NATIONAL VEGETATION

CLASSIFICATION

field guide to
mires and heaths
National Vegetation Classification:
Field guide to mires and heaths
Cover photograph:
Studland Heath
National Nature Reserve, Dorset.
© English Nature
National Vegetation Classification:
Field guide to mires and heaths
T. Elkington, N. Dayton, D.L. Jackson and I.M. Strachan
Joint Nature Conservation Committee
Monkstone House
City Road
Peterborough
PE1 1JY
UK
ISBN 1 86107 526 X
© JNCC 2001
Contents
1. Introduction 7
National Vegetation Classification 7
Mires and heaths: scope of this guide 7
Using this guide 7
References 8
2. Dendrogram keys to mire communities 9
3. Mire community descriptions and sub-community keys 19
M1 Sphagnum auriculatum bog pool community 19
M2 Sphagnum cuspidatum/recurvum bog pool community 20
M3 Eriophorum angustifolium bog pool community 21
M4 Carex rostrata – Sphagnum recurvum mire 22
M5 Carex rostrata – Sphagnum squarrosum mire 23
M6 Carex echinata – Sphagnum recurvum/auriculatum mire 24
M7 Carex curta – Sphagnum russowii mire 26
M8 Carex rostrata – Sphagnum warnstorfii mire 27
M9 Carex rostrata – Calliergon cuspidatum/giganteum mire 28
M10 Carex dioica – Pinguicula vulgaris mire 30
M11 Carex demissa – Saxifraga aizoides mire 32
M12 Carex saxatilis mire 33
M13 Schoenus nigricans – Juncus subnodulosus mire 34
M14 Schoenus nigricans – Narthecium ossifragum mire 34
M15 Scirpus cespitosus – Erica tetralix wet heath 36
M16 Erica tetralix – Sphagnum compactum wet heath 38
M17 Scirpus cespitosus – Eriophorum vaginatum blanket mire 40
M18 Erica tetralix – Sphagnum papillosum raised & blanket mire 42
M19 Calluna vulgaris – Eriophorum vaginatum blanket mire 44
M20 Eriophorum vaginatum blanket & raised mire 46
M21 Narthecium ossifragum – Sphagnum papillosum valley mire 47
M22 Juncus subnodulosus – Cirsium palustre fen-meadow 48
M23 Juncus effusus/acutiflorus – Galium palustre rush-pasture 50
M24 Molinia caerulea – Cirsium dissectum fen-meadow 52
M25 Molinia caerulea – Potentilla erecta mire 54
M26 Molinia caerulea – Crepis paludosa mire 56
M27 Filipendula ulmaria – Angelica sylvestris mire 58
M28 Iris pseudacorus – Filipendula ulmaria mire 60
M29 Hypericum elodes – Potamogeton polygonifolius soakway 62
M30 Related vegetation of seasonally inundated habitats 62
M31 Anthelia julacea – Sphagnum auriculatum spring 63
M32 Philonotis fontana – Saxifraga stellaris spring 64

5
 M33 Pohlia wahlenbergii var. glacialis spring 66
M34 Carex demissa – Koenigia islandica flush 66
M35 Ranunculus omiophyllus – Montia fontana rill 67
M36 Lowland springs and streambanks of shaded situations 67
M37 Cratoneuron commutatum – Festuca rubra spring 68
M38 Cratoneuron commutatum – Carex nigra spring 69
4. Dendrogram keys to heath communities 70
5. Heath community descriptions and sub-community keys 76
H1 Calluna vulgaris – Festuca ovina heath 76
H2 Calluna vulgaris – Ulex minor heath 78
H3 Ulex minor – Agrostis curtisii heath 80
H4 Ulex gallii – Agrostis curtisii heath 82
H5 Erica vagans – Schoenus nigricans heath 84
H6 Erica vagans – Ulex europaeus heath 86
H7 Calluna vulgaris – Scilla verna heath 88
H8 Calluna vulgaris – Ulex gallii heath 90
H9 Calluna vulgaris – Deschampsia flexuosa heath 92
H10 Calluna vulgaris – Erica cinerea heath 94
H11 Calluna vulgaris – Carex arenaria heath 96
H12 Calluna vulgaris – Vaccinium myrtillus heath 98
H13 Calluna vulgaris – Cladonia arbuscula heath 100
H14 Calluna vulgaris – Racomitrium lanuginosum heath 102
H15 Calluna vulgaris – Juniperus communis ssp. nana heath 104
H16 Calluna vulgaris – Arctostaphylos uva-ursi heath 104
H17 Calluna vulgaris – Arctostaphylos alpinus heath 106
H18 Vaccinium myrtillus – Deschampsia flexuosa heath 108
H19 Vaccinium myrtillus – Cladonia arbuscula heath 110
H20 Vaccinium myrtillus – Racomitrium lanuginosum heath 112
H21 Calluna vulgaris – Vaccinium myrtillus – Sphagnum
capillifolium heath 114
H22 Vaccinium myrtillus – Rubus chamaemorus heath 116

6
1 Introduction
National Vegetation Classification dune slack communities and Epilobium hirsutum
stands are described in Volume 5 (Rodwell 2000).
Since its development in the 1980s, the National ‘Grass heaths’, Dryas heaths and related lichen
Vegetation Classification (NVC) has become the and bryophyte dominated vegetation are included
standard classification used for describing vegeta- in Volume 3 (Rodwell 1992), as are certain wet
tion in Britain. Whereas many other classifications grasslands. Volume 1 (Rodwell 1991b) encompasses
are restricted to particular types of vegetation, the wet woodland and scrub vegetation. Companion
NVC aims to describe all the vegetation of Great guides to volumes 1 and 3 have also been pub-
Britain. This means that it is possible to analyse, lished by JNCC (Hall et al. 2001; Cooper 1997).
and map, a complex site, composed of several Users of this guide should also note that most
habitat types (e.g. woodland, scrub, heathland and ‘wet heath’ vegetation is described in the NVC
bog) using the same classification system. with the mires rather than the heaths (as M15
The NVC is a ‘phytosociological’ classification, Scirpus cespitosus – Erica tetralix wet heath and
classifying vegetation solely on the basis of the M16 Erica tetralix – Sphagnum compactum wet
plant species of which it is composed. The result- heath), because of its floristic affinities. Helpful
ing communities can usually be correlated to other insight into the floristic relationships of NVC
factors, such as geology and soils, climate, water types can be gained from the Phytosociological
chemistry and management; but the plant species Conspectus in Volume 5 of British Plant
alone are used to assign the vegetation to a Communities. This places all NVC communities
community. within a hierarchical framework of European
The NVC breaks down each broad vegetation vegetation.
type (e.g. heath, mire, woodland) into communities, Various gaps in coverage of the NVC have been
designated by a number and name (e.g. H4 Ulex gal- identified at community and sub-community level
lii – Agrostis curtisii heath, M10 Carex dioica – subsequent to the publication of British Plant
Pinguicula vulgaris mire, W4 Betula pubescens – Communities. These include several mire and
Molinia caerulea woodland). Many (but not all) of heath types, as outlined in JNCC Report No. 302
these communities contain two or more sub-com- Review of coverage of the National Vegetation
munities, designated by a letter (e.g. H4b Ulex gal- Classification (Rodwell et al. 2000). No attempt
lii – Agrostis curtisii heath, Festuca ovina sub-com- has been made to incorporate these here, pending
munity). Sub-communities in a few cases are fur- further analysis and formal description.
ther divided into variants (e.g. M10bi and ii).

Mires and heaths: Using this guide

scope of this guide
The summary descriptions provided here are
The second volume of British Plant Communities derived directly from the full accounts prepared
was published in 1991 (Rodwell 1991a). It provides by John Rodwell, but are in no way a substitute for
a detailed account of 38 mire communities and 22 them. Rather they are intended as an aide-mem-
heath communities, giving information on their oire to assist surveyors in the field or for anyone
composition, structure and distribution, their else wishing to familiarise themselves with the
affinities to other types of vegetation, both in overall scheme of classification for mires and
Britain and on the Continent, and the relation of heaths. Anyone who uses this book should always
the communities described within the NVC to check their results against the frequency tables
those previously described by other authors. The and full descriptions for each community in
scope of this field guide is identical to that of Volume 2 of British Plant Communities. The
Volume 2. descriptions are not intended to take account of
A number of vegetation types which might also the results of recent survey work undertaken by
be considered as ‘mires’ or ‘heaths’ are described the three country agencies (Countryside Council
in other volumes of British Plant Communities, for Wales, English Nature, Scottish Natural
and so are not included here. Aquatic, swamp and Heritage) which may help circumscribe some of
tall-herb fen communities can be found in Volume the communities more tightly and improve our
4 (Rodwell 1995), whilst inundation communities, understanding of community distributions.

