I lex and Blechnum in the Brit ish Isles


W11 Quercus petraea-Betula pubescens-Oxalis acetosella woodland

W17 Quercus petraea-Betula pubescens-Dicranum majus woodland

The W11 and W17 woodland here has a canopy dominated by birch. This does not accord strictly withthe current JNCC definition of this Annex I type which is defined as having oak present or showing evidenceof past or future potential presence of oak. However, the woods on the northern slopes of the Three Sistersand from Allt Coire Gabhail and the Lost Valley boulder-field down to and along the River Coe are worthyof mention because they are of great international importance for oceanic bryophytes. They are among thebest British (and European) woods for these plants. Most other patches of W11 and W17 at this site aresmall.

Alluvial forests with Alnus glutinosa and Fraxinus excelsior


W7c Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland, Deschampsia cespitosa sub-community

This habitat is confined here to the extreme north-west of the site, on level ground in the valley floor: it isalder woodland with a lush, species-poor grassy ground layer.

One additional ‘non-qualifying’ Annex I habitat type considered by Scottish Natural Heritage to be presentin the site is the aquatic habitat: ‘Oligotrophic to mesotrophic standing waters with vegetation of theLittorelletea uniflorae and/or the Isoeto-Nanojuncetea’. We did not record any vegetation which seemed tous to belong to this EC habitat type, though our survey concentrated on terrestrial vegetation and we did notventure into lochs to look at the aquatic flora.

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5.3 Flora

This site has a rich flora. We found 275 vascular plant species and 243 bryophytes. These species arelisted in Appendix 2. Some other species have been found here by other people.

5.3.1 Phytogeography of the flora

Appendix 2 shows the placement of each species into the European phytogeographical groups by Preston& Hill (1997) (for vascular plants) and Hill & Preston (1998) (for bryophytes). The flora of Glen Coeencompasses a wide range of European phytogeographical groups (Table 3). About half of the vascularplants species and about a third of the bryophytes found in this survey are widespread in Europe, with nonorthern, southern, oceanic or continental tendency. Many others have a moderate northern, southern orwestern bias in their European distributions, and a significant minority are more distinctly northern, southernor western. Species in these last three groups are listed below.

17% of vascular plants and 16% of bryophytes have strongly northern distributions in Europe:

Vascular plants: Alchemilla alpina, Arabis petraea, Athyrium distentifolium, Carex bigelowii,C. capillaris, C. dioica, C. saxatilis, Cerastium alpinum, C. arcticum, Cochlearia pyrenaica,Cornus suecica, Diphasiastrum alpinum, Draba incana, Dryas octopetala, Empetrum nigrum,Epilobium alsinifolium, E. anagallidifolium, Eriophorum vaginatum, Festuca vivipara,Gnaphalium supinum, Huperzia selago, Juncus biglumis, J. trifidus, J. triglumis, Luzulaspicata, Lycopodium annotinum, Oxyria digyna, Persicaria vivipara, Poa alpina, P. glauca,Salix herbacea, S. myrsinites, S. phylicifolia, Saussurea alpina, Saxifraga aizoides,S. cernua, S. nivalis, S. oppositifolia, S. rivularis, S. stellaris, Sedum rosea, Sibbaldiaprocumbens, Silene acaulis, Thalictrum alpinum, Tofieldia pusilla, Vaccinium uliginosum, V. vitis-idaea.

Mosses: Andreaea rupestris, Aulacomnium turgidum, Bartramia ithiphylla, Blindia acuta,Brachythecium glaciale, Calliergon sarmentosum, C. stramineum, C. trifarium, Conostomumtetragonum, Dichodontium pellucidum, Dicranum fuscescens, Distichium capillaceum,Drepanocladus revolvens, Fissidens osmundoides, Hypnum callichroum, Kiaeria blyttii, K. falcata, K. starkei, Oligotrichum hercynicum, Orthothecium rufescens, Philonotis seriata,Plagiobryum zieri, Pogonatum urnigerum, Pohlia ludwigii, Polytrichum alpinum, P. strictum,Racomitrium lanuginosum, R. sudeticum, Scorpidium scorpioides, Sphagnum fuscum, S. girgensohni.

Liverworts: Anastrophyllum minutum, Anthelia julacea, A. juratzkana, Blepharostomatrichophyllum, Gymnomitrion concinnatum, Marsupella alpina, M. sphacelata, Preissiaquadrata, Ptilidium ciliare, Tritomaria quinquedentata.

3% of vascular plants and 18% of bryophytes have markedly western, oceanic distributions in Europe:

Vascular plants: Carex binervis, Conopodium majus, Erica cinerea, Hyacinthoides non-scripta, Hymenophyllum wilsonii, Myosotis secunda, Narthecium ossifragum, Saxifragahypnoides, Ulex europaeus.

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Mosses: Andreaea alpina, Breutelia chrysocoma, Campylopus atrovirens, C. setifolius,Glyphomitrium daviesii, Hymenostylium insigne, Hyocomium armoricum, Isothecium holtii, I. myosuroides var. brachythecioides, Molendoa warburgii, Oedipodium griffithianum,Paraleptodontium recurvifolium, Ptychomitrium polyphyllum, Rhabdoweisia crenulata,Sphagnum strictum, Trichostomum hibernicum, Ulota phyllantha.

Liverworts: Anastrophyllum donnianum, Aphanolejeunea microscopica, Bazzania pearsonii,Colura calyptrifolia, Douinia ovata, Frullania teneriffae, Gymnomitrion crenulatum,Harpalejeunea molleri, Herbertus aduncus, H. stramineus, Lejeunea patens, Lepidoziapearsonii, Leptoscyphus cuneifolius, Mastigophora woodsii, Metzgeria leptoneura,Plagiochila carringtonii, P. exigua, P. killarniensis, P. punctata, P. spinulosa, Pleuroziapurpurea, Radula aquilegia, Saccogyna viticulosa, Scapania gracilis, S. nimbosa, S. ornithopodioides.

Species with distinctly southern European distributions are much scarcer than northern species here. The onlyones which we have seen are the moss Funaria obtusa and the liverworts Calypogeia arguta, Porellaarboris-vitae and Scapania compacta. They make up only 2% of the bryophyte flora in this survey.

We did not find any species with distinctly continental European distributions here; there are not many suchspecies in Britain.

We found only five non-native species in this survey. There are a few trees of Acer pseudoplatanus atAchtriochtan. Epilobium brunnescens is moderately common on damp, flushed ground and along the stonypaths. Juncus tenuis is rare and was found on a track. Mentha spicata occurs on disturbed ground aroundthe new car-park near the road in the main glen. There is a single stunted plant of Picea sitchensis in thescrees of upper Coire nan Lochan. The moss Campylopus introflexus grows on bare peat in dried-outTrichophorum-Eriophorum bog M17b in the north of the site. There does not seem to be a threat of any ofthese species increasing in quantity and causing a reduction in the botanical interest of the site. Acerpseudoplatanus is a fast-growing tree which is commonly planted as shelter around houses. Unless it occursin woodland it is unlikely to spread. Epilobium brunnescens originates from New Zealand. It was firstrecorded in 1904 in Edinburgh (Preston et al. 2002) and is now well-established in wet and stony habitatsthroughout Britain. Mentha spicata – the common spearmint which is used to make mint sauce – is a gardenescape which has been naturalised in Britain for hundreds of years. Juncus tenuis is a native of Americawhich has spread rapidly since the late 19th century (Preston et al. 2002). It grows on damp, open ground,especially on paths and tracks. Picea sitchensis is a common tree of coniferous plantations and the plant inthe study area probably originated as a wind-sown or bird-sown seed. Campylopus introflexus is a colonistof bare peat and is native in the southern hemisphere. It was first recorded in Britain in 1941 (Hill et al.1992) but has now spread all over Britain and Ireland.

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Table 3 Number of species in each phytogeographical group, and their percentage

representation of the total f lora recorded in this sur vey at Glen Coe in 2002

Vascular plants Bryophytes

No. % No. %

11 Oceanic Arctic-montane 0 0 0 0

12 Suboceanic Arctic-montane 0 0 1 <1

13 European Arctic-montane 13 5 1 <1

14 Eurosiberian Arctic-montane 1 <1 0 0

15 Eurasian Arctic-montane 2 <1 0 0

16 Circumpolar Arctic-montane 18 7 8 3

21 Oceanic Boreo-arctic Montane 0 0 0 0

22 Suboceanic Boreo-arctic Montane 0 0 0 0

23 European Boreo-arctic Montane 1 <1 2 1

24 Eurosiberian Boreo-arctic Montane 0 0 1 <1

26 Circumpolar Boreo-arctic Montane 12 4 28 12

32 Suboceanic Wide-boreal 0 0 1 <1

34 Eurosiberian Wide-boreal 1 <1 0 0

35 Eurasian Wide-boreal 1 <1 0 0

36 Circumpolar Wide-boreal 10 4 12 5

41 Oceanic Boreal-montane 1 <1 16 7

42 Suboceanic Boreal-montane 2 <1 9 4

43 European Boreal-montane 3 1 12 5

44 Eurosiberian Boreal-montane 3 1 0 0

45 Eurasian Boreal-montane 1 <1 0 0

46 Circumpolar Boreal-montane 11 4 11 5

51 Oceanic Boreo-temperate 2 <1 1 <1

52 Suboceanic Boreo-temperate 2 <1 11 5

53 European Boreo-temperate 19 7 22 9

54 Eurosiberian Boreo-temperate 24 9 1 <1

55 Eurasian Boreo-temperate 16 6 0 0

56 Circumpolar Boreo-temperate 26 9 33 14

63 European Wide-temperate 0 0 0 0

64 Eurosiberian Wide-temperate 3 1 0 0

65 Eurasian Wide-temperate 2 <1 0 0

66 Circumpolar Wide-temperate 2 <1 5 2

70 Hyperoceanic Temperate 0 0 7 3

71 Oceanic Temperate 6 2 3 1

72 Suboceanic Temperate 10 4 12 5

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Table 3 (cont inued)

Vascular plants Bryophytes

No. % No. %

73 European Temperate 41 15 18 7

74 Eurosiberian Temperate 9 3 0 0

75 Eurasian Temperate 2 <1 0 0

76 Circumpolar Temperate 4 1 3 1

80 Hyperoceanic Southern-temperate 0 0 13 5

81 Oceanic Southern-temperate 1 <1 3 1

82 Suboceanic Southern-temperate 4 1 0 0

83 European Southern-temperate 8 3 1 <1

84 Eurosiberian Southern-temperate 4 1 1 <1

85 Eurasian Southern-temperate 1 <1 1 <1

86 Circumpolar Southern-temperate 1 <1 1 <1

91 Mediterranean-Atlantic 0 0 0 0

92 Submediterranean-Subatlantic 0 0 4 2

93 Mediterranean-montane 0 0 0 0

? Uncertain 3 1 0 0

i Introduced 5 2 1 <1

Total 275 100 243 100

5.3.2 Uncommon plant species

Many plant species which we found in this survey are uncommon at least locally in Great Britain. Thesespecies are listed in Table 4. The ‘target notes’ for the locations of species of particular interest are given inAppendix 3, and their locations are shown on the 1:10,000 maps in Appendix 4.