7
A series of dendrograms have been produced to
show the broad floristic relationships between the
References
main communities and between the sub-commu-
nities for each community where these exist. Cooper, E A (1997) Summary descriptions of Nat-
These dendrograms are only intended as guides ional Vegetation Classification grassland and
and should not be followed slavishly. Details of montane communities. Joint Nature Con-
variants, if indicated, can be found in Volume 2 of servation Committee, Peterborough (UK Nature
British Plant Communities. Conservation No. 14).
The amount of any particular species is Hall, J E, Kirby, K J and Whitbread, A M (2001)
referred to both in terms of its frequency and National Vegetation Classification: field guide
abundance. ‘Frequency’ refers to how often a plant to woodland. Joint Nature Conservation Com-
is found in moving from one sample or vegetation mittee, Peterborough.
to the next, irrespective of how much of that Rodwell, J S, (ed) (1991a) British Plant Commun-
species is present in each sample. This is sum- ities. Volume 2. Mires and heaths. Cambridge
marised in the published tables as classes denoted University Press, Cambridge.
by the Roman numerals I to V: 1-20% frequency Rodwell, J S, (ed) (1991b) British Plant Commun-
(that is, up to one sample in five) = I, 21-40% = II, ities. Volume 1. Woodlands and scrub. Cambridge
41-60% = III, 61-80% = IV, and 81-100% = V. The University Press, Cambridge.
summary descriptions follow the usual conven- Rodwell, J S, (ed) (1992) British Plant Commun-
tion of referring to species of frequency classes IV ities. Volume 3. Grasslands and montane com-
and V in a particular community as its constants, munities. Cambridge University Press, Cambridge.
with those species of class III as common or fre- Rodwell, J S, (ed) (1995) British Plant Commun-
quent, of class II as occasional and of class I as ities. Volume 4. Aquatic communities, swamps
scarce or rare. The term ‘abundance’, on the other and tall-herb fens. Cambridge University Press,
hand, is used to describe how much of a plant is Cambridge.
present in a sample, irrespective of how frequent Rodwell, J S, (ed) (2000) British Plant Commun-
or rare it is among the samples. It is summarised ities. Volume 5. Maritime communities and
in the published tables as bracketed numbers for vegetation of open habitats. Cambridge
the Domin ranges, and is referred to in the text University Press, Cambridge.
here, as in the published descriptions, using such Rodwell, J S, Dring, J C, Averis, A B G, Proctor, M
terms as dominant, abundant, frequent and sparse. C F, Malloch, A J C, Schaminée, J N J and
The nomenclature for plant species used in Dargie, T C D (2000) Review of coverage of the
British Plant Communities has been followed in National Vegetation Classification. JNCC Report
this publication for consistency. Botanists more No. 302.
familiar with Stace’s New flora of the British Isles Stace, C (1997) New flora of the British Isles. 2nd
(Stace 1997) may not recognise names such as ed. Cambridge University Press, Cambridge.
(Stace equivalent in brackets): Scirpus cespitosus
(Trichophorum cespitosum), Carex demissa
(C. viridula ssp. oedocarpa), C. lepidocarpa
(C. viridula ssp. brachyrrhyncha) and Silene vul-
garis maritima (S. uniflora). Amongst cryptogams,
the common lichen of heaths and bogs referred to
here as Cladonia impexa is now generally known
as C. portentosa.

8
 A Key to mires

1. Sphagnum spp. 1a. Sphagnum spp. not

2
constant or very frequent constant in the vegetation.
in the vegetation, either Occasional small patches
continuous or patchy. may be present but these
are never prominent.
2. Calluna 2a. Sub-shrubs
vulgaris not constant;
and/or though Calluna 7. Spring or flush vegeta- 7a. More bulky
Erica vulgaris and/or tion; an open bryophyte- vegetation in
tetralix Erica tetralix rich mat in which any of: which sedges,
constant. may be present at Cratoneuron commutatum, rushes, grasses
Usually low abundance. Philonotis fontana, or tall herbs are
quite exten- Anthelia julacea, Pohlia prominent.
sive mires wahlenbergii or Koenigia
over peat. islandica can be prominent.

3. Dominated by 3a. Sedges may 8. Vegetation 8a. Sedges may

sedges be present but dominated by be frequent in the
Eriophorum, are subordinate sedges, Carex sward but not
Carex spp., or to bryophytes or or Eriophorum dominant and
tall rushes to other vascular spp., either in never structurally
Juncus spp. species. low, open mats important.
or taller, more
dense mires.
4. Vegetation 4a. Vege- 5. Mossy 5a. Open
9a. Tall rushes,
dominated by tation domi- carpet springs or rills
Juncus spp.,
Eriophorum nated by dominated with constant,
and/or tall fen
angustifolium other sedges by semi- but patchy
herbs dominant.
mire communities

and/or E. or tall aquatic Sphagnum

Molinia caerulea
vaginatum. rushes. Sphagnum auriculatum.
rare but may be
spp., includ-
locally abundant
ing S. cuspi-
Dendrogram keys to

in western
datum with
mires.
vascular 6. Semi-
species aquatic rills 6a. Open
typically and flushes bryophyte
sparse. dominated by mats around 9. Molinia caerulea
Bog pools. Potamogeton springs. constant and usually
polygonifolius Sphagnum structurally important
and/or auriculatum in the vegetation; with
Montia can be or without Schoenus
fontana. constant. nigricans.

9
See Fig 1. See Fig 2. See Fig 3. See Fig 4. See Fig 5. See Fig 6. See Fig 6. See Fig 7. See Fig 8. See Fig 9.
Mires Figure 1

Fig 1. Key to vegetation with constant Sphagnum spp. and constant ericoid sub-shrubs

1. Eriophorum vaginatum 1a. See next page

(either as prominent tussocks
or sparse fronds) and/or
Sphagnum papillosum and/
or S. magellanicum constant.
Deep peat > 1 m. Blanket
and raised mires.

2. Molinia caerulea 2a. Scirpus cespitosus

and/or Scirpus cespitosus not constant and rarely
constant with at least prominent, though can
some: Narthecium be frequent in patches.
ossifragum, Eriophorum Molinia caerulea
angustifolium, Potentilla generally absent (can
erecta, Sphagnum form mosaics with tus-
papillosum and socky Eriophorum vagi-
S. capillifolium. natum in degraded forms
of M19.)

3. Sphagnum spp. 3a. Sphagnum spp. patchy,

prominent, especially though S. capillifolium
S. papillosum, S. capilli- constant. S. tenellum,
folium and S. tenellum, S. magellanicum and
often S. magellanicum. Odontoschisma sphagni
Associates include at generally absent or very rare.
least some of Vaccinium
oxycoccos, Drosera
rotundifolia and 4. Calluna vulgaris and 4a. Vegetation dominated
Odontoschisma sphagni. Eriophorum vaginatum form by tussocks of Eriophorum
the bulk of the vegetation, vaginatum with Calluna
E. vaginatum usually tus- vulgaris and Sphagnum
socky, with Sphagnum capil- spp. at best infrequent
lifolium, E. angustifolium and patchy. Mounds of
and pleurocarpous mosses: Empetrum nigrum and/or
Pleurozium schreberi, Vaccinium myrtillus with
Hylocomium splendens constant Eriophorum
and Hypnum jutlandicum. angustifolium and
Deschampsia flexuosa.

M17 Scirpus cespitosus – M18 Erica tetralix – M19 Calluna vulgaris – M20b Eriophorum
Eriophorum vaginatum Sphagnum papillosum Eriophorum vaginatum vaginatum blanket and
blanket mire raised and blanket mire blanket mire raised mire, Calluna
vulgaris – Cladonia
spp. sub-community

Burning can increase the The Eriophorum mire

proportion of Calluna is apparently biotically
vulgaris or Deschampsia derived from M19 blanket
flexuosa and Vaccinium bog or M18 raised mire;
myrtillus. Grazing usually the degree of floristic
decreases the abundance of impoverishment depend-
C. vulgaris, ultimately to its ing on the intensity and
disappearance, especially if duration of destructive
combined with large-scale management practice.
burns. Such disturbance can There is therefore a con-
shift the floristics of this tinuous gradation between
community towards the richer and poorer stands.
impoverished M20 mire. In the wet north-west
Lower altitude stands some Sphagnum spp.
(<400 m) to the west usually can persist even in
include constant Erica tetral- highly degraded stands,
ix which is replaced in high- especially S. recurvum
er and more eastern stands which is relatively toler-
by Empetrum nigrum and ant to some disturbance.
Rubus chamaemorus.

10
From previous page:
1a. Both Eriophorum vaginatum
and Sphagnum papillosum
generally absent, though may
be patchily present at low
frequency in the wetter sub-
communities of the M15
Erica – Scirpus mire.

5. Calluna vulgaris and Erica 5a. Erica tetralix constant.

tetralix both prominent with Calluna vulgaris absent or
Eriophorum angustifolium present at low frequency in
and Molinia caerulea constant. the vegetation.

6. Narthecium ossifragum 6a. Narthecium

constant and abundant ossifragum present at
with Drosera rotundifolia, low frequency. Ericoid
Eriophorum angustifolium, sub-shrubs and at least
Sphagnum papillosum, one of Scirpus cespitosus,
plus S. auriculatum and/ Molinia caerulea and
or S. recurvum. Schoenus Eriophorum angustifolium
nigricans absent or make up the bulk of the
very rare. vegetation over patchy
sphagna, S. capillifolium
or S. compactum.
Wet heaths.

7. Erica tetralix usually 7a. Calluna vulgaris usu- 8. Schoenus 8a. Molinia
the most prominent ally the most prominent nigricans caerulea
sub-shrub. Sphagnum sub-shrub except where constant with dominant, dense
layer dominated by grazing pressure has prominent and tussocky.
S. compactum and favoured the expansion Narthecium Sparse Erica
S. tenellum. Molinia of E. tetralix. Sphagnum ossifragum tetralix and
caerulea usually domi- spp. dominated by and Sphagnum Potentilla
nant amongst the vascu- S. capillifolium and subnitens. erecta constant.
lar monocot associates. S. subnitens with Sphagnum
occasional S. papillosum palustre and
in the wetter S. recurvum can
sub-communities. be prominent
in wet north-
western stands.

M21 Narthecium M16 Erica tetralix – M15 Scirpus cespitosus – M14 Schoenus M25a Molinia
ossifragum – Sphagnum Sphagnum compactum Erica tetralix wet heath nigricans – caerulea –
papillosum valley mire wet heath Narthecium Potentilla
ossifragum erecta mire,
May be confused with the Wet heath primarily of This is a very variable mire Erica tetralix
flushed wet heath, M15a; the south and east of vegetation type and, of sub-community
however the sedges Britain, this community the major components,
Carex panicea, C. echinata provides the major locus any can be dominant with Burning and
are constant in the latter for Scirpus cespitosus up to two of the others grazing of M15
and Myrica gale is more and Eriophorum missing. A community wet heath can
frequent. Pinguicula vul- vaginatum (though rare) primarily of north-west favour an
garis is common in M15a in this part of the Britain, it includes most increase in
but is replaced in the country. of the heather-dominated Molinia which
M21 community by less vegetation intermediate in shifts the
frequent P. lusitanica. character between the dry vegetation
heath and blanket mire towards that of
types. Where heavily the Erica tetralix
grazed and/or burnt, the sub-community
ericoids can become very of M25. The
sparse and these stands similarities
are often transitional to between M25a
the Molinia-dominated and Molinia-
community M25, or rich M15 show
drier stands to the Juncus the transition
squarrosus grassland U6. between these
two communities
through distur-
bance.

11
Mires Figure 2

Fig 2. Key to vegetation with constant Sphagnum spp. and dominated by Eriophorum vaginatum and/or E. angustifolium

1. Vegetation dominated by 1a. Vegetation dominated by

Eriophorum angustifolium in Eriophorum vaginatum in
which Sphagnum cuspidatum which E. angustifolium
and/or Drepanocladus fluitans can be constant but is
can be prominent. Bog pools always subordinate to
and erosion hagg runnels with- E. vaginatum.
in blanket peat.
2. Tussocky E. vaginatum 2a. It can be difficult to
with very patchy Sphagnum separate richer forms of the
capillifolium, constant M20b sub-community from
Deschampsia flexuosa poor and degraded stands of
and mounds of Vaccinium the M19 blanket mire, since
myrtillus and/or Empetrum there is a continuum of
nigrum nigrum. Calluna vegetation types between
vulgaris usually absent but these two communities with
can be present as scattered increasing disturbance to the
sprigs at low frequency. latter. Generally, if C. vulgaris
North-western stands also is constant with a reasonably
feature prominent Poly- intact Sphagnum flora this
trichum commune. community can be categorised
with the M19 mires, though
E. vaginatum may be visually
dominant.

M3 Eriophorum M20b Eriophorum vagina- M19 Calluna vulgaris –

angustifolium bog pool tum blanket and raised mire, Eriophorum vaginatum
Calluna vulgaris – Cladonia blanket mire
spp. sub-community
This community is typical of
recent or disturbed bog pools or
forms an early seral stage in the
transition from exposed peat
back to mire vegetation. Poorer
stands of the M2b Sphagnum
recurvum bog pool can be very
similar to Sphagnum-rich M3
stands, reflecting the transitional
continuum between these
communities.

12
 Fig 3. Key to communities with constant Sphagnum spp. and dominated by sedges or tall rushes

1. Carex rostrata constant 1a. Carex rostrata absent or

and prominent. present only at low frequency.

2. Moss layer dominated 2a. Sphagnum recurvum 4. Vegetation dominated by 4a. Tall rushes absent or
Mires Figure 3

by Sphagnum recurvum, absent or can be present but tall rushes; Juncus effusus only very sparse and patchy.
S. cuspidatum and is always subordinate to and/or Juncus acutiflorus
Polytrichum commune, other Sphagnum spp. usually with Carex echinata
never S. squarrosum. Carex but this sometimes very
curta can be abundant in 3. Sphagnum squarrosum 3a. Sphagnum warnstorfii sparse. Associates include: 5. Vegetation dominated 5a. Carex curta absent
some stands (DOMIN 2-9) constant with at least three constant with at least three Polytrichum commune, by Carex curta with con- or very rare. Carex
but S. warnstorfii never of: Potentilla palustris, of Carex nigra, Epilobium Potentilla erecta, Viola stant Sphagnum russowii echinata constant
present. Carex nigra, Eriophorum palustre, Selaginella palustris over a moss and at least two of Carex over a moss layer
angustifolium, Succisa selaginoides, Viola palustris, layer dominated by echinata, Eriophorum dominated by
pratensis, Aulacomnium Aulacomnium palustre, Sphagnum palustre and angustifolium, Viola Sphagnum palustre
palustre. Calliergon cuspidatum, either S. recurvum or palustris, S. papillosum. and either S.recurvum
Rhizomnium pseudopuncta- S. auriculatum. or S. auriculatum.
tum and Sphagnum teres. Other sedges frequent
including C. panicea,
C. nigra and Eriophorum
angustifolium. Nardus
stricta and Juncus
squarrosus can be
very frequent in
some stands.

M4 Carex rostrata – M5 Carex rostrata – M8 Carex rostrata – M6 Carex echinata – M7 Carex curta – M6 Carex echinata –
Sphagnum recurvum mire Sphagnum squarrosum mire Sphagnum warnstorfii mire Sphagnum recurvum/auric- Sphagnum russowii mire Sphagnum recurvum/
ulatum mire, M6c Juncus auriculatum mire,
Flush or floating raft over Montane flush vegetation, effusus sub-community M6a Carex echinata
Montane flush vegetation,
liquid peat. Often forms almost always between and M6d J. acutiflorus sub-community and
almost always between
stage in the transition to 400 m and 800 m asl. sub-community M6b Carex nigra –
650 m and 1100 m asl in
open water or as part of a Nardus stricta sub-
central Scotland.
complex mosaic with other community
C. rostrata dominated com-
munities (M9, S9, S27). Stands with abundant
Carex nigra, Erio-
phorum angustifolium
and Juncus squarrosus
may be transitional
to wetter forms of the
U6 J. squarrosus –
Festuca ovina
grassland.

13
Mires Figure 4

Fig 4. Key to bog pools dominated by open Sphagnum carpets

1. Bog pool dominated by 1a. Bog pool dominated by

Sphagnum auriculatum and Sphagnum cuspidatum and/or
S. cuspidatum, usually with S. recurvum. S. auriculatum
some Menyanthes trifoliata never present. Scattered
and Eriophorum angustifolium. Eriophorum angustifolium
throughout pool; Erica tetralix,
Drosera rotundifolia and other
vascular species around
margins.

M1 Sphagnum auriculatum M2 Sphagnum cuspidatum

bog pool /recurvum bog pool

As temporary pools infill,

Eriophorum angustifolium
increases in prominence. Such
pools can exist in all transition-
al stages between M2 and M3
bog pool floristics.

Mires Figure 5

Fig 5. Key to semi-aquatic rills and flushes dominated by

Potamogeton polygonifolius and/or Montia fontana

1. Flush vegetation dominated 1a. Rills and spring-head

by Potamogeton polygonifolius vegetation dominated by Montia
with Hypericum elodes, Ranun- fontana and Ranunculus omio-
culus flammula and Juncus phyllus, usually with Ranun-
bulbosus. culus flammula and some of:
Juncus bulbosus, J. articulatus,
Potamogeton polygonifolius,
Myosotis secunda and Agrostis
stolonifera.

M29 Hypericum elodes – M35 Ranunculus omiophyllus –

Potamogeton polygonifolius Montia fontana rill
soakway

This community has been This community has been

poorly sampled from the north. poorly sampled from the north
Hypericum elodes is abundant of Britain. Stands similar in
in this community across south- floristics to the M35 rills but
west Britain but declines to the lacking R. omiophyllus are
north and is absent from most occasional along ditch-lines at
stands north of the Scottish the margins of improved land
Borders except for occasional in lowland areas.
outliers in the Western Isles.

14
 Fig 6. Key to open, bryophyte-rich springs and flushes

1. Montane or sub-montane 1a. Montane springs or rills

bryophyte-rich spring-heads in which Cratoneuron
dominated by Cratoneuron commutatum and Philonotis
commutatum and/or fontana are absent or very
Mires Figure 6

Philonotis fontana. sparse.

2. Moss carpet dominated by 2a. Cratoneuron commutatum 4. Open mat dominated 4a. Montane springs and
Cratoneuron commutatum absent or only present at low by Anthelia julacea with rills lacking Anthelia
or occasionally by frequency. Philonotis fontana constant Sphagnum julacea.
C. filicinum. constant with two or more of: auriculatum, Marsupella
Saxifraga stellaris, Scapania emarginata, Scapania 5. Spongy bryophyte 5a. Bryophyte mat
undulata, Deschampsia undulata and Deschampsia carpets dominated by Pohlia with open areas of
cespitosa, Dicranella palus- cespitosa. wahlenbergii var. glacialis. stony ground. Carex
tris, Stellaria alsine, Constants include: Saxifraga demissa and Koenigia
Sphagnum auriculatum. stellaris, Deschampsia cespi- islandica constant
3. Species-rich springs in 3a. Springs overwhelmingly tosa and Pohlia ludwigii. with Deschampsia
which Carex spp. are fre- dominated by Cratoneuron Cerastium cerastoides cespitosa, Saxifraga
quent, though sparse, espe- commutatum with can be frequent. stellaris and
cially C. nigra, C. panicea Philonotis fontana rare to Scapania undulata.
and C. demissa. Diverse occasional. Small sedges are
herbs including some of: scarce and the diversity of
Cardamine pratensis, vascular plants quite poor,
Leontodon autumnalis, usually including Cardamine
Trifolium repens and pratensis, Festuca rubra
Polygonum viviparum. and Agrostis stolonifera.
Philonotis fontana can be
frequent in the moss carpet.