5.3.2.1 Nationally scarce and nationally rare vascular plants

Many vascular plant species recorded at this site are nationally scarce (recorded in 16–100 10km squaresin Great Britain since 1950) or nationally rare (<16 squares). Staff of Scottish Natural Heritage and theNational Trust for Scotland have a special interest in these plants, so this survey included making a specialsearch for them. We re-found all of the previously recorded species except for Carex lachenalii, C. atrata,Juncus castaneus, Potentilla crantzii and Veronica alpina. Of these, only J. castaneus and P. crantzii havebeen seen in recent years. Details of the populations of the plants, and their locations and habitats, are givenin Table 5. The locations of the montane rare species in upper Coire nam Beitheach are shown inPhotographs 56 and 57. The following paragraphs give a summarized account of the distribution andhabitats of each of the species which we found here in this survey.

Athyrium distentifolium is a montane fern which grows in snow-beds among block scree. It resembles thecommon A. filix-femina of the lowlands but is smaller and the sori or spore-cases on the backs of the leaves

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are round rather than rectangular or comma-shaped. Most of the A. distentifolium in the study area growsamong boulders in the upper part of Coire nam Beitheach, but there are also smaller quantities inCryptogramma-Athyrium snow-beds U18 in Coire nan Lochan and Coire Gabhail. The total population inthe study area is perhaps a few hundred plants.

Carex capillaris is a sedge with yellow-green leaves and small green flowers borne on very thin, droopingstalks. It grows in grassland on the north-eastern slopes of Meall Mór. We saw it here on the lower slopesin the north of the surveyed part of the hill, and also further south within and just downslope from the fencedarea on the upper slopes. We saw about 50 plants. The species is rather inconspicuous and can be hardto find in dense grassland: there are probably many more plants here, perhaps in places in between thetwo areas where we found it. The total population might be a few hundred plants. C. capillaris is not a tallplant. Here and elsewhere in Scotland it grows mainly in grassland where grazing keeps the associatedvegetation from growing so tall and dense as to out-compete it. Within the fenced, ungrazed exclosure onthe upper slopes it seems possible that C. capillaris will suffer as the ungrazed vegetation grows taller.

Carex saxatilis is an uncommon montane sedge which grows in a few base-rich flushes in upper Coire namBeitheach, at about 825m. The leaves are a rich dark green, with russet tips, and the black flowers aredistinctive. It grows in dense swards. At least 100 rosettes were seen in this survey; no flowers were seenon any of these plants. Although C. saxatilis is nationally scarce it is quite plentiful in the Breadalbane hillswith their base-rich rocks, and also in areas of late snow-lie in the central and north-western Highlands. It issurprising that it occurs so locally in the study area.

Cerastium alpinum is the rarer of the two montane Cerastium species in the study area. It is an attractiveplant with grey leaves covered all over with a dense silvery-white felt of fine silky hairs. Again, it grows inCoire nam Beitheach at altitudes over 850m. It grows in the tall-herb ledge vegetation U17 on the lowestband of the cliffs to the west of Collie’s Pinnacle, and on the eastern cliffs of Stob Coire nam Beith, also inU17. The plants are very thinly-scattered. Only 12 individuals were seen in 2002. None were flowering atthe time of the survey.

Cerastium arcticum is the commoner of the two montane Cerastium species in the study area. It is a robustplant with widely-spaced pairs of yellow-green leaves and with large white starry flowers. The leaves andstems are thinly-clad with long hairs. The plant is common in the upper part of Coire nam Beitheach. Itscrambles among taller plants in the tall-herb ledge vegetation U17 and forms pure mats and patches onscree and gravel, on cliff ledges and among boulders. It occurs in most of the montane grassland, snow-bed and spring communities in the corrie. It is so numerous that it is impossible to estimate the populationsize; there are many hundreds if not thousands of plants.

Cystopteris montana is an easily-recognised small fern with clusters of grey-green, 3-pinnate leaves(Photograph 16). Because the leaves are so finely divided they look very lacy and delicate. C. montanagrows in the upper parts of Coire nam Beitheach, above about 900m. It grows in at least four places in thecorrie: on the cliffs and screes between Collie’s Pinnacle and Hourglass Gully and on the spur of rockseparating the two large gullies in the western half of the corrie. In all the sites there are several large, bushyand healthy plants and the total population probably consists of several tens of individuals.

Draba norvegica is another of the montane rarities in Coire nam Beitheach. Its tight rosettes of small leavesare scattered on ledges and beneath overhangs on the cliffs between Collie’s Pinnacle and Hourglass Gully.

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There are at least nine plants. Doubt has been cast on the identity of this species in the past, because onone occasion someone reported seeing some fruiting plants with twisted pods rather like those of D. incana.This could not be checked in 2002 because none of the plants were flowering. The basal rosettes, however,do look more like those of D. norvegica. The general consensus of opinion seems to be that the plants areD. norvegica.

Dryas octopetala grows on the limestone ledges on the eastern and northern sides of Meall Mór. It is asplendid plant. The tiny, wavy-edged tough leaves are dark glossy green on top and silver beneath, the largeflowers are a pale cream-gold and the seed-heads consist of a twisted tuft of long white silken hairs. D. octopetala is reasonably plentiful over a wide area of the hillside; there are certainly more than the 28 plants or clumps recorded by Halcrow (1996). Most of the plants are on the upper slopes betwen 500mand 620m, but it also grows on many outcrops lower down, reaching about 400m on the northern slopes.On the east-facing slopes there are several places where D. octopetala actually grows under trees, sharingthe limestone crags with birch and rowan.

Juncus biglumis is a small rush with stiff, yellow-green grass-like leaves and stout stems each bearing a pairof oval black flowers. It grows in Coire nam Beitheach on the steep, broken ground below the main cliffsbetween Collie’s Pinnacle and Hourglass Gully. Around 30 plants were seen in each of two locations onthese cliffs, but the ground is steep and dangerous and there may well be more. At one site the plant wasgrowing with the rare moss Aulacomnium turgidum.

Pinus sylvestris, though a common sight in plantations and in parks and gardens, is scarce as a nativespecies. It is a tall and statuesque conifer. Its rich red bark, flecked with large greyish scales, contrasts withits dark-green needles. There is a solitary tree in amongst the birches at the stream confluence known as theMeeting of Three Waters, three trees in the Clachaig gully and a fragment of pine woodland together withscattered trees in Glen Etive. Old pine roots are exposed in peat hags in the Lairig Eilde. The patch ofwoodland in Glen Etive has recently been deer-fenced in an attempt to allow the trees to regenerate.

Poa alpina is a montane grass which has pale, almost blue-grey leaves. It is almost always viviparous, soinstead of a spike of flowers there is a cluster of tiny plantlets on a long stalk. P. alpina grows in thin swardsand tufts in tall-herb ledge vegetation on the cliffs of upper Coire nam Beitheach. It is reasonably commonhere, and there are hundreds of plants.

Poa glauca is far less common than P. alpina in the study area. It, too, has glaucous blue-grey leaves andgrows in tufts or thin patches. Unlike P. alpina, this species flowers in Britain and bears panicles of plump,oval, glossy blue-grey flowers. There are several patches of this grass on the lowest tier of the cliffs betweenCollie’s Pinnacle and Hourglass Gully, below the terrace where the rare saxifrages grow. It is also scatteredmore thinly on the lower ledges of the upper band of cliffs. The total population probably numbers severaltens of plants.

Salix myrsinites is a montane willow. It has a tangle of brown twigs bearing bright green leaves which arehairless and shiny on both sides; and in summer it has yellow furry catkins. It grows on limestone crags onthe upper east-facing slopes of Meall Mór, at altitudes between 550m and 620m (Photograph 8). The areain which most of the willows grow was stock-fenced in 2000 and, in two growing seasons, the groundvegetation and that on the smaller outcrops has grown tall and lush. This means that it is less easy than it

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was to see the willows on their ledges, and some of the smaller plants are now quite obscured by tall grassesand herbs. The larger bushes, measuring up to about a metre high and two metres across, are scattered onthe ledges and outcrops and only locally thicken up into a few square metres of scrub. There are also manysmaller, possibly younger plants, which are up to about 50cm tall. The willows have not been browsed fortwo years, but those in the more accessible spots have contorted lower stems which suggests that they havebeen browsed in the past. Halcrow’s (1996) figure of 40 mature plants is probably a conservative estimateand there are many more smaller individuals. There is no evidence to suggest that the population size haschanged since 1995. Not all of the willows were flowering but those that were, in June 2002, were allfemales. Halcrow (1996), visiting the site between July and September, saw no flowers at all.

Saxifraga cernua is a beautiful little plant with lobed, almost hairless, grey-green leaves set off by thebrilliant red bulbils which form in clusters in the axils of the leaves (Photograph 20). It grows in the upperpart of Coire nam Beitheach at altitudes above 900m. All the populations are on the base-rich andesiteoutcrops, either in crevices and on small ledges or in the screes immediately below. S. cernua grows inseveral places on the cliffs between Collie’s Pinnacle and Hourglass Gully, at the top of Hourglass Gully, onthe base-rich rock outcrop below the west top of Bidean and beneath an overhang on the rocky spur whichdivides this gully from the next one to the west. The characteristic habitat is on damp rock among mossunderneath an overhang. We re-found the plant in what appears to be all of its known stations, togetherwith a few more. In the summer of 2002 there were at least 250 plants of S. cernua in seven discretepopulations. No flowering plants were seen but almost all, even the small individuals, bore the red bulbils.One known site was not re-visited because to do so would have necessitated crossing a new rock-fall andthen a steep, wet, smooth slab of rock; both on the lip of a precipice. The estimate of population size is aconservative one. The species grows in clumps and patches and it can be hard to tell where one individualends and another begins, especially when you are standing up to your ankles in wet scree on a 60-degreeslope and holding on to slippery wet rock ledges with your fingernails!

Saxifraga rivularis also grows in upper Coire nam Beitheach but descends to about 825m on the corriefloor. It is a small plant with clumps of pale green lobed leaves and inconspicuous white flowers (Photograph 18).Like S. cernua, it grows in crevices on the andesite rock outcrops and on the screes beneath. It has, however,a wider range of habitats than S. cernua. It grows along the sides of the streams and rills in the corrie,among Pohlia ludwigii and other snow-bed species on the steep screes, in Philonotis-Saxifraga springs M32and Pohlia wahlenbergii var. glacialis springs M33, and among boulders in the gullies. The populations arealongside the rills and among boulders on the floor of the corrie, on the cliff-ledges between Collie’s Pinnacleand Hourglass Gully, on the steep upper walls of the corrie and in the scree-filled gullies. S. rivularis is moreplentiful than S. cernua and there are hundreds of plants. Most of these were in flower.

Saxifraga nivalis is yet another rarity of upper Coire nam Beitheach. The stout rosettes of succulent purple-backed leaves grow on small rock ledges and among scree above about 900m, and indeed the habitathere is very similar to that of S. cernua. S. nivalis grows on the cliffs between Collie’s Pinnacle and HourglassGully, on the base-rich rock outcrop below the west top of Bidean, among boulders in the wide gully belowthis, beneath an overhang on the rocky spur which divides this gully from the next one to the west, and onthe eastern cliffs of Stob Coire nam Beith. Many of the ledges are inaccessible but there are at least ahundred plants in the corrie. About half those seen were flowering.

Sibbaldia procumbens is a distinctive plant with trailing dark red stems and silvery-green, 3-lobed, toothedleaves. It is one of the more widespread of the rare species in the study area. It grows in places where snow

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lies late in spring and where the ground is churned up by the plucking movements of the packed snow asit slips down the steep slopes. S. procumbens grows in Coire Eilde, Coire Gabhail, Coire nan Lochan, inboth branches of Coire nam Beitheach, in Coire Corcaig on the northern side of Aonach Eagach and inCoire Clioche Finne on the south-eastern side of Buachaille Etive Mór. The total population must numberthousands of plants.

There is no doubt at all that the site is of immense value for its rare montane vascular plant species. It isconsidered (Slack & Farrell, unpublished notes) to have the largest populations in Britain of Saxifraga cernuaand S. rivularis. There are hundreds of plants of both. There are also good populations of Cerastiumarcticum, Poa alpina, Athyrium distentifolium, Cystopteris montana, Deschampsia cespitosa ssp. alpina,Dryas octopetala, Salix myrsinites and Sibbaldia procumbens, and smaller ones of Saxifraga nivalis,Polystichum lonchitis, Juncus biglumis, Carex saxatilis, C. capillaris, Cerastium alpinum, Draba norvegicaand Poa glauca. Most of these rare species grow in types of vegetation which are themselves uncommonand are Qualifying EC Annex I Habitats for the proposed Special Area for Conservation. They all grow inLuzula-Geum tall-herb-vegetation U17. Saxifraga cernua, S. rivularis, Cerastium arcticum, Athyrium distentifolium,Cystopteris montana, Deschampsia cespitosa ssp. alpina and Sibbaldia procumbens also grow in thevarious snow-bed communities. Dryas octopetala and Salix myrsinites grow in Dryas-Silene heath CG14and Salix-Luzula scrub W20. Most of the remaining Pinus sylvestris in the study area grows along the banksof the River Etive. There are many scattered trees and a small patch of pine woodland W18.

Carex lachenalii, C. atrata, Juncus castaneus, Potentilla crantzii and Veronica alpina were not re-foundduring this survey. Carex lachenalii was previously recorded on steep broken ground below the main terracein upper Coire nam Beitheach. This is a dangerous area, specially when wet and slippery, and was notthoroughly searched during our survey. However, the species has not been re-found since its first discovery– even by the original recorder. Carex atrata was recorded in 1986 by a party from the BSBI on the cragson the north-eastern side of Stob Coire nam Beithe but has not been seen there since. Only two plants ofJuncus castaneus have been seen in recent years (Halcrow 1996). The plant is inconspicuous when notflowering or when the flowers are grazed and may still occur more widely. Veronica alpina was recordedin 1987 but no location was given and it has not been seen since then. 12 plants of Potentilla crantzii werefound in 1995 on the upper north-eastern slope of Meall Mór (Halcrow 1996). If this was in the ungrazedexclosure it would be hard to see it now in the tall vegetation, unless it was flowering.

One of the mysteries of Saxifraga cernua on Bidean nam Bian is the variation in what the numerous visitingbotanists have found. Many of the records are drawn onto sketched maps which are hard to relate tolandmarks in the field. The best records were those left by Halcrow (1996), who marked the populationsonto photographs. Our findings correlate well with those of Halcrow (1996) – we re-found all her sitestogether with two further small populations. The sizes of the populations seem to vary over the years. Forexample, in 1977 Paul Harrold found 12 plants at the foot of Collie’s Pinnacle. In 1985 Lynne Farrell,Andrew Bachell and Alf Slack found 6 plants here. In 2002 there were 14 plants. In 1974, a party led byHugh McAllister found 24 plants under the overhang where we found at least 50 plants in 2002. PaulHarrold in 1977 found about 60 plants on the outcrop below the west top of Bidean; in 2002 there wereat least 125 plants here.

Saxifraga cernua is a perennial herb which over-winters as bulbils, putting out shoots each spring (Clapham,Tutin & Moore 1987). The bulbils can start to develop into new plants while still attached to the parent, but

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usually they detach from the stem and roll a short distance before they start to grow. They can also bedispersed more widely by wind or melting snow (Lusby & Wright 1996). Some of the plants on Bidean aregrowing upside-down in crevices below the roofs of overhangs and it is hard to imagine how the bulbilsever reach such spots.

5.3.2.2 Oceanic bryophyte species

The site is of great importance for western, oceanic bryophytes. We have found 43 oceanic bryophytespecies here:

Mosses: Andreaea alpina, Breutelia chrysocoma, Campylopus atrovirens, C. setifolius,Glyphomitrium daviesii, Hymenostylium insigne, Hyocomium armoricum, Isothecium holtii, I. myosuroides var. brachythecioides, Molendoa warburgii, Oedipodium griffithianum,Paraleptodontium recurvifolium, Ptychomitrium polyphyllum, Rhabdoweisia crenulata,Sphagnum strictum, Trichostomum hibernicum, Ulota phyllantha.

Liverworts: Anastrophyllum donnianum, Aphanolejeunea microscopica, Bazzania pearsonii,Colura calyptrifolia, Douinia ovata, Frullania teneriffae, Gymnomitrion crenulatum,Harpalejeunea molleri, Herbertus aduncus, H. stramineus, Lejeunea patens, Lepidoziapearsonii, Leptoscyphus cuneifolius, Mastigophora woodsii, Metzgeria leptoneura,Plagiochila carringtonii, P. exigua, P. killarniensis, P. punctata, P. spinulosa, Pleuroziapurpurea, Radula aquilegia, Saccogyna viticulosa, Scapania gracilis, S. nimbosa, S. ornithopodioides.

This is a very rich assemblage of oceanic bryophytes. There are particularly high concentrations of theseplants in the rocky woods along the Allt Coire Gabhail and along the nearby River Coe, in the woods onthe steep north-facing slopes of the Three Sisters, among rocky snow-beds in the higher corries of Bideanand in rocky Vaccinium heath in Coire na Tulaich on Buachaille Etive Mór. The woodlands are among therichest in Great Britain – and thereby among the richest in Europe – for oceanic bryophytes.

5.3.2.3 Uncommon bryophyte species

Although we were not asked specifically to look at uncommon bryophytes, many such species do occur hereand we recorded a number of them in the course of our work. The Nationally Rare moss Brachytheciumglaciale occurs in snow-beds and among boulders in upper Coire nam Beitheach, as do the NationallyScarce Aulacomnium turgidum, Kiaeria starkei, K. falcata, K. blyttii, Philonotis seriata, Pohlia ludwigii,Marsupella alpina and the oceanic liverworts Anastrophyllum donnianum, Bazzania pearsonii andPlagiochila carringtonii. Most of these species occur also in the other high corries. The Nationally Scarceoceanic moss Dicranodontium uncinatum and the liverwort Herbertus aduncus grow in liverwort-rich Calluna-Vaccinium-Sphagnum heath H21b and in the bryophyte-rich woodlands. The Nationally Scarce liverwortsColura calyptrifolia, Sphenolobopsis pearsonii, Lejeunea patens and Microlejeunea ulicina grow in thewoods. The Nationally Scarce mosses Bartramia hallerana and Rhabdoweisia crenulata grow on rockledges.

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Vascular plants

Alchemilla alpinaAntennaria dioicaArabis petraea *Asplenium virideAthyrium distentifolium *Botrychium lunariaCarex bigelowiiCarex capillaris *Carex paucifloraCarex saxatilis *Carum verticillatumCerastium alpinum *Cerastium arcticum *Cirsium heterophyllumCochlearia pyrenaicaCornus suecicaCrepis paludosaCryptogramma crispaCystopteris montana *Diphasiastrum alpinumDraba norvegica *Drosera longifoliaDryas octopetala *Dryopteris oreadesEmpetrum nigrum hermaphroditumEpilobium alsinifoliumEpilobium anagallidifoliumEriophorum latifoliumGalium borealeGalium sterneriGeranium sylvaticumGeum rivaleGnaphalium supinumHuperzia selagoHymenophyllum wilsoniiJuncus biglumis *Juncus trifidusJuncus triglumisJuniperus communisKoeleria macranthaListera cordataLuzula spicataLycopodium annotinumLycopodium clavatumOxyria digynaParnassia palustrisPersicaria viviparaPinus sylvestris *Poa alpina *Poa glauca *Polystichum lonchitis

Populus tremulaPseudorchis albidaPyrola minorRhynchospora albaRubus saxatilisSalix herbaceaSalix myrsinites *Salix phylicifoliaSaussurea alpinaSaxifraga aizoidesSaxifraga cernua **Saxifraga hypnoidesSaxifraga nivalis *Saxifraga oppositifoliaSaxifraga rivularis **Saxifraga stellarisSedum roseaSibbaldia procumbens *Silene acaulisSilene unifloraThalictrum alpinumTofieldia pusillaTrollius europaeusVaccinium uliginosum