M38 Cratoneuron M37 Cratoneuron M32 Philonotis fontana – M31 Anthelia julacea – M33 Pohlia wahlenbergii M34 Carex demissa –
commutatum – Carex commutatum – Festuca Saxifraga stellaris spring Sphagnum auriculatum var. glacialis spring Koenigia islandica
nigra spring rubra spring spring flush

Light grazing is important in Typically from base-rich Very variable community, This community is
keeping the mat open and areas inaccessible to grazing though presence of P. fontana only recorded from
maintaining diversity. as this can destroy the reliable. Lower altitude the Trotternish Ridge
bryophyte mat. examples often lack on Skye. It is described
Saxifraga stellaris and from just 4 samples.
grade into the Cratoneuron
commutatum springs,
M37, M38.

15
 Fig 7. Key to vegetation dominated by sedges, Carex and Eriophorum spp., in which Sphagnum spp. are absent or infrequent in the sward

16
1. Vegetation 1a. Vegetation dominated
overwhelmingly dominated by Carex spp. Eriophorum
by Eriophorum spp; Carex angustifolium can be
spp. never prominent. constant but never dominant.
Mires Figure 7

2. Eriophorum angustifolium 2a. Eriophorum vaginatum 3. Carex rostrata prominent 3a. Open vegetation
dominant on open eroded dominant and usually in herb-rich swards over dominated by short sedges
peat, especially around tussocky. Often extensive bulky mosses, often forming including at least three of:
haggs. mire. floating rafts. Constants Carex panicea, C. demissa,
include: Eriophorum C. dioica, C. lepidocarpa,
angustifolium, Galium C. pulicaris, C. saxatilis,
palustre, Menyanthes C. echinata, Drepanocladus
trifoliata, Potentilla palustris revolvens.
and Calliergon cuspidatum.
4. Carex saxatilis dominant, 4a. Carex saxatilis
other sedges subordinate. absent. Frequent species:
Constants include: Carex panicea, C. pulicaris,
Thalictrum alpinum, Drepanocladus revolvens,
Hylocomium splendens, Pinguicula vulgaris, Aneura
Polygonum viviparum pinguis, Campylium
and Aneura pinguis. stellatum and Selaginella
selaginoides.

5. Saxifraga aizoides 5a. Two or more of

and Carex demissa constant. Carex dioica, C. pan-
icea, C. pulicaris and
C. lepidocarpa con-
stant. Saxifraga
aizoides at low
frequency.

M3 Eriophorum M20 Eriophorum vaginatum M9 Carex rostrata – Calliergon M12 Carex saxatilis mire M11 Carex demissa – M10 Carex dioica –
angustifolium bog pool blanket and raised mire cuspidatum/giganteum mire Saxifraga aizoides mire Pinguicula vulgaris
mire

This community usually Often occurs in complex Montane flushes over Carex dioica may be
results from degraded blan- mosaics with M5 and the 700 m asl. absent or difficult to
ket mire, M19. Species-poor, S27 swamp in which the find in poorer stands.
but a range of transitional diagnostic herbs are very C. demissa may
M20-M19 stands occurs. similar, though lacking the replace C. lepidocarpa
moss carpet of M9. in higher altitude/less
base-rich stands.
 Fig 8. Key to vegetation in which Molinia caerulea is constant and structurally important but Sphagnum spp. are generally lacking

1. Schoenus nigricans 1a. Schoenus nigricans

constant and abundant. absent or rare, though may
be locally frequent in
flushed wet heath.
Mires Figure 8

2. Juncus subnodulosus 2a. Schoenus nigricans 3. Calluna vulgaris 3a. Calluna vulgaris
constant, usually with usually abundant with some and Erica tetralix constant absent or present at low
other Juncus spp., Succisa of: Erica tetralix, Narthecium and usually abundant with frequency. Erica tetralix
pratensis, Potentilla erecta, ossifragum, Anagallis Scirpus cespitosus and may be frequent in the M25a
Carex panicea and tenella, Drosera rotundifolia Potentilla erecta. sub-community
Calliergon cuspidatum. and generally some
Sphagnum auriculatum
and/or S. subnitens.
J. subnodulosus absent.
4. Molinia caerulea – 4a. Cirsium dissectum
dominated fen with constant and Juncus subnodulosus
Cirsium dissectum, usually absent or rare.
with Juncus subnodulosus,
sometimes J. articulatus
or J. inflexus. Associates 5. Rich Molinia caerulea 5a. Molinia caerulea
include poor fen herbs, fen with Carex nigra and at and Potentilla erecta
particularly Valeriana dioica least three of: Crepis only constants in gen-
and Succusa pratensis. paludosa, Carex panicea, C. erally quite poor vege-
pulicaris, Valeriana dioica, tation. Associates can
Succisa pratensis, include frequent
Filipendula ulmaria Juncus acutiflorus
and Ranunculus acris. and sparse herbs such
as Succisa pratensis,
Cirsium palustre
and/or Erica tetralix.

M13 Schoenus nigricans – M14 Schoenus nigricans – M15 Scirpus cespitosus – M24 Molinia caerulea – M26 Molinia caerulea – M25 Molinia caerulea
Juncus subnodulosus mire Narthecium ossifragum mire Erica tetralix wet heath Cirsium dissectum fen-meadow Crepis paludosa mire – Potentilla erecta mire

This community is mainly Existing records limit this

in south and east Britain, community to the North
though sedge-rich stands of Pennines and Lake District,
M25 found in south-west though it may extend into
Scotland and the Western the Southern Uplands.
Isles may resemble the
Juncus acutiflorus – Erica
tetralix sub-community M24c.

17
18
Fig 9. Key to Juncus spp. and tall herb dominated fens in which Sphagnum spp. are never prominent

1. Iris pseudacorus constant 1a. Iris pseudacorus absent

and generally dominant, or at low frequency in the
usually with constant sward.
Oenanthe crocata, Filipendula
Mires Figure 9

ulmaria and Poa trivialis.

2. Filipendula ulmaria 2a. Filipendula ulmaria can

constant and prominent be occasional to frequent but
with tall fen herbs including never prominent. Vegetation
Angelica sylvestris, dominated by tall rushes
Valeriana officinalis usually with constant Holcus
and Rumex acetosa. lanatus and Lotus uliginosus.

3. Fen-meadow dominated 3a. Vegetation dominated by

by Juncus subnodulosus Juncus effusus and/or J. acu-
often with frequent J. tiflorus with constant Galium
inflexus and/or J. articulatus. palustre and other poor-fen
Constants include: Mentha associates, most usually
aquatica, Cirsium palustre Cirsium palustre, Rumex
and Calliergon cuspidatum. acetosa, Epilobium palustre,
Mentha aquatica and
Ranunculus flammula.

M28 Iris pseudacorus – M27 Filipendula ulmaria – M22 Juncus subnodulosus – M23 Juncus effusus/
Filipendula ulmaria mire Angelica sylvestris mire Cirsium palustre fen-meadow acutiflorus – Galium palustre
rush-pasture

Oenanthe crocata is often In western Britain Molinia

absent from stands of this caerulea can be locally abun-
community in the north and dant in this community.
west of Scotland.
3 Mire community descriptions
and sub-community keys
M1 Sphagnum auriculatum in flowering. Around the pool margins Molinia
caerulea can extend down from the mire surface
bog pool community although its cover is generally low.
This community is confined to pools and wetter
This bog pool community typically consists of hollows on ombrogenous and topogenous mires
floating masses or soft wet carpets of Sphagnum with base-poor and oligotrophic raw peat soils in
spp., mainly Sphagnum auriculatum (including the more oceanic parts of Britain. It is a widespread
var. inundatum) and S. cuspidatum, with scat- component in the Scirpus cespitosus – Eriophorum
tered vascular plants growing on or through them vaginatum blanket mire (M17) in the far west of
or in areas of open water between. Locally, the Britain including western Scotland, parts of the
bright orange-yellow S. pulchrum is conspicuous. Lake District, Wales, and the South-West Peninsula,
S. recurvum is rare in contrast with the Sphagnum and the Narthecium ossifragum – Sphagnum papil-
cuspidatum/recurvum bog pool community (M2). losum valley mire (M21) in south-western valley
Other bryophytes are generally scarce, but mires with a high water table, particularly in the
Cladopodiella fluitans is characteristic at low New Forest and Dorset.
frequencies and Gymnocolea inflata can also be The wetness gives some protection to this
present. vegetation where mires are grazed or burned, but
The commonest vascular plants are Meny- it has been reduced on many sites by draining and
anthes trifoliata and Eriophorum angustifolium cutting of the peat. It has been widely lost where
which together make up a cover of less than 30%. Erico-Sphagnion communities have been con-
In open water Sphagnum cover is reduced and verted to Ericion heaths or their degraded deriva-
Utricularia species, usually U. minor or locally tives. Shallow peat-digging can create flooded hol-
U. intermedia, are sometimes present. In shallow lows which become suitable for recolonisation by
water Rhynchospora alba is characteristic, and R. Sphagnum spp., Rhynchospora alba and Drosera
fusca is found occasionally in this community. spp., but such locally reconstituted stands often
Narthecium ossifragum and Drosera spp., particu- lie in much-modified mire contexts.
larly D. rotundifolia, are also occasionally present.
In some areas Carex limosa is frequent, but is shy No sub-communities.