Mosses

Andreaea alpinaAntitrichia curtipendulaAulacomnium turgidum *Bartramia hallerana *Brachythecium glaciale **Calliergon trifarium *Campylopus setifolius *Conostomum tetragonum *Dicranodontium uncinatum *Distichium capillaceumDryptodon patensGlyphomitrium daviesii *Grimmia funalisHylocomium umbratumHymenostylium insigne **Hypnum callichroumIsothecium holtiiIsothecium myosuroides

brachythecioidesKiaeria blyttii *Kiaeria falcata *Kiaeria starkei *Molendoa warburgiiOedipodium griffithianum *Orthothecium rufescens *Paraleptodontium recurvifolium *

Philonotis seriata *Plagiobryum zieriPohlia ludwigii *Pohlia wahlenbergii glacialis*Ptilium crista-castrensisRhabdoweisia crenulata *Sphagnum fuscumSphagnum girgensohniSphagnum magellanicumSphagnum strictumSplachnum ampullaceumTrichostomum hibernicum *

Liverwor ts

Anastrepta orcadensisAnastrophyllum donnianum *Anthelia juratzkana *Aphanolejeunea microscopicaBarbilophozia atlantica *Bazzania pearsonii *Bazzania tricrenataCololejeunea calcareaColura calyptrifoliaDouinia ovataFrullania teneriffaeGymnomitrion concinnatumGymnomitrion crenulatumGymnomitrion obtusumHarpalejeunea molleriHerbertus aduncusHerbertus stramineusKurzia trichocladosLepidozia pearsoniiLeptoscyphus cuneifolius *Marsupella alpina *Marsupella sphacelata *Mastigophora woodsii *Metzgeria leptoneuraMoerckia hibernica *Nardia compressaPlagiochila carringtonii *Plagiochila exiguaPlagiochila killarniensisPlagiochila punctataPleurozia purpureaPorella arboris-vitaeRadula aquilegiaRiccardia palmataScapania nimbosa *Scapania ornithopodioides *Sphenolobopsis pearsonii *Tritomaria exsectiformis

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Table 4 Plant species recorded in this sur vey at Glen Coe in 2002 which are uncommon at

least in most par ts of Great Bri tain

Nationally rare and nationally scarce vascular plants recorded here but not seen in this survey: Carex atrata *, C. lachenalii **, Juncus castaneus *, Potentilla crantzii * and Veronica alpina *. Additional uncommon bryophytes,found by British Bryological Society, include the mosses Mnium thomsonii *, Pseudoleskea patens * andIsopterygiopsis muelleriana *, and the liverworts Diplophyllum taxifolium *, Marsupella boeckii * and Moerkia blyttii *

Note: * = nationally scarce (recorded in 16–100 10 x 10km squares in Great Britain since 1950)** = nationally rare (recorded in <16 10 x 10km squares in Great Britain since 1950)

Table 5 Locations where nationally rare and nationally scarce vascular plant species were

found in this sur vey at Glen Coe in 2002

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Species Grid ref Target Habitat Population Other notes(all NN) note no.

Athyrium 1447 46 fern/bryophyte several tens of plants no signs of grazing seendistentifolium 5457 snow-bed vegetation scattered through

large area

1487 53 fern/bryophyte several tens of plants fronds too young to note5527 snow-bed vegetation scattered through any effects of grazing

large area

1473 50 fern/bryophyte several tens of plants fronds too young to note5412 snow-bed vegetation scattered through any effects of grazing

large area

Carex capillaris 1112 5 species-rich grassland c.20 plants seen flowering well5706 (U4F/CG10) around here

1111 7 Dryas-Silene heath a few plants seen; flowering well5662 (CG14) not counted because

they were on ratherinaccessible crags

1091 9 Salix myrsinites a few plants seen; flowering well5638 scrub (W20) not counted because

they were oninaccessible crags

1105 10 species-rich grassland 6 plants seen; flowering well5610 (U4F/CG10)) probably more

1114 11 species-rich grassland 10 plants seen; flowering well5607 (U4F/CG10) probably more; seen

mostly just outside(downslope of)fenced area

Carex saxatilis 1441 49 Carex saxatilis mire over 100 rosettes no flowers seen5472 (M12) seen within an area

of about 20 squaremetres

Cerastium 1397 29 wet tall-herb 10 plants seen not floweringalpinum 5458 vegetation U17a

1429 44 wet tall-herb a few plants seen not flowering5454 vegetation U17a

Cerastium 141 545– includes wet tall-herb abundant; thousands flowering wellarcticum 145 547 29 and vegetation U17a, of plants in upper

44 snow-beds (U11, U12, Coire nam BeitheachU14 and Pohlialudwigii snow-beds),on barish gravel andscree, and amongboulders

Table 5 (continued)


Cystopteris 1412 34 lower ledges of cliffs 3 clumps near base lush, healthy, well grownmontana 5441 of cliff plants, with spores; no

signs of grazing seen

1410 35 lower ledges of cliffs dense clump about lush, healthy, well grown5444 60cm wide and 2m plants, withspores; no

long signs of grazing seen

1410 36 at base of cliff, where a few plants lush, healthy, well grown5449 cliff base adjoins plants, with spores; no

boulder-field downslope signs of grazing seen

1429 44 wet tall-herb plentiful in clumps; lush, healthy, well grown5454 vegetation U17a also populations of plants, with spores; no

more scattered fronds signs of grazing seen

Draba 1420 40 on ledges beneath 9 rosettes not floweringnorvegica 5443 overhang at foot of

cliff

Dryas 1044 2 Dryas-Silene heath scattered patches floweringoctopetala 5678 CG14

1085 4 Dryas-Silene heath plentiful on steep, Because of difficulty of5638 to CG14, and rocky banks but also physical access on these1100 scattered among scattered further east slopes, combined with5594 more open downslope; certainly difficulty in pinpointing

vegetation on crags more than the particular locations on a28 plants or clumps map we found detailedrecorded by Halcrow mapping and counting of(1996) Dryas within this area to

be impracticable

1111 7 Dryas-Silene heath scattered patches flowering5662 CG14

1114 8 growing under trees scattered patches flowering5652 on wooded crag

1091 9 Salix myrsinites scrub scattered patches flowering5638 W20

1114 11 on rocky banks among several plants some plants had flowered5607 grassland; some also in this year

grassland below crags

Juncus biglumis 1410 33 on flushed ledges two small but healthy flowering well5437 below ‘The Terrace’ populations, each of

about 25–30 plants

Pinus sylvestris 1968 83 in ravine one tree seen here5173

1972 84 in ravine one tree seen here5180

2055 85 wooded ravine of several trees (locally5125 River Etive dominant in woodland

canopy) on steep slopes,with birch. Some youngregeneration in fencedexlosure just to N

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Table 5 (continued)


Poa alpina 141545– includes tall-herb ledge quite common viviparous inflorescences 145 547 29, 35, vegetation on the throughout this area: common

37, 38 cliffs of upper Coire hundreds of plantsand 44 nam Beitheach in total

Poa glauca 1412 34 scattered thinly on fewer than 20 plants flowering5441 to cliff ledges seen14205443

1472 43 scattered on cliff just W fewer than 20 plants flowering5450 of dark streaks where seen

water flows over rock

1429 44 wet tall-herb vegetation fewer than 20 plants flowering5454 U17a on cliff ledges seen

Salix myrsinites 1075 4 steep crags with plentiful; at least a Larger bushes to c.1m5650 to herb-rich vegetation few tens of bushes high x 2m across;1100 scattered on rocky banks;5595 thickening up in places

into a few square metresof W20 scrub; manysmaller (younger?) plantsup to c.50cm tall.Willows unbrowsed for2 years but those in moreaccessible places havecontorted lower stems –probably browsed in thepast. Some smaller plantsnow quite obscured bytall ungrazed grasses andherbs. Difficulty ofphysical access andpinpointing locations onmap makes detailedcounting and mappingimpracticable. Halcrow’s(1996) figure of 40mature plants is aconservative estimate:there are many moresmaller individuals. Noevidence to suggest thatpopulation size haschanged since 1995.Not all willows floweringbut those that were inJune 2002 were allfemales. (Halcrow (1996),reported no flowers insummer 1995).

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Table 5 (continued)


Saxifraga 1411 30 lower part of NE- habitat too steep, most plants with bulbils;cernua 5429 facing rock face on unstable and no flowers seen

SW side of gully dangerous to getbelow summit ridge close enough to

count the plants

1407 31 Luzula-Geum at least 125 spikes most plants with bulbils;5430 vegetation U17a on and vegetative no flowers seen

mossy ledges and in clumpsgravelly soil on rockoutcrop at head oflong gully just belowwest top of Bidean

1410 32 western cliff at foot two spikes most plants with bulbils;5433 of Hourglass gully no flowers seen

1412 34 along base of cliffs at least 30 spikes most plants with bulbils;5441 in ‘The Terrace’ and vegetative clumps no flowers seen

1410 36 beneath overhang ten clumps most plants with bulbils;5449 W of long gully no flowers seen

1420 40 beneath a big at least 50 plants most plants with bulbils;5443 overhang at the no flowers seen

corner of the cliff

1435 42 on east side of base at least 14 plants most plants with bulbils;5444 of Collie’s Pinnacle no flowers seen

Saxifraga 1397 29 wet tall herb several tens of plants most plants flowering;rivularis 5458 vegetation (U17a)

1407 31 wet tall herb scattered some plants flowering5430 vegetation (U17a)

and on gravelly soil

1410 32 wet gravel and common here some plants flowering5433 Pohlia snow-beds/

springs (incl. M33)on scree at foot ofHourglass gully andacross to rock outcropat top of long gully

1410 36 beneath an overhang scattered rosettes some plants flowering5449

1416 37 along base of a fringe of plants flowering5443 shallow overhang just along base of

W of conspicuous overhangvertical crack knownas The Gash

1418 39 scattered among a few plants all plants flowering5453 boulders in the long

gully

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Table 5 (continued)


1440 48 in Philonotis-Saxifraga plentiful many plants flowering5465 spring M32 in the

bed of the streamand among bouldersbeside the stream

Saxifraga 1397 29 wet Luzula-Geum a few tens of plants some plants floweringnivalis 5458 tall-herb vegetation

U17a on cliffs

1407 31 wet Luzula-Geum scattered plants no flowers seen5430 tall-herb vegetation

U17a and mossyledges on cliffs

1410 36 beneath an overhang scattered rosettes no flowers seen5449

1416 37 scattered on small several tens of plants no flowers seen5443 ledges, to above

head-height

1418 39 scattered among one plant seen flowering5453 boulders in the long

gully

1420 40 on lower ledges of 13 rosettes no flowers seen5443 ‘The Terrace’

Sibbaldia 133 582– too summit and north side thousands of plants many plants floweringprocumbens 162 583 common of Aonach Eagach scattered through

here for ridge large areaseparatetargetnotes

135 554– too top and NE side of thousands of plants many plants flowering160 533 common main Bidean ridge scattered through

here for large areaseparatetargetnotes

2120 87 species-rich grassland several plants plants are low-grown;5377 (CG11) no flowers seen

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5.4 Summar y of the total botanical interest

The Glen Coe survey area has a great diversity of vegetation and habitat types including several which areuncommon in Britain or Europe, and a rich flora including several rare species. These combine to make thesite one of the most botanically interesting places in the British uplands. The most notable habitats here are:

● Snow-beds (U11, U12, U14, U18 and Pohlia ludwigii types) on the north side of the main Bideanridge. Diverse, extensive and containing several nationally uncommon species, these are some of themost impressive examples of snow-bed vegetation in the west Highlands. They are nationally important.

● Basic igneous crags and associated tall herb vegetation (U15 and U17) and flushes (M11 and M12)on the north side of Bidean. Nationally important populations of some rare montane plants, includingwhat may be the largest populations of Saxifraga cernua and S. rivularis in Britain.

● Bryophyte-rich heaths (H20c and H21b) on the north-facing slopes on Bidean, the Three Sisters andBuachaille Etive Mór. These heaths contain excellent, internationally important assemblages of northern,sub-montane and montane oceanic liverworts. This is around the southern limit of the very rich northernoceanic liverwort assemblages which occur from here northwards through the western Highlands.

● Rocky, bryophyte-rich woodland (W17) on the very steep N slopes on the NE end of Gearr Aonach,and stretching from the Allt Coire Gabhail boulder-field and ravine down to the confluence with the RiverCoe (‘Meeting of Three Waters’) and further downstream along the River Coe. Taken together, these twoareas of woodland form one of the richest sites for oceanic bryophytes in Britain.

● Excellent, internationally important examples of western, oceanic blanket bog (M17 and M18 with M1and M3) in two areas in the NE of the survey area – NE of Buachaille Etive Beag and Buachaille EtiveMór. This is ‘top quality’ bog because of its floristic diversity, microtopography, good Sphagnum carpetsand apparent lack of disturbance in the past.

● Limestone area on the steep NE and N slopes of Meall Mór: grassland (CG10, CG11, CG12, U4F,U5c and M25c), flushes (M10 and M11), tall herb vegetation (U15 and U17), woodland (W9), Dryasheath (CG14) and Salix myrsinites scrub (W20). A diverse mixture of vegetation types, with a very richflora of calcicole species, some of which are nationally uncommon. Rather like a north-western outlier ofBreadalbane-type calcicolous vegetation. Nationally important.

Among the various ecological divisions in the Scottish Highlands, two are particularly important with regardto vegetation. One is between the east and the west. The west has a more oceanic climate and the east amore continental one. There are corresponding differences in the typical vegetation: most notably the greaterabundance and diversity of bryophytes in the west. Glen Coe is on the western side of this division, withvery good bryophyte-rich heaths and woods, and fine examples of wet blanket-bog. The other ecologicaldivision is between areas with predominantly acid rocks and those with a greater extent of base-rich rocks.The Glen Coe hills have both acid and basic rocks: there are very good examples of acidophilous woods,heaths, bogs and snow-beds, and some of the best examples of basiphilous vegetation outside theBreadalbanes.

Other parts of the western Highlands have mixtures of vegetation types similar to that on the Glen Coe hills.On the Mamore Forest, to the north across Loch Leven, there is a similar range of dwarf-shrub heathsincluding Dryas octopetala heath CG14 and liverwort-rich Calluna-Vaccinium-Sphagnum heaths H21b.There are herb-rich Festuca-Agrostis-Thymus CG10 and Nardus U5c grasslands, Rhytidiadelphus loreus

snow-beds U13b, Trichophorum-Eriophorum blanket bog M17 and birch woodland W11 and W17. Thesetypes of vegetation also occur on Ben Nevis. Ben Nevis is higher than Bidean nam Bian and has a largeplateau where snow collects; snow-bed vegetation is more extensive there and on the adjacent peaks thanit is on Bidean. Saxifraga cernua and S. rivularis also grow on Ben Nevis, although they are not knownfrom any other hills in this part of the country. The hills around Glens Affric and Cannich also have muchsnow-bed vegetation, including large patches of Rhytidiadelphus loreus snow-bed U13b andCryptogramma-Athyrium snow-bed U18. They have tall-herb ledge vegetation U16 and U17, montanewillow scrub W20 on cliffs, pine woodland W18, western Trichophorum-Eriophorum blanket bogs W17and bryophyte-rich heaths. They are as far west as Glen Coe but are further from the sea. Though there areoceanic elements in the flora and vegetation, there is not such a rich flora of oceanic bryophytes, and manyof those that do occur grow in the montane heaths and snow-beds where they are protected from frost by alayer of snow in winter. There are also Calluna-Cladonia heaths H13, Vaccinium-Cladonia heaths H19,Carex-Polytrichum sedge heaths U8 and Juncus trifidus heaths U9: all eastern, boreal types of montanevegetation. There are hills further west, such as Ladhar Bheinn in Knoydart, which are almost as high asBidean and have similarly rich assemblages of oceanic plants and vegetation (e.g. Averis 2001). However,although the precipitation is still very great, these more western hills have less snow-bed vegetation, none ofthe eastern vegetation types such as Calluna-Cladonia heath H13 and Carex-Polytrichum heath U8, andfewer montane species than the Glen Coe hills.

The hill slopes between Sgorr nam Fiannaidh and the Pap of Glen Coe, outwith the study area to the west,are made of Dalradian quartzite. Although this ground is adjacent to the study area, its flora and vegetationare quite different. There is much more heather and very little grassland. There is Calluna-Vaccinium-Sphagnum damp heath H21, Calluna-Eriophorum blanket bog M19 with much Rubus chamaemorus, and on the upper slopes large patches of Calluna-Racomitrium heath H14 on gravelly, windswept ground.The montane species Loiseleurea procumbens is common here.

The Polytrichum-Kiaeria and Salix-Racomitrium snow-beds U11 and U12, the Alchemilla-Sibbaldia snow-bedU14 and the Pohlia ludwigii snow-bed occur in intricate and quite bewildering mosaics in the upper corriesof the study area. These communities have many plants in common, and the distribution of individual speciesseems to be quite random, though there may be subtle differences of shelter, shade, wetness of the soil andduration of snow-lie on a very small scale. Much of the vegetation in the upper corries does not fit neatlyinto an NVC category. Though the individual types can be recognised here and there, all possibleintermediates also occur and, in fact, cover more of the ground. In the west Highlands, only Ben Nevis and the adjacent hills are known to have more extensive and better-developed snow-beds than those in Glen Coe. These two ranges of hills, because of their great height and the high precipitation they receive,have the best examples of these types of vegetation outwith the eastern and central Highlands. On both sitesthese acid snow-beds and tall-herb vegetation on base rich ledges occur in unusually close proximity.

Another interesting element in the vegetation of both Bidean nam Bian and the Ben Nevis range isRacomitrium heath in which the dominant species is R. ericoides rather than R. lanuginosum. We found thistype of vegetation – identical to the normal Carex-Racomitrium heath except for the dominant species – onthe summit of Stob Coire nan Lochan on gravel and shattered stone. Similar heath was discovered byMcVean & Ratcliffe (1962) on the summit of Aonach Mór east of Ben Nevis. On that hill it is (or was) moreextensive, covering some tens of square metres where wind-blown sand from nearby eroded surfaces wascontinually being added to the vegetation. R. ericoides can tolerate repeated burying by sand and shingle.

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It is exactly this which enables it to colonise riverside shingle and the edges of tracks. On Stob Coire nanLochan there is no obvious sand, and no obvious reason why R. ericoides should take the place of R. lanuginosum. Its occurrence here is interesting, though, because R. ericoides is the dominant species insome moss heaths in Iceland and Jan Mayen Island (McVean & Ratcliffe 1962) and over small areas in thehills of the Faroe Islands (Hobbs & Averis 1991).

The Salix myrsinites scrub W20 on Meall Mór is very interesting. Pure scrub of S. myrsinites is rare. Thespecies is more common as a component of more mixed willow scrub including such species as S. lapponumand S. arbuscula. Pure S. myrsinites scrub in Britain seems to be a community of limestone. Elsewhere itoccurs at Inchnadamph in Sutherland and at Rassal in Wester Ross. In these two places it occurs at lowaltitudes on dry limestone, without the rich underlayer of tall herbs. The scrub on Meall Mór, with its tallswards of herbs on dripping limestone ledges, is more like the mixed willow scrub in the Breadalbane hillsthan it is like the low-altitude scrub in the far north west. Dryas octopetala is one of the few species commonto both. D. octopetala is a plant of open ground and it is rare to see it growing under trees and shrubs, as it does here. It also grows under birch trees on the limestone in Strath Suardal on Skye (Averis & Averis2000), and under hazel in the Burren in western Ireland (Mabey & Evans 1980).

The birch woods in Glen Coe have a rich flora of oceanic bryophytes. There are woods with a similar arrayof species in Glen Nevis and Glen Creran (Averis 1991) and in Knoydart (Averis 2001). These woods arenotable because northern liverworts such as Scapania ornithopodioides and Plagiochila carringtonii,generally plants of moderate to high altitudes, grow in them. It is also notable that more southern species,such as Bazzania trilobata and Saccogyna viticulosa, grow in mixed mats with the northern and montaneplants. This tends to happen most often where the climate is very oceanic. Species of contrastingphytogeographical distributions are able to grow together because the winters are not too cold for thesouthern species and the summers are not too warm for the northern species. Glen Coe is an especiallygood place for this. Another example is the lowland, southern species Silene dioica. It grows in the upperpart of Coire nam Beitheach, at over 900m, in the company of montane plants such as S. acaulis, Saxifragacernua, S. rivularis, S. nivalis, Poa alpina and Cystopteris montana. Yet another example is the normallymontane Empetrum nigrum ssp. hermaphroditum. This species occurs here not only at high altitudes on themountains but also low down in the wooded boulder-field of the Allt Coire Gabhail.