19
M2 Sphagnum cuspidatum/ community. There may be some sedges including
Carex limosa, C. curta and C. magellanica.
recurvum bog pool community The community is typically found in pools and
lawns on very wet and base-poor raw peats on
This community is typically dominated by soft ombrogenous and topogenous mires in the less
wet carpets of Sphagnum cuspidatum or oceanic parts of Britain. Its range coincides
S. recurvum, or both. S. pulchrum occurs very closely with that of the Erica tetralix – Sphagnum
locally, occasionally with S. tenellum, S. magel- papillosum mire (M18) and it typically forms the
lanicum or S. papillosum. Sphagnum auriculatum pool, wet hollow and lawn elements in that
is rare in contrast with the Sphagnum auricula- community (and its degraded derivatives) on
tum bog pool community (M1). Other bryo- lowland raised bogs, on locally raised areas
phytes are scarce but Polytrichum commune or within low altitude blanket mires and in base-poor
Aulacomnium palustre can form occasional basin mires. It occurs from Wales up through the
patches and there may be scattered leafy hepatics. Scottish Borders and south-west Scotland with
Vascular plants occur as scattered individuals some localities in north-east Scotland.
with Eriophorum angustifolium and Erica tetralix This community has been reduced by wide-
both constant; the former often extending into spread drainage and cutting of mires, so that often
deeper pools and the latter confined to drier areas. just small and modified fragments remain within
Drosera rotundifolia is frequent and Narthecium predominantly agriculture landscapes. However
ossifragum occasional. Andromeda polifolia, this community readily colonises shallow flooded
where present, is distinctive of this vegetation workings and appears to have expanded its cover-
type particularly around pool margins, and togeth- age in sites where there has been some agricultural
er with Rhynchospora alba it forms a clear sub- enrichment of the water.

M2

Sphagnum recurvum usually Sphagnum recurvum constant

absent but Rhynchospora alba and often abundant with no
and Andromeda polifolia very Rhynchospora alba and only
frequent. occasional Andromeda polifolia.

M2a M2b
Rhynchospora alba
sub-community Sphagnum recurvum/
cuspidatum sub-community
Sphagnum cuspidatum is
the typical dominant in the Sphagnum recurvum is
Sphagnum carpet. Eriophorum constant with S. cuspidatum
angustifolium and Erica tetralix and often the more abundant
join Rhynchospora alba and of the two. Eriophorum angusti-
Andromeda polifolia as con- folium, Erica tetralix and
stants, Drosera rotundifolia is Drosera rotundifolia maintain
frequent and D. anglica or their high frequency and
D. intermedia occasional. Vaccinium oxycoccos is a
good preferential. Polytrichum
This sub-community is more commune and Aulacomnium
widely distributed on active, palustre can occur sporadically
undisturbed raised mires. in slightly drier areas with
some Calluna vulgaris and
Eriophorum vaginatum.
Molinia caerulea may be
locally prominent.
This sub-community is more
restricted to soligenous areas
and disturbed basin mires.

20
M3 Eriophorum angustifolium This community is typically found as small stands
on barer exposures of acid raw peat soils in
bog pool community depressions, erosion channels or shallow peat
cuttings on a wide range of mire types. It can be
Eriophorum angustifolium is dominant here in found in natural hollows on surfaces of more or
swards where other vascular species and less intact mires but is more common among
Sphagnum spp. play a relatively minor role. Its erosion features where the peat has been worn
shoot density is very variable, as is the sward height down in gullies or redistributed. It is also some-
which may be very short or up to half a metre or times associated with abandoned peat workings
more. Usually shoots reach approximately 30 cm. on lowland mires. The community is particularly
Other vascular plants attain only occasional associated with the eroded blanket mire in the
frequency but there can be scattered small tus- north-west of Britain, being a common feature in
socks of Eriophorum vaginatum or Molinia tracts of the Calluna vulgaris – Eriophorum vagi-
caerulea or sparse individuals of Drosera rotundi- natum and Eriophorum vaginatum mires (M19
folia, Erica tetralix or Empetrum nigrum ssp. and M20), and it is widespread but local in low-
nigrum. Bryophyte cover is also very variable and land Erico-Sphagnion mires (M18, M21) and
there are no constant species but Drepanocladus Ericion wet heaths (M15, M16). This community
fluitans may be frequent, often growing sub- may represent a seral stage in the redevelopment
merged. Sparse shoots or small tufts of Sphagnum of active mire vegetation following disruption.
spp. may be present, usually S. cuspidatum but
sometimes S. recurvum or S. papillosum. No sub-communities.

21
M4 Carex rostrata – abundant as stoloniferous mats), Molinia caerulea,
Potentilla erecta, Galium palustre, Rumex ace-
Sphagnum recurvum mire tosa, Viola palustris, Succisa pratensis and
Stellaria alsine. Usually only one or two of these
This mire typically has a cover of sedges over a are present in any one stand. Potentilla palustris,
carpet of semi-aquatic Sphagnum spp. Carex ros- Menyanthes trifoliata and Equisetum fluviatile
trata is the commonest sedge, usually forming a also may occur occasionally.
rather open cover of shoots, but it can be accom- This community is characteristic of pools and
panied by C. curta, C. lasiocarpa, C. limosa or seepage areas on raw peat soils of topogenous and
C. nigra (the first two especially can be locally soligenous mires where the waters are fairly acid
prominent). Carex chordorrhiza is a rare associate. and only slightly enriched. It can occur in bog
Occasionally the taller element of the vegetation pools on the surface of basin (and sometimes
also has Eriophorum angustifolium, Juncus raised) mires, but is more common in obviously
effusus or J. acutiflorus. There is generally an soligenous areas as in mire laggs and the wettest
extensive wet carpet of Sphagnum spp. parts of water-tracks. Enrichment is slight and the
S. recurvum and S. cuspidatum are usually the pH is typically around 4. The community is of
most frequent and abundant species and S. auric- widespread but local occurrence throughout the
ulatum is also common. Sphagnum palustre is north-west of Britain and probably remains as
occasional, with sparse records for Sphagnum remnants in drained mire systems in the lowlands.
subnitens and S. papillosum. S. squarrosum and The place of this community in the terrestrial-
S. teres are characteristically rare, which provides ising succession is not clear and the vegetation
a good contrast with Carex rostrata – Sphagnum may be very stable provided the high water table
squarrosum mire (M5). Other bryophytes are few, and modest irrigation are maintained. Drainage
but Polytrichum commune is very frequent form- results in the demise of the more aquatic
ing scattered patches. Aulacomnium palustre and Sphagnum spp. and perhaps a transition to
Calliergon stramineum are very sparse. the Carex echinata – Sphagnum recurvum/ auric-
Scattered through the ground cover are indi- ulatum mire (M6), and with grazing, may result in
viduals of an impoverished poor-fen herb flora. a spread of Juncus dominance.
The commonest species are Agrostis canina ssp.
canina and A. stolonifera (which may be locally No sub-communities.

22
M5 Carex rostrata – Sphagnum latum are occasionally found. Sphagnum contor-
tum is rare in contrast with the Carex rostrata –
squarrosum mire Calliergon cuspidatum/giganteum mire (M9).
Other common bryophytes are Aulacomnium
This mire is fairly heterogeneous and is charac- palustre and Calliergon stramineum.
terised overall by the dominance of sedges with This mire is typically found as a floating raft or
scattered poor-fen herbs over a patchy carpet of on soft, spongy peats in topogenous mires and in
moderately base-tolerant Sphagnum spp. The soligenous sites with mildly acid, only moderate-
commonest species throughout are Carex rostrata ly calcareous and rather nutrient-poor waters; the
and C. nigra, with the former generally more pH range is from about 4 to above 6. It is charac-
extensive. Carex lasiocarpa can be locally promi- teristically found in zonations and mosaics, the
nent and C. curta is occasionally found. Carex simplest being open water transitions around
limosa and C. diandra are typically absent in con- lakes. It can also be found around springs, seepage
trast with the Carex rostrata – Calliergon cuspida- lines and streams where it can form part of a mix-
tum/giganteum mire (M9). ture of poor- and rich-fen communities. The com-
Other vascular plants are often limited to scat- munity has a widespread but fairly local distribu-
tered individuals, but the most frequent overall tion in north-western parts of Britain. It was prob-
are Potentilla palustris, Eriophorum angustifoli- ably once much more widespread in the lowland
um, Menyanthes trifoliata, Galium palustre and south and east where relic stands may still occur.
such typical poor-fen herbs as Succisa pratensis, The peat under this community is often very
Viola palustris, Ranunculus flammula, Epilobium soft which gives the vegetation a measure of pro-
palustre and Lychnis flos-cuculi. Juncus effusus tection against the trampling and grazing effects of
can be frequent, as can Molinia caerulea and larger herbivores, although damage may occur
Myrica gale. during periodic dry spells. Where the community
The bryophyte carpet helps define the Carex – runs onto firmer peats around the margins of
Sphagnum squarrosum mire against closely relat- lakes or basins, the vegetation tends to pass to the
ed vegetation types. Sphagnum spp. are at least Carex echinata – Sphagnum recurvum/auricula-
patchily prominent. Especially distinctive is the tum mire (M6). The effect of grazing on these tran-
presence of Sphagnum squarrosum and S. teres. In sitions may favour the spread of Juncus effusus.
addition S. recurvum and S. palustre are frequent-
ly encountered and S. cuspidatum and S. auricu- No sub-communities.

23
M6 Carex echinata – Sphagnum ossifragum, Drosera rotundifolia and Erica tetralix
recurvum/auriculatum mire are found. The rushes Juncus acutiflorus and
J. effusus may each be dominant in particular sub-
This community has a distinct general character communities.
but includes a wide variation in composition, A ground carpet of Sphagnum spp. is promi-
expressed here in four sub-communities. Ess- nent and it is most frequently composed of
entially it is a poor-fen with small sedges or rush- S. recurvum and S. auriculatum, with occasional
es dominating over a carpet of oligotrophic and occurrence of S. subnitens and S. papillosum.
base-intolerant Sphagnum spp. The constants are There are only a few other commonly occurring
very few. Among vascular plants only Carex echi- bryophyte species. Polytrichum commune is very
nata has a uniformly high frequency, but frequent, Rhytidiadelphus squarrosus is occasion-
C. nigra and C. panicea are common, and C.demis- al and Calliergon stramineum and Aulacomnium
sa occasional. There are two negative characters palustre are patchy throughout. Calliergon cuspi-
which aid definition of this community. Firstly, datum and Plagiothecium undulatum are conspic-
the general absence of more calcicolous Carex uously rare.
species, e.g. C. dioica, C. pulicaris, C. lepidocarpa This mire is the major soligenous community
and C. flacca, helps to separate this community of peats and peaty gleys irrigated by rather base-
from the Caricion davallianae rich fens (M9 to poor waters in the sub-montane zone of northern
M12), and secondly, only local occurrence of and western Britain. The soils and water are quite
species like C. rostrata and C. curta marks the veg- acidic with a superficial pH usually between 4.5
etation off from communities like the Carex ros- and 5. It typically occurs as small stands among
trata – Sphagnum recurvum mire (M4). other mire communities, grassland and heaths and
The most common vascular associates are sometimes with swamp and spring vegetation. It is
grasses and poor-fen dicotyledons. Among the commonly found in tracts of unenclosed pasture
grasses, Agrostis canina ssp. canina and Molinia on upland fringes, particularly between 200 m and
caerulea are the most common but Anthoxanthum 400 m (although it may be found much higher)
odoratum is also frequent. Commonly occurring and is ubiquitous in the upland fringes of Britain.
poor-fen dicotyledons include Viola palustris and The community is frequently grazed. This, espe-
Potentilla erecta, and occasionally one or more of cially where combined with drainage, can convert
Galium saxatile, G. palustre, Cirsium palustre, the community to grassland. The exclusion of her-
Epilobium palustre, Succisa pratensis, Ranun- bivores would be expected to permit progress to
culus flammula or Cardamine pratensis may be wet scrub and woodland, although in many cases
present. Sometimes species such as Narthecium this would probably be slow and patchy.

24
 M6

Mixtures of Carex Sedge cover less

echinata, C. nigra and varied and extensive
C. panicea usually domi- and Eriophorum
nate with Eriophorum angustifolium usually
angustifolium sometimes sparse; Juncus effusus
abundant but Juncus spp. or J. acutiflorus
at most sparse. dominant.

Carex echinata Carex nigra,

usually the most C. panicea and
abundant sedge, C. echinata equally Juncus effusus is Juncus acutiflorus
C. panicea and frequent and the most abundant usually totally
C. nigra subordi- mixtures of these rush among an replacing J. effusus
nate and Nardus usually dominate. extensive with Molinia
stricta and Juncus Nardus stricta and Sphagnum carpet caerulea frequent
squarrosus occa- Juncus squarrosus usually dominated and a generally
sional in an exten- very common by S. recurvum more varied
sive Sphagnum among a sometimes with S. palustre. Sphagnum carpet.
carpet. patchy Sphagnum
cover.

M6a M6b M6c M6d

Carex echinata sub-community Carex nigra – Nardus stricta Juncus effusus sub-community Juncus acutiflorus sub-community
sub-community
The vegetation is generally dominated Sedges are less frequent and abundant Juncus acutiflorus is dominant over an
by mixtures of sedges with Carex echina- Together with the Carex spp. and other and the vegetation is dominated by extensive Sphagnum carpet and Molinia
ta generally most common; grasses are species mentioned above, Eriophorum Juncus effusus. Frequent vascular caerulea becomes more consistently
rather scarce except for Molinia caerulea angustifolium is very common and associates are few but there is often some frequent. Most of the same poor-fen herbs
and Agrostis canina ssp. canina which Anthoxanthum odoratum frequent. Agrostis canina ssp. canina, Potentilla as listed in M6c are frequent. Sphagnum
are very common. Eriophorum angusti- Juncus squarrosus is a good preferential, erecta, and (rather diagnostic here) spp. are generally abundant with
folium is frequent and may dominate. but J. effusus is scarce and J. acutiflorus Galium saxatile. Carex echinata, Molinia S. palustre common throughout and
Rushes are typically scarce and of low absent. In some stands Sphagnum caerulea and Viola palustris are also S. recurvum or S. auriculatum with
cover. The Sphagnum carpet is typically recurvum and S. palustre are the fairly common. The Sphagnum carpet S. subnitens, S. papillosum and S. capilli-
extensive and luxuriant. S. palustre is commonest Sphagnum species with poor is generally extensive and luxuriant and folium having dominance in the carpet.
common but S. recurvum and S. auricula- fen herbs such Ranunculus flammula, S. recurvum is almost always dominant.
tum show a pattern of replacement, the Epilobium palustre and Cirsium palustre. Polytrichum commune remains frequent This sub-community is found throughout
latter becoming more prominent in the In contrast other stands have a greater and sometimes abundant. the range of M6.
oceanic far west of Britain. Drosera abundance of Sphagnum spp., with
rotundifolia, Narthecium ossifragum, S. auriculatum, S. subnitens and S. This sub-community is found throughout Two variants.
Erica tetralix, Juncus bulbosus/kochii and papillosum being preferential and often the range of M6.
Menyanthes trifoliata show some prefer- some Molinia caerulea present among
ence for the S. auriculatum type of flush. the other grasses. Two variants.

This sub-community is found throughout This sub-community is found throughout

the range of M6. the range of M6.

25
Two variants. Two variants.
M7 Carex curta – Sphagnum mune, Calliergon stramineum or C. sarmentosum.
russowii mire Grasses play a minor role, although Nardus stricta
is very common and Agrostis canina ssp. canina
This mire community has prominent cyperaceous frequent. Among dicotyledons Viola palustris and
and Sphagnum components with a distinct north- Galium saxatile are most common, but are typi-
ern and montane character. Eriophorum angusti- cally of low cover.
folium and Carex echinata are very frequent and This community is confined to high altitude
provide a floristic link with the Carex echinata – sites, usually above 650 m, forming small stands
Sphagnum recurvum/auriculatum mire (M6) com- where peaty soils are irrigated by oligotrophic and
munity which occurs at lower altitudes, but in base-poor waters. It is characteristic of hollows
contrast, C. curta is a constant often with high and drainage channels in blanket mires or flushes
cover. It is often accompanied by C. bigelowii or and seepage areas in tracts of montane moss
C. aquatilis and C. rariflora. Carex nigra can also heaths. It is an altitudinal replacement for Carex
occur, sometimes abundantly. Larger Juncus spp., echinata – Sphagnum recurvum/auriculatum
e.g. Juncus effusus and J. acutiflorus, are very mire (M6) with a preponderance of montane
scarce, again in contrast to M6. plants. The community is mainly confined to the
The Sphagnum carpet is typically extensive. central Highlands of Scotland, but extends south
Sphagnum papillosum is common and often into the Pennines and perhaps also into Wales.
abundant, and S. subnitens, S. auriculatum, Most of the occurrences of the Carex curta –
S. capillifolium or S. recurvum may be frequent. Sphagnum russowii mire are close to or above the
The high altitude species S. russowii is constant as potential forest limit in the Scottish Highlands
is S. lindbergii in one of the sub-communities. and the community is probably an essentially sta-
The rare S. riparium also grows in this community. ble component of the vegetation pattern under
Other frequent bryophytes are Polytrichum com- present-day conditions.

M7

Sphagnum lindbergii very Sphagnum lindbergii absent with

frequent and usually co-domi- S. recurvum abundant among
nant with S. papillosum.Carex frequent S. papillosum and
echinata usually co-dominant S. russowii. Carex bigelowii very
with C. bigelowii in the vascular scarce and C. echinata usually
tier with Nardus stricta dominant.
common.

M7a M7b

Carex bigelowii – Sphagnum lindbergii sub-community Carex aquatilis – Sphagnum recurvum sub-community

Carex curta can be common with C. echinata and C. Carex curta is often co-dominant with C. echinata. Some
bigelowii but other sedges are scarce. Among the few stands have the rare C. aquatilis and C. rariflora. C. nigra
dicotyledons Saxifraga stellaris is preferential. As well as is more common in this sub-community and can be locally
the Sphagnum spp. mentioned above, S. subnitens, S. auric- dominant. There is also more diversity among vascular
ulatum and S. capillifolium are frequent and S. recurvum associates with Festuca vivipara, Agrostis stolonifera,
scarce. Among other bryophytes Polytrichum commune is Deschampsia flexuosa, Luzula multiflora and Galium sax-
frequent, but more distinctive are Calliergon sarmentosum, atile all preferential. Nardus stricta is only occasional. Only
Drepanocladus exannulatus, Polytrichum alpestre and the Sphagnum spp. mentioned above are frequent. Other
P. alpinum. bryophytes are few, but Polytrichum commune is better
represented and Calliergon stramineum replaces
This sub-community is found throughout the range of M7. C. sarmentosum.

This sub-community is more local being concentrated

around the Clova-Caenlochan area of the east Highlands.