Several of the oceanic bryophytes which grow so prolifically in the woods also occur on the open slopes.Herbertus aduncus, Pleurozia purpurea, Scapania gracilis, S. ornithopodioides, Plagiochila spinulosa, P. carringtonii, Mastigophora woodsii, Mylia taylorii, Bazzania tricrenata, B. pearsonii, Anastrepta orcadensisand Lepidozia pearsonii form the hepatic mat community which makes up the characteristic bryophyte layerof liverwort-rich Calluna-Vaccinium-Sphagnum heath H21b and, with the addition of the more montaneScapania nuimbosa and Anastrophyllum donnianum, of liverwort-rich Vaccinium-Racomitrium heath H20c.More unusually, many of these species grow in their characteristic mixed mats in damp Festuca-Agrostis-Galium grassland U4d, Trichophorum-Erica wet heath M15, Oreopteris limbosperma and Pteridiumaquilinum fern communities U19 and U20, and in Crypyogramma-Athyrium snow-beds U18. They occur fromthe foot of the main glen at about 100m to over 900m in the high corries. Only in the far west Highlands arethese species so widely distributed through so many types of vegetation and over such an altitudinal range.

The hillsides in Glen Coe are green and grazed, with little heather except on the cliffs. This pattern of vegetation is common in this part of Scotland: for example on the hills in Glen Etive, the Black Mount,

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Beinn Dòrain, Beinn an Dòthaidh, Beinn Cruachan, Ben Lui and the western Breadalbanes. This is almostcertainly because of the great intensity of grazing where the rocks are rich and the herbage nutritious. All these hills have similar vegetation on their long green slopes: mixtures of sub-montane Vaccinium-Deschampsia heath H18, Nardus-Galium U5 and Festuca-Agrostis-Galium U4 grasslands, Oreopterislimbosperma fern beds U19 and bracken U20.

One of the more interesting types of grassland which we found on the slopes of Meall Mór is the herb-richform of Festuca-Agrostis-Galium grassland U4F. This is a rare grassland community which seems to occuronly on steep slopes which are flushed with moderately base-rich water and which are not grazed too hard.We have found it on Mull (Averis & Averis 1995b, 1999b), Beinn Eighe (Averis & Averis 1998) and onBen Lui (Averis & Averis 1999a). We have seen more of it on Ben Lui than anywhere else: it has becomevery extensive on the north-facing slopes since the sheep flock was sold. It was also good to see examplesof herb-rich Juncus squarrosus heath U6R and Deschampsia cespitosa grassland U13ahr. Both occur in theBreadalbane hills and over richer rocks in the north-west Highlands, but neither is described in the NVC.

The Glen Coe hills are not far from Ben Lui and the western Breadalbanes. Bidean nam Bian is only about15km west of Ben Lui, but its vegetation and flora are considerably more oceanic. Many of the oceanicbryophytes which are so plentiful in Glen Coe do not grow on Ben Lui at all, showing the very steep gradientof oceanicity inland from the west coast. Ben Lui, like Glen Coe, is within the range of native pine woodlandand there are some fine surviving examples of Pinus-Hylocomium woodland W18 in lower Glen Cononish.There is Trichophorum-Eriophorum bog M17 there too, but it is much more degraded by burning and heavygrazing than that in Glen Coe. The oceanic snow-bed communities Rhytidiadelphus loreus snow-bed U13band Cryptogramma-Athyrium snow-bed U18 are much less extensive on Ben Lui and the adjacent hills thanthey are in Glen Coe, but there are more lichen-rich heaths, and more Calluna-Eriophorum blanket miresM19 on Ben Lui. Carex saxatilis mires M12 are common on Ben Lui as they are throughout theBreadalbanes. There seems no reason why this type of vegetation should be so scarce on the Glen Coehills. The tall-herb ledge vegetation on Ben Lui, as elsewhere in the Breadalbanes, is tall, lush and luxuriant,forming vertical gardens of grasses, ferns and flowering herbs. In the Glen Coe hills, only the limestone cragsof Meall Mór have anything like the typical Breadalbane tall-herb ledges and herb-rich woodland and scrubon crags. Alhough there is much Luzula-Geum vegetation on Bidean nam Bian and its satellite peaks, mostof it, and certainly the patches with the more interesting rare species, belongs to the Alchemilla-Bryumsub-community U17a. U17a has a more open, patchy and shorter sward than the Geranium sub-communityU17b which is so noticeable on Ben Lui and on the other Breadalbane hills with basic rocks.

5.5 Relationships between vegetation and environment

5.5.1 Geology

There are great geological contrasts over the study area, from basic limestone to acid granite. However,with a few exceptions the plants and plant communities fall into two groups: those on acid substrata andthose on basic substrata. We found three vegetation types – Dryas-Silene heath CG14, Salix myrsinitesscrub W20 and Saxifraga-Alchemilla banks U15 – only on the limestone. The other basiphilous communitiesoccur both on the limestone and on the basic volcanic rocks.

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5.5.2 Climate

The study area has a cool and oceanic climate, especially on the high tops where winds are stronger, mistand rain more frequent and temperatures lower. There is much variation on a small scale, with localrainshadow effects resulting in almost bewildering changes of weather over the length of the study area.Almost whichever way the clouds come, they will be forced up over high ridges. It is not uncommon for thesun to be shining on Meall Mór at the same time that torrential rain is falling at the top of the glen, or forSgorr nam Fiannaidh and the Aonach Eagach to be clear and dry while the other side of the glen iswrapped in cloud and rain.

Many of the vegetation types which occur are distinctively western ones such as mossy Quercus-Betula-Dicranum woodland W17a, Calluna-Erica dry heath H10, liverwort-rich forms of Calluna-Vaccinium heathH20c and Calluna-Vaccinium-Sphagnum heath H21b, Trichophorum-Eriophorum blanket mire M17a,Racomitrium-rich Trichophorum-Erica heath M15c, herb-rich Carex-Racomitrium heath U10b andRhytidiadelphus loreus snow-beds U13b. In contrast, there are only a few more eastern, boreal types ofvegetation, such as Calluna-Cladonia heath H13 and Carex-Polytrichum snow-bed U8.

An interesting effect of an oceanic climate is that cold-tolerant and thermophilous species and vegetationtypes can occur in close proximity. So Juncus acutiflorus mires M6d and M23a, Iris pseudacorus mires M28and Quercus-Betula-Oxalis woodland with ferns and honeysuckle W11a, all of which need a mild climateand have a south-western distribution in Scotland, grow within sight of late snow-bed vegetation U11, U12and U14, Calluna-Cladonia heath H13 and Carex-Polytrichum heath U8, all of which are more associatedwith the cold and more continental climate of the central and eastern Highlands. The summers are not toowarm for the northern and eastern types, nor the winters too cold for the southern types.

There is a great range of habitats from the woodlands on the lower slopes with luxuriant layers of tall herbsand ferns under the trees and a mild and temperate microclimate, to the moss heaths of the highest groundwhere the conditions are near-arctic. As is usual in the oceanic climate of the far west, the ecologicalgradients are extremely steep and it is possible to stand in snow-bed vegetation and look down on shelteredwoodlands only one or two kilometres below. In the high montane zone there is an enormous contrastbetween the exposed summits and ridges and the sheltered, shaded corries. Bryophytes predominate inboth, but the species and the communities they form are very different. The Carex-Racomitrium heaths U10and Carex-Polytrichum heaths U8 on the bare summit ridges are able to tolerate extremes of exposure frombitter winter cold to the scorching sun on clear days in summer, and are resistant to wind and ice. Thebryophytes of the snow-beds on the high sunless slopes of the corries are able to grow in an environmentwhere they may be covered by snow for up to eleven months of the year. Mosses and liverworts are ableto grow in these places because they need so little light for photosynthesis (Russel 1990) that they are ableto do so even under snow. Many of them contain chemicals which have antifungal and antibiotic properties(Averis 1994). This means that they do not succumb to moulds even though they grow in still, damp andshaded conditions.

The amount of snow and the time it lies can vary a great deal from year to year. In 2002, there was a lotof snow in the corries and on the summit ridges in early June, and the long gullies in upper Coire namBeitheach were almost filled with it (see photographs). There was still a narrow wreath of snow at the topof the corrie on 20 July, but by early August all had disappeared. However, in 1994 there was still muchsnow in the upper corries in late August, and by October there were fresh accummulations of new snow(Averis & Averis, personal observation).

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Vegetation/ basic basichabitat type (limest.) (volc.) acidic

W3 . . .W4 . . X

W7b X . .W7c . . .W9b X . .W11a . . X

W11b X X X

W17a X X X

W17b . . X

W17c . . X

W18d . . X

W20 X . .W24 . . .W25 . . .W25D . . .H10a . . X

H10b . . X

H10c . . X

H10d . . X

H12b . . X

H12c . . X

H13a . . X

H14 . . X

H18a X X X

H18b X X X

H20a . X X

H20b . X X

H20c . X X

H21a . . X

H21b . . X

H22a . . X

HX . . X

M1 . . X

M3 . . X


M4 . . X

M6a . . X

M6b . . X

M6c . . X

M6d X X X

M7 . . .M10a X X .M11a X X .M11b X X .M12 . X .M15a X X X

M15b X X X

M15c X X X

M15d . X X

M17a . . X

M17b . . X

M18a . . X

M18b . . X

M19a . . X

M23a X . .M25a X X X

M25b X X X

M25c X . .M28a . . .M31 . X X

M32a . . X

M32b X X .M33 . . .M37 . X .MX X . .MG9 . . X

CG10a X X X

CG10b X X .CG10c X . .

Table 6 Occurrence of vegetation types in relation to broad rock and soil type ( l imestone,

basic igneous rock and acidic rock)

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Table 6 (continued)

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CG10 shingle . X .CG11a X X X

CG12 X X .CG14 X . .U4a X X X

U4b . . .U4d . X X

U4e . X X

U4eA . X X

U4F X . .U5a X X X

U5b X X X

U5c X . .U5e X X X

U6a X X X

U6c X X X

U6d X X X

U6Cv . . .U6R . . .U7a . X X

U7b . X X

U7c . X X

U8 . . X

U10a . . X

U10b . . X

U10c . X .U11a . . X

U12a . . X

U12b . . X

U13a . X X

U13ahr . X .


U13b . . X

U14 . X X

U15 X X .U16a . . .U16b . . .U16c . . .U17a . X .U17b X X .U17d . X .U18 . . .U19 X X X

U20a X X X

U20c . . .U21 . . X

S9a . . X

S9b . . X

S10a . . X

OW + Menyanthes . . X

Je . . X

Rac.eric. shingle . . .Rac.eric. heath . . .Fell-field . . X

Garden . . .Acid Rock . . X

Basic Rock X X .Acid Scree . . X

Basic Scree . . .Boulder-field . . .Pohlia ludwigii . . .Shingle . . .Open Water . . X

Note: No ‘X’ in any column = we do not know what the bedrock type is at the precise location of this vegetation type.