26
M8 Carex rostrata – Sphagnum Selaginella selaginoides. Common grasses are
warnstorfii mire Festuca ovina (and F. vivipara), Nardus stricta,
Anthoxanthum odoratum and Agrostis stolonifera,
This community has a dominant cover of sedges all of which are generally present as scattered
over an extensive carpet of Sphagnum spp. and a shoots or small tussocks.
fairly numerous and diverse assemblage of herbs. This community typically occurs as small
Carex rostrata and C. nigra are the commonest stands and is strictly confined to raw peat soils in
sedges, the former usually more abundant and of waterlogged hollows in the montane zone of
high cover. Other poor-fen sedges, C. panicea, Britain where there is moderate base-enrichment
C. echinata and C. demissa occur frequently by drainage from calcareous rocks. These condi-
and sometimes abundantly and C. pulicaris is tions are not common. The peat deposits on which
occasional. the community is found are typically quite deep,
The Sphagnum carpet is typically extensive, usually more than 1 m, with a high and stagnant
and the prominence of the base-tolerant water table. The pH of the waters and the peat is
Sphagnum teres and S. warnstorfii is quite dis- usually between 5.5 and 6. The small but distinct
tinctive. Sphagnum recurvum occurs frequently montane element in the flora of this community
and S. subsecundum sensu stricto, although only helps separate it from the Carex rostrata –
occurring occasionally, is also very characteristic Sphagnum squarrosum mire (M5) which is found
of this community. in analogous situations in the lowlands. The com-
Other bryophytes are numerous and in particu- munity is generally confined to altitudes between
lar Aulacomnium palustre and Rhizomnium 400 m and 800 m in the central Highlands except
pseudopunctatum are frequent. Also distinctive for a few examples in southern Scotland and
are Calliergon cuspidatum, C. stramineum and northern England.
less frequently C. sarmentosum. Homalothecium The frequent presence of seedlings of Salix
nitens is quite common and a good diagnostic aurita in stands of the community may indicate a
species. Hylocomium splendens and Rhytidiad- tendency towards the development of montane
elphus squarrosus are frequently recorded. willow scrub but such successions have never
Herbaceous associates are common but typically been seen to progress further.
of low total cover. Constant species are Epilobium
palustre, Potentilla erecta, Viola palustris and No sub-communities.

27
M9 Carex rostrata – Calliergon the larger Mniaceae are also common. Campylium
stellatum is a distinctive species of this communi-
cuspidatum/giganteum mire ty together with Scorpidium scorpioides and
Drepanocladus revolvens.
This community has a diverse composition and This community is characteristic of soft,
physiognomy, even within individual stands, but spongy peats kept permanently moist by at least
is generally characterised by a fairly rich moderately base-rich and calcareous waters.
assemblage of sedges and vascular plants over a Waters and substrates always have a pH above 5
carpet of bulky mosses and localised patches of and usually above 6. It is commonest in wetter
Sphagnum spp. The commonest large sedge is parts of topogenous mires in hollows or old peat-
Carex rostrata, which is often abundant and workings, but also around springs, laggs of raised
sometimes dominant. Carex diandra is frequent, mires and mowing marshes. The community is
and C. lasiocarpa may accompany one or both of widespread but local, ranging from south-east
these species. Carex paniculata or C. appropin- England, particularly in Broadland, to Wales and
quata is present in some localities. Small sedges northern England and through most of Scotland.
which commonly occur in this community The community is limited by the fairly sparse
include C. panicea and C. nigra. occurrence of suitable natural situations and in
Intermixed with these species, or fringing the lowland south-east by wetland drainage and
patches of them, are a variety of associates. the cessation of shallow peat-digging. It is
Potentilla palustris and Menyanthes trifoliata are typically too wet to be grazed but in some areas it
common and are particularly important when the occurs within mowing marsh that is periodically
vegetation forms floating rafts. Also common are cropped. Throughout its range, at least at the
Eriophorum angustifolium, Equisetum fluviatile, lower altitudes and in less remote sites, the Carex
E. palustre, Succisa pratensis, Pedicularis rostrata – Calliergon cuspidatum/giganteum mire
palustris, Cirsium palustre and Ranunculus is probably a successional stage to Salix pentan-
flammula often with Galium palustre. Less dra – Carex rostrata woodland (W3) in the north
evenly distributed and usually present as and Salix cinerea – Betula pubescens – Phragmites
scattered individuals are Mentha aquatica, Caltha australis woodland (W2) in the south-east,
palustris, Valeriana dioica, Angelica sylvestris, although development to woodland may be hin-
Epilobium palustre and Lychnis flos-cuculi. The dered by a high water table or by the periodic
commonest grass to occur in this community is mowing of the vegetation. There is also the possi-
Molinia caerulea, particularly in drier stands. bility that under certain conditions this mire type
Bryophytes are almost always conspicuous. is seral to the development of poor-fen and
Calliergon cuspidatum is constant and C. gigan- ombrogenous mire through the local formation of
teum and C. cordifolium frequent. One or more of Sphagnum nuclei.

28
 M9

Carex rostrata usually dominant, Carex rostrata and C. diandra

sometimes with C. lasiocarpa or constant, either or both dominant
more locally C. diandra or with or without C. lasiocarpa.
Schoenus nigricans. Calliergon Herbaceous associates often
cuspidatum common but other numerous and lush. Campylium
Calliergon species and larger stellatum only occasional and
Mniaceae at most local. Ground Scorpidium scorpioides and
carpet usually dominated by Drepanocladus revolvens rare,
mixtures of Campylium stellatum, but Calliergon cordifolium and
Scorpidium scorpioides and C. giganteum common, often
Drepanocladus revolvens. with large Mniaceae.

M9a M9b

Campylium stellatum – Scorpidium scorpioides Carex diandra – Calliergon giganteum sub-community

sub-community
The pattern of dominance is more variable here as indicated
Smaller sedges are more numerous than in M9b; Carex pan- above. Juncus subnodulosus is locally abundant in eastern
icea and C. nigra retain their high frequency and C. limosa England. Herbaceous associates are more numerous;
and C. echinata are strongly preferential. Herbaceous associ- Potentilla palustris, Menyanthes trifoliata and Filipendula
ates are variable and not very rich, and total cover is low, ulmaria can be prominent with Angelica sylvestris,
giving an open community. Menyanthes trifoliata and Epilobium palustre, Lychnis flos-cuculi, Valeriana dioica,
Potentilla palustris can be prominent, but most species Caltha palustris, Cardamine pratensis and Mentha aquatica
occur as scattered plants. Bryophytes are usually prominent all frequent. Bryophytes are often extensive with Calliergon
as indicated above. Clumps of Sphagnum spp. are occasion- giganteum, C. cordifolium, Plagiomnium rostratum and
al, notably so for the base-tolerant Sphagnum contortum. P. affine showing their maximum development in this
sub-community.
This sub-community is largely north-western in range.
This sub-community occurs throughout the range mostly in
topogenous mires.

29
M10 Carex dioica – Pinguicula calcicolous species provide a strong definition for
vulgaris mire the community against its counterparts in the
Caricion nigrae (M5-M7) and in flushed Oxycocco
The community includes a range of distinctive – Sphagnetea mires (M14-M21) which occupy
calcicolous flush vegetation in which the bulk of similar but more base-poor situations.
the sward is composed of small sedges, dicotyle- This community is typically a soligenous mire
dons and bryophytes. There are marked variations of mineral soils and shallow peats kept very wet
in proportions of frequent species, as indicated in by base-rich, calcareous and oligotrophic waters.
the three sub-communities and their variants. The pH of flushing waters is high, usually
Essentially this is a small sedge mire with Carex between 5.5 and 7.0 or sometimes higher, and the
dioica, C. hostiana, C. lepidocarpa, C. panicea and composition of this community is one of the most
C. pulicaris as constants which are also often calcicolous of British mires. It is found in small
abundant. Carex nigra is frequent and C. echinata stands, often associated with spring and rill
and C. flacca common. Other prominent Cyper- vegetation, within grasslands and more occasion-
aceae are Eriophorum angustifolium, a constant, ally in ombrogenous mires and around topoge-
and Eleocharis quinqueflora. nous mires. Typically the in situ formation of peat
Some rushes and grasses occur frequently. is limited, a feature which helps to distinguish the
Juncus articulatus is a constant and J. bulbosus/ habitat of the community from that of base-rich
kochii is typical of less base-rich sites. Molinia basins where it is replaced by vegetation like the
caerulea is the commonest grass with Festuca Carex rostrata – Calliergon cuspidatum/giganteum
ovina more variable in occurrence. Other herbs mire (M9) and the Carex rostrata – Sphagnum
generally occur as scattered plants. The frequent warnstorfii mire (M8). It is predominantly a com-
occurrences of Pinguicula vulgaris and Selaginella munity of north-west Britain from Wales and the
selaginoides are very distinctive of this communi- Pennines northwards, developed in the cool, wet
ty. Potentilla erecta and Succisa pratensis are climate.
also common species in this community but The community typically occurs in unenclosed
Equisetum palustre and Euphrasia officinalis agg. uplands and most of the stands are grazed and
are more variable in occurrence. Other species are trampled by large herbivores. It is probably these
characteristic of particular sub-communities. factors, combined with nutrient impoverishment
Bryophytes are always obvious, often compris- and the often strong and scouring effect of the
ing 50% or more of the ground cover. Calcicolous irrigation, which play a major part in maintaining
species such as Campylium stellatum, Aneura the community in its generally rich, varied and
pinguis, Drepanocladus revolvens, Ctenidium open state. Most stands would probably progress
molluscum, Fissidens adianthoides and Craton- to Alno – Ulmion scrub or woodland (W7, W9) if
euron commutatum are frequent, together with grazing were withdrawn. However, at higher
Bryum pseudotriquetrum. Such assemblages of altitudes the vegetation may be a climatic climax.

30
 M10

Carex hostiana, C. pulicaris and Species listed opposite scarce

C. nigra all frequent with or absent. Gymnostomum
Eriophorum angustifolium and recurvirostrum or less commonly
Molinia caerulea. Potentilla Catascopium nigritum forming
erecta and Succisa pratensis hummocks with Drepanocladus
common with Ctenidium revolvens and Cratoneuron
molluscum and Fissidens commutatum often abundant.
adianthoides frequent.