5.6 Land-use and management

5.6.1 Histor y of vegetation, land-use and management

These hills have been vegetated from the time the last Pleistocene glaciers began to retreat about 10,000years ago. One of the first colonists was probably Dryas octopetala, and this species may have occurredcontinuously on Meall Mór from that day to this. Saxifraga cernua and S. rivularis are also plants whichgrow on wet gravel and rocks around the margins of glaciers, and these, too, along with the snow-bedbryophytes, may have been in continuous occupancy since late-glacial times.

Trees moved northwards as the climate became warmer, and woodland would have reached its greatestextent about 7000 to 8000 years ago (Birks 1988, Tipping 2003). Even at this time there was probablynot continuous, wall-to-wall tree cover. There is more likely to have been a mixture of tall dense forest ondeeper soils in sheltered spots, scrubby woodland on the more exposed slopes and much open groundwhere the soils were thin and rocky or peaty and wet. Such mixtures of woodland and scrub possiblyextended up to over 700m on sheltered slopes (Birks 1988), as birch woods still do today in isolatedfragments in Glen Nevis a few kilometres to the north. It is more likely that the general limit of closed, densewoodland in this part of Scotland would have been around 500m (Ward 1987 in Tipping 1997).

Most of the woodland on the more acid soils would have consisted of mixtures of birch and pine (Birks1988), probably with a shrub layer of rowan and holly and with a heathy ground flora similar to that whichpersists today under the birch woods in the main glen and in the pine woods in Glen Etive. Mosses,liverworts and lichens, including the choice oceanic species still recorded here, would have grown over therocky ground, clothing boulders in thick, richly-coloured mats and patches and festooning the trunks andbranches of the trees. On the richer soils there would have been pockets of ash and elm woodland, withan understorey of hazel and perhaps hawthorn, and a flowery herb-rich ground layer below. A similar typeof woodland clings today to the limestone cliffs of Meall Mór. It occurs in the woods of the Allt Coire Gabhailas well, and there are still a few large ash trees scattered on the slopes of the main glen. In the boggy glens,along the streams and on the floodplain of the River Coe there would have been wet woodland with alder,birch and willows and an underlayer of herbs and grasses.

There is still much woodland in the lower glen of the Allt Coire Gabhail and on the steep inaccessible facesof Gearr Aonach and Aonach Dubh, and there are scattered rowans up to about 700m on the cliffs. Thereare even a few dwarfed rowans on the ridges at over 900m, presumably the result of seeds dropped bybirds. Above the altitudinal limit of tall woodland the trees would have thinned out. There would have beenan open zone of stunted rowan, birch and willow, either with a heathy understorey or with ferns, Luzulasylvatica and tall herbs similar to that which occurs in western Norway today. There may also have beenareas of juniper scrub. Juniperus communis ssp. nana (Photograph 46) still grows on the summit ridges ofAonach Dubh, Gearr Aonach and, especially, Beinn Fhada as well as on the cliffs. It is sensitive to burningand may once have been much more common. On the upper slopes of Meall Mór there may once havebeen substantial areas of montane willow scrub, with tall herbs under a low tangled canopy of Salixmyrsinites. This, however, is by no means certain, as the palynological records do not show much evidencefor extensive montane scrub in most of Scotland (Tipping 1997).

The earliest large-scale clearance of woodland in this part of Scotland is believed to have taken place about400 years ago (Birks 1988). This is when the amount of tree pollen in peat cores drops to about 50% of

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its post-glacial maximum level. People have been living in these parts for much more than 400 years, andthere must have been smaller-scale clearings for pasturing animals and raising crops. It is possible that theoriginal pines were felled preferentially to use for buildings as they must have been the tallest, straightest andmost easily worked of the native trees. The place names support this. Many of the Gaelic names in the glenincorporate beith, birch, but there are none with ghuibhais, pine, nor with any other tree species. The FionnGhleann leading up between Bidean nam Bian and Meall Mór is the white glen, which suggests neithertrees nor heather.

At the time of the infamous massacre in 1692, there were about 150 people living in hamlets and groupsof cottages scattered on the floor of the glen. The survivors remained in the glen and soon the populationwas high once more. Then the sheep were introduced, and with them came the Clearances, and the landwas left desolate again. Dorothy Wordsworth, travelling through in 1803, was able to note that the glenwas uninhabited and the houses ruined.

5.6.2 Grazing

The Glen Coe hills are grazed by sheep and red deer. There are small numbers of mountain hares and roedeer, and wild goats are seen occasionally. Highland cattle are kept on the lower ground of Meall Mór andin the fields below Achtriochtan in the main glen (National Trust for Scotland 1998).

The Red Deer Commission count for 1995 estimated 130 deer on the land owned by the National Trust forScotland and 94 on the Glen Etive estate (National Trust for Scotland 1998). The NTS policy is to maintaina stable low population and a healthy ratio of sexes, although deer numbers are currently rising (JamesFenton, pers. comm.). During our survey we saw no deer on Meall Mór or in the Fionn Ghleann. We sawtwo hinds in Coire nan Lochan, and over 100 animals in both the Lairig Eilde and the Lairig Gartain. Therewere several large herds of hinds and calves and smaller ones of stags.

Although the sheep on the land owned by the National Trust for Scotland are now confined to Meall Mórand the fenced parks in the main glen, ewes and lambs from the Blackmount Estate were still grazing theunenclosed northern slopes and corries of the main glen until the end of 2002 (James Fenton, pers. comm.).Sheep are able to get onto the base-rich cliffs in Coire nam Beitheach, and in summer 2002 were grazingin Luzula-Geum tall-herb vegetation U17 on the outcrops below the west top of Bidean, as well as on thegrassy terrace and broken slopes below the main cliffs of Church Door Buttress. Small numbers of sheepwere seen in Coire nan Lochan in June but none in Coire nan Gabhail – the Lost Valley is surrounded bycliffs and boulder-fields and it cannot be easy for sheep to get in it.

There are a great many sheep in the Fionn Ghleann and on the south side of the main ridge of Bidean. Theyare grazing up onto the high montane ground and it may be some of these sheep that are dropping downinto upper Coire nam Beitheach.

There are many sheep on Meall Mór. Although the vegetation on the steep slopes is not grazed hard, theintensity of grazing is obviously enough to prevent a succession to woodland or scrub, and woodland, scruband Calluna heaths are confined to the crags and steeper ground. There are numerous bushes of Salix auritaon the gentle ground between the foot of the north-east face and the A82 road, but they are all short andsuppressed by browsing. There are also sheep on the summit of Meall Mór, but again the vegetation is notgrazed hard, although about 30% of the shoot-tips of Vaccinium myrtillus have been nipped off.

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Little specific management can be done for the rare montane plants on Bidean, beyond keeping sheepnumbers sufficently low. Most of them grow among boulders, on cliffs or in unstable gravel and fluctuationsin their populations may be natural.

The fence on Meall Mór is not really a good idea. Smaller species such as Potentilla crantzii, Carex capillaris,Tofieldia pusilla and perhaps even Dryas octopetala may be out-competed and over-shaded by tall grassesand herbs. The willows will certainly grow taller, but without patches of bare ground are unlikely to re-establish from seed, even assuming that plants of both sexes occur. Only female ones were seen duringthe survey.

In 1984 an experimental exclosure was erected on the slopes just to the west of the confluence of the Allt Coire Gabhail and the River Coe (Photograph in Executive Summary and Photographs 29–31). The heatherin this exclosure has come back mightily. There is now much Trichophorum-Erica heath M15 with a densetall sward of dwarf shrubs. Tree regeneration is more patchy, even though there is such a good source ofseed in the woods nearby. Young birches and rowans are growing up in the exclosure but not as denselyor as fast as one might expect.

In recent years sheep numbers on the north-facing slopes of the main glen have been progressively reduced,in an attempt to allow the woodlands on the cliffs to expand and to allow the vegetation of the open groundto become more natural. There is little obvious change yet, and in fact the vegetation was still sheep-grazedat the time of our study, as ewes and lambs from neighbouring estates were trespassing onto these slopesuntil the end of 2002 (James Fenton, pers. comm.). The vegetation in the experimental exclosure suggeststhat heather will re-grow on these open north-facing slopes and that woodland will spread, albeit slowly.

The area on Meall Mór with the willow scrub was stock-fenced in 2000, in an attempt to allow the Salixmyrsinites to recover from browsing and if possible to regenerate. After one and a half growing seasons theincrease in size and flowering of the species in the grasslands and mires is very noticeable. Many of whatwere open flushes are losing their distinctness. Grasses and the larger herbs are growing tall in them, sothat they blend into the surrounding grasslands. The grasses and sedges are growing tall and rank on themore accessible outcrops. There is a risk that smaller species may be overwhelmed, perhaps including theDryas octopetala and Carex capillaris which are important elements of the vegetation here.

It is important to remember that no grazing at all is as unnatural as the current overgrazing by unsustainablenumbers of sheep and deer. There would always have been natural herbivores: red and roe deer, voles,hares and a wide range of plant-eating invertebrates. In remote antiquity there would also have beenbeavers, brown bears, horses, wild boar, wild cattle and more species of deer (Tipping 2003). Theseanimals would have been preyed on by an array of large predators such as wolves and lynx, and in anatural ecosystem it is unlikely that the numbers of herbivores would have risen above the carrying-capacityof the land. The effects of no grazing at all are often fascinating, but can result in vegetation which has lessvalue for nature conservation than the grazed, much-modified precursor. For example, ungrazed grasslandscan become tall and rank, and a few robust species can come to dominate at the expense of smallergrasses, herbs and bryophytes. It can also be difficult for dwarf shrubs and trees to establish in such thickturf. Ungrazed woodlands can develop such a dense canopy that the ground vegetation is almost entirelyshaded out, and rich assemblages of grasses, ferns and herbs can be replaced by a sprinkling of etiolatedplants over expanses of bare soil. There are good examples of this in the exclosures at Rassal NNR

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(Averis 2002). In milder climates the woodland understorey can develop into thickets of bracken andbramble which themselves may shade out many more interesting species. Ungrazed woods can become sodark that important populations of bryophytes and, especially, lichens are unable to survive.

Although it may be necessary to exclude all grazing animals for a short period, in order to allow theregeneration of dwarf shrubs and trees, it is not advisable to exclude them indefinitely but rather to reducetheir numbers to a level at which trees and shrubs can survive. The ideal is to find a balance where deerand even sheep can live as part of a healthy and sustainable ecosystem.

5.6.3 Human views and recreation

Glen Coe has been a popular destination for climbers, walkers and holiday-makers since the early 20th century. This was not always so. At one time, the places where tourists now stop their cars and flockfrom buses to gaze at the peaks around them were the scene of fearful hurrying on atrocious roads througha huge and intimidating landscape. The Rev. John Lettice, a clergyman from Sussex, travelled through theregion in 1792 and described the Buachaille Etive Mór as ‘hideously disgustful’, and as late as the 19th century the author Charles Dickens described Glen Coe as ‘... perfectly terrible ... an awful place ...scores of glens high up, which form such haunts as you might imagine yourself wandering in, in the veryheight and madness of a fever’. But by the 1930s the mountaineer and writer W H Murray could not praisethe region enough: ‘... Glencoe, when the frozen towers of Bidean burned in the moon. In the last resort, itis the beauty of the mountain world ... that holds us spellbound, slaves till life ends.’ The packed car-parksand well-trodden footpaths show that many people today share his feelings. Several of the footpaths leadingup from the main glen have been restored and surfaced with stone blocks. These are hard wearing andblend into the slopes. There is little unsightly erosion here. The paths are rougher, looser and more widely erodedhigher up in the corries, and some of the scree-runs and gullies used as routes up onto the ridge are badlyworn. On the main Bidean ridge the ground is so stony that only locally are there noticeable signs of wear.

The paths through the Lairig Eilde and the Lairig Gartain are badly eroded where they traverse peat,especially in the Lairig Gartain where the path is all but unusable except in the driest conditions. There is awide swathe of eroded and trampled peat. Even so, although eroded paths and foot-worn summits are verynoticeable, their effects on the vegetation and flora are negligible compared with the depredations of thesheep. There has been some concern that the Saxifraga cernua and S. rivularis may be susceptible todamage by climbers, especially at the base of Collie’s Pinnacle and in Hourglass Gully. This may be so.However, both species are plants of unstable habitats and are able to tolerate the disturbance caused bysnow movements and solifluction. The bulbils of S. cernua might possibly be spread into new sites onclimbers’ boots. People walk up Coire nam Beitheach to reach the ridge of Bidean, but they tend to ascenda few well-used gullies, dry scree-runs and boulder-fields, and avoid the wetter, less stable slopes where therare plants grow. At the time of our survey there was no sign of any human activity at or near any of theplaces where the rare plants grow in Coire nam Beitheach.

Rocks fall frequently in the high corries. They are dislodged by walkers and by sheep grazing the steepunstable slopes and lower crags, but they also move with no obvious cause. There were several minorrockfalls during the days I spent up there. Fryday (1993) found evidence in the lichens for much recent re-distribution and re-sorting of the scree-fans in the main glen.

The Lost Valley of lower Coire Gabhail was described by the climber and writer W H Murray in the 1930sas ‘a green mountain lawn ... a man might ... pitch a tent on that meadow’, giving the impression that

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the glen floor was covered in grassland. Now it is almost entirely stony and there are only isolated patchesof grassland; most of them on thin stony soils where it would be most uncomfortable to camp.

5.7 Assessment of the Upland Vegetation Sur vey vegetation map of Glen Coe

The vegetation of the western part of our survey area – Bidean nam Bian SSSI – was surveyed by the Nature Conservancy Council’s Upland Vegetation Survey (UVS) team in 1981, 1982 and 1987. Theirvegetation map was produced as a series of transparent overlays to black-and-white vertical aerialphotogrpahs at a nominal scale of 1:25,000. They mapped most of the vegetation before the NationalVegetation Classification was produced, so they used the simpler classification devised by Birks & Ratcliffe(1980). They added some mire NVC types in 1987. Altogether they found 29 types of vegetation. Thesecould be equated with up to 53 NVC communities or sub-communities. In contrast, we found 129 vegetationand habitat types in this same area. The scheme of Birks & Ratcliffe is not as detailed as the NVC, but itdoes distinguish most of the types of vegetation which occur in this area.

The UVS vegetation map did not adequately describe the vegetation of this site. The mapped vegetationunits are much larger than those in our survey, and the labels do not show the full range of vegetation types.Some vegetation was misclassified, and some vegetation types and much of the small-scale variation invegetation was overlooked. For example, the north-eastern slope of Meall Mór was mapped as ‘C1a/e(R1a + T)’ which equates with U4a acid Festuca-Agrostis grassland and CG10 herb-rich Festuca-Agrostisgrassland with scattered cliffs and trees. This omits many other vegetation types which occur here, some ofwhich are hard to miss: extensive Molinia grassland (M25b/c), Nardus grassland (U5a/c/e), patches ofOreopteris limbosperma fern vegetation (U19), base-enriched flushes (M10 and M11), superb examples oftall herb vegetation (U15 and U17) and numerous patches of woodland (W7b/W9b/W17a).

Another example is an area on the north side of Bidean nam Bian mapped in the UVS survey as‘C1a/R2a(D6)’ which equates to U4a acid Festuca-Agrostis grassland with scree, and smaller areas of U16 Luzula sylvatica or U19 Oreopteris vegetation or both. This area spans a large altitudinal range(100–930m), and, not suprisingly, a large range of variation in vegetation: Festuca-Agrostis grassland (U4),bracken (U20), Oreopteris fern beds (U19) and Trichophorum-Erica wet heath (M15c) on the lower slopes;losing bracken but gaining Luzula (U16) and Vaccinium heaths (H18) higher up, and passing further upslopeinto complex mosaics of Festuca-Agrostis, Nardus and Deschampsia cespitosa grasslands (U4, U5, U7 and U13),wet heath (M15c), Vaccinium heath (H18), fern snow-beds (U18) and bryophyte snow-beds (U11 and U12).

Bryophyte-dominated late snow-beds (U11 and U12) and bryophyte-rich Calluna heath (H21b) are amongthe most notable vegetation types at this site, but were completely missed in the UVS survey, even thoughthey can be identified as separate types in the Birks & Ratcliffe classification. Another notable habitat – tallherb vegetation (U15 and U17) – is widespread and conspicuous on the northern slopes of Bidean namBian and Meall Mór but was completely missed here by the UVS survey; they mapped just one tiny patchof ‘D1’ (= U17), on the south side of Bidean.

Other examples could be mentioned, but it is clear that the UVS survey was inadequate because themapping was too simple and missed a lot of the particularly interesting vegetation. There are severalpossible reasons for the inadequacy of the UVS survey.

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One possible reason is that the results of the UVS surveys were presented at a small scale. Although the fieldmapping was done on enlarged aerial photographs at a scale of 1:10,000 the final maps were drawn upat 1:25,000. Surveyors did not take the trouble to delineate small patches of vegetation which they knewcould not be fitted onto the final map. Nevertheless some of the larger units really should have beensubdivided, and the labels could have been more detailed.

Second, the Birks & Ratcliffe classification is neither as comprehensive nor as definitive as the NVC. Havingsaid that, the Birks & Ratcliffe scheme is adequate for describing upland vegetation; it was, after all, basedon McVean & Ratcliffe (1962) whose samples form the backbone of most of the upland NVC types.Nevertheless the NVC is a more precise tool. It opens one’s mind to the possibility of greater variation in vegetation than that inherent in the earlier scheme. Many Birks & Ratcliffe types equate with NVC communities rather than sub-communites. For example, grassland described as ‘C1e’ using the Birks & Ratcliffe scheme could be any one of the three sub-communities of CG10 in the NVC. Similarly theBirks & Ratcliffe heathland type ‘B1’ is represented here by the NVC types H10 (with four sub-communities),H12 (three sub-communities, two of which were found in our survey) and H21 (two sub-communities, bothfound in our survey).

A third possible reason for the poor quality of the UVS maps is that some of the vegetation at this site canlook quite uniform even at fairly close range, but is in fact a very complicated mosaic of several types. A good example of this is the grasslands on the north-eastern slopes of Meall Mór. It is a common experienceto look at a hillside only a few hundred metres distant and not to see any clear boundaries. Small patchesof many types of vegetation can form intricate mosaics. These do not show up as distinct tones on the aerialphotographs which were used for mapping by the earlier surveyors. It was found during UVS surveys thatthe aerial photographs of some hills – for example in the Breadalbanes – showed little but featureless hillsidesof a uniform texture and pale-grey tone. Cliffs with tall-herb ledges can be so thickly vegetated that they donot stand out well from the smooth slopes in the photographs. And since there are no contours on thephotographs it was particularly difficult both to navigate and to plot patches of vegetation with anyaccuracy. Given this it is hardly surprising that the UVS surveyors found themselves unequal to the task ofreconciling the vegetation and the photographs. Even with good-quality 1:10,000 maps and the guidanceof the NVC the vegetation of parts of Glen Coe is still bewilderingly complex and we, too, had to mapmany stands of vegetation as large mosaics of numerous different types.

Finally, most of the UVS survey work was done in the early years of the Upland Vegetation Survey project,at a time when most of the surveyors had insufficient experience to do justice to a place such as Glen Coe.

5.8 Extra note on birds and animals seen during this sur vey

During the course of this vegetation survey we saw several bird species: grey heron, mallard, golden eagle(a pair), buzzard, kestrel, ptarmigan (female with 9–10 chicks and a pair with chicks), common sandpiper,snipe, black-headed gull, herring gull, sand martin, swallow, skylark, meadow pipit, grey wagtail, piedwagtail, dipper, wren, wheatear, whinchat, stonechat, ring ouzel, raven, hooded crow, willow warbler,chaffinch and twite. We also saw red deer and fox.

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6 ACKNOWLEDGMENTS

Dominic Sargent and Debbie Greene of Scottish Natural Heritage organised the contract, providedbackground information and made comments on an earlier draft of this report. James Fenton of the NationalTrust for Scotland also provided information. We would like to thank Scott McCombie and other local staffof the National Trust for Scotland for assistance and encouragement, the Maps Office, SNH, Edinbugh forthe base-maps, and Judie Drever and Archie Gibb of Glendevin, Inchree and Mr and Mrs Banks of Brudair,Ballachulish, for their hospitality at their bed and breakfast establishments.

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7 REFERENCES

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Averis, A.M. (1994). The ecology of a Atlantic liverwort community. PhD thesis. University of Edinburgh.

Averis, A.M. (2001). Vegetation survey of Lì & Coire Dhorrcail, Knoydart, June 2001. Report for the John Muir Trust.

Averis, A.M. (2002). Vegetation survey of Rassal SSSI & cSAC, June 2001. Scottish Natural Heritagereport HT/LD09/01/02/28.

Averis, A.B.G. & Averis, A.M. (1995). The vegetation of north Mull. Scottish Natural Heritage report5/F2B/650.

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a survey of the vegetation of glen coe, scotland, summer 2002

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