Carex demissa and Carex demissa and M10c

C. echinata occasional C. echinata scarce,
to frequent; C. lepido- but C. lepidocarpa, Gymnostomum recurvirostrum
carpa and C. flacca C. hostiana, C. pulicaris sub-community
scarce and C. pulicaris and C. flacca frequent.
patchy. Juncus bulbo- Juncus bulbosus and Vascular plants have low
sus and Erica tetralix Erica tetralix only of individual and total cover;
quite common. local significance. much more prominent are the
conspicuous moss hummocks,
particularly of Gymnostomum
recurvirostrum which are up to
M10a M10b 30 cm high and 60 cm across.
There is much bare ground with
Carex demissa – Juncus Briza media – Primula a fragmentary cover of vascular
bulbosus/kochii sub-community farinosa sub-community plants; good preferentials are
Plantago maritima, Sagina
This comprises the less Vascular plants are also nodosa and Minuartia verna.
calcicolous types of M10 with prominent but many swards M. stricta is restricted to this
vascular plants predominant. are open with extensive bare sub-community in its only
Carex panicea, C. dioica, ground. Calcicoles and more British locus.
C. hostiana and C. nigra remain mesophytic herbs well
very frequent with the species represented. Carex lepidocarpa, This striking vegetation is only
listed above. More calcicolous C. hostiana and C. pulicaris are recorded from Upper Teesdale.
herbs such as Briza media, consistently frequent and com-
Primula farinosa, Linum monly accompanied by C. flac-
catharticum and Sesleria ca. Among the preferentials
albicans are usually poorly Briza media, Primula farinosa,
represented. Bryophytes are Linum catharticum, Sesleria
generally less prominent in the albicans and Equisetum
turf; Bryum pseudotriquetrum, variegatumare frequent. Juncus
Fissidens adianthoides and bulbosus/ kochii, Erica tetralix,
Ctenidium molluscum are Narthecium ossifragum and
rather uncommon and Drosera rotundifolia are
Campylium stellatum and reduced in their occurrence.
Scorpidium scorpioides Among the bryophytes Aneura
rather more prominent. pinguis, Ctenidium molluscum
and Fissidens adianthoides are
This sub-community is largely consistently frequent.
restricted to Scotland and the
Lake District with outlying This sub-community is
occurrences in north-west predominantly found in
Wales and Upper Teesdale. northern England.

Three variants. Three variants.

31
M11 Carex demissa – Saxifraga very common at higher altitudes, and occasionally
Saxifraga stellaris, S. oppositifolia and Alchemilla
aizoides mire filicaulis ssp. filicaulis. Typically all these species
occur in an uneven and broken sward.
This vegetation is typically open with rich mix- Bryophytes are frequent and varied. Aneura
tures of small sedges, other herbs and bryophytes pinguis, Campylium stellatum, Drepanocladus
among water-scoured runnels and with much revolvens, Bryum pseudotriquetrum, and at lower
exposed silt and rock debris. There is a strong altitudes, Cratoneuron commutatum, Fissidens
floristic link with Carex dioica – Pinguicula vul- adianthoides, Ctenidium molluscum and Scor-
garis mire (M10) but the Arctic-Alpine element of pidium scorpioides are all common. The montane
the vegetation is much more pronounced in this moss Blindia acuta can be prominent at higher
community than it is in M10. Typically there is no altitudes and is a good preferential for this com-
single dominant. Carex demissa, C. panicea and munity.
C. pulicaris are very frequent throughout and This community is characteristic of open,
C. flacca and C. dioica are common in some vari- stony flushes, strongly irrigated with moderately
ants. Juncus articulatus is a constant, Eriophorum base-rich waters, on generally steep slopes in sub-
angustifolium is frequent as is Eleocharis quin- montane and montane parts of Britain. Although
queflora at lower altitudes. At higher altitudes the community can occur almost at sea level in the
Juncus triglumis is constant and Tofieldia pusilla far north-west of Scotland, it is generally confined
becomes frequent. By comparison with M10, to high altitudes. It is always associated with cal-
C. lepidocarpa and C. hostiana are much less com- careous bedrocks having a soil pH range of 5.5 to
mon and C. nigra and C. echinata also rather 7.0. Flushing is vigorous and erosion of the sur-
scarce. At higher altitudes there is an Arctic- face is therefore often pronounced and the soil
Alpine element with Juncus triglumis being con- cover little more than scoured accumulations of
stant and Tofieldia pusilla becoming more fre- silt and organic matter with exposed rock debris.
quent. Grasses are typically low in cover. Festuca It is largely confined to Scotland, but also present
ovina/vivipara is common and Agrostis stolonifera in the Lake District, and more locally in the
occasional. At higher altitudes Deschampsia Southern Uplands, the northern Pennines and
cespitosa (including D. alpina), Nardus stricta, north Wales.
Anthoxanthum odoratum, Agrostis canina ssp. The community is normally grazed and this
canina and Festuca rubra may be present. grazing may help maintain the open structure and
Other herbs found in this community include help prevent the development of a woody cover.
Pinguicula vulgaris and Saxifraga aizoides which However, colonisation of trees and scrubs would
are both constant and Selaginella selaginoides be slow due to the climatic conditions in which
which is very frequent. The montane element of this the community occurs and at higher altitudes the
community includes Thalictrum alpinum, which is mire is probably a climatic climax.

M11

Juncus triglumis and Thalictrum alpinum constant, Eleocharis quinqueflora constant with Juncus triglumis
but Eleocharis quinqueflora uncommon. Deschampsia and Thalictrum alpinum becoming more frequent at higher
cespitosa, Nardus stricta and Anthoxanthum odoratum altitudes. The grasses found in M11a at most occasional.
frequent and Alchemilla alpina occasional. Cratoneuron Cratoneuron commutatum and Scorpidium scorpioides very
commutatum, Scorpidium scorpioides and Fissidens common and abundant, often with Fissidens adianthoides.
adianthoides all scarce.
M11b
Cratoneuron commutatum – Eleocharis quinqueflora
sub-community
M11a
Thalictrum alpinum – Juncus triglumis sub-community In this sub-community M11 grades into M10 with more
extreme montane plants, except Saxifraga aizoides and
There is an obvious montane element in the vegetation as Blindia acuta, much more poorly represented; and in more
indicated in the species above. Saxifraga aizoides, Carex southerly stands even these become rare. Eleocharis quin-
demissa and C. panicea are usually the most abundant queflora is constant and sometimes abundant, rivalling the
vascular plants with Blindia acuta, Campylium stellatum or sedges, among which Carex hostiana and, in wetter stands,
Drepanocladus revolvens predominating among the mosses. C. rostrata are sometimes found. Vascular plant cover
typically more extensive than in M11a.
This is the typical form of M11 at higher altitudes and is
virtually confined to Scotland. This sub-community is also frequent in Scotland at lower
altitudes and in most of the English and Welsh stands.
Two variants.

32
 sional. Hylocomium splendens is also a constant
M12 Carex saxatilis mire as is Scapania undulata. There can also be some
small patches of Sphagnum spp.
Carex saxatilis is typically dominant in this This mire type is strictly confined to margins of
montane mire with a distinctive assemblage of high-montane flushes irrigated with base-rich and
associates. The sward is generally less than 20 cm calcareous waters perhaps influenced by long
high and rather open with patches of soil. Carex snow-lie. It typically occurs as small stands bor-
demissa, C. echinata and C. nigra are very fre- dering rills or more strongly irrigated soligenous
quent and can be abundant. Carex bigelowii is fair- mires. The soils that this community is found on,
ly consistent, especially in grassy transitions to though continuously irrigated, are not of especial-
surrounding swards. ly high pH, ranging from 4.6 to 6.3. The commu-
Eriophorum angustifolium is also frequent nity is fairly widespread but local on peaks above
attaining a cover of more than 10%. Almost all 750 m through the southern and central Scottish
other herbs occur as scattered individuals. Highlands with scattered localities in north-west
Selaginella selaginoides and Pinguicula vulgaris Scotland.
are both very common as in other calcicolous The physical effects of flushing, snow-melt,
flushes, but more distinctive are Thalictrum cryoturbation, and solifluctional flow result in the
alpinum, Polygonum viviparum, and Juncus trig- continual instability of the substrate on which this
lumis. Saxifraga aizoides is infrequent, in contrast community is found and this is important in
to Carex demissa – Saxifraga aizoides mire (M11). maintaining open stony areas where rare
Also common are the poor-fen herbs Viola palus- Arctic-Alpine sedges and rushes find a niche. It is
tris, Caltha palustris and Agrostis canina ssp. canina. possible that grazing prevents colonisation by
Bryophytes are an important element of the Arctic-Alpine willows; however, in the extreme
vegetation although apart from the constant environment in which it occurs the community is
Drepanocladus revolvens, cover of individual probably a climatic climax.
species is low. Aneura pinguis is frequent and
Bryum pseudotriquetrum, Blindia acuta, Campy- No sub-communities.
lium stellatum and Calliergon trifarium are occa-

33
M13 Schoenus nigricans – Juncus
frequently the constant Potentilla erecta and Erica
subnodulosus mire tetralix.
Bryophytes vary in cover and species but can
In this community Schoenus nigricans is typically be very extensive. The commonest throughout are
very frequent and consistently associated with Campylium stellatum and Calliergon cuspidatum.
other distinctive floristic features. It is generally Other frequent species include Drepanocladus
dominant (although it may be absent from frag- revolvens, Aneura pinguis, Cratoneuron commu-
mentary stands) giving a grey-green appearance to tatum and C. filicinum.
the vegetation. Commonly it is intermixed with This community is confined to peat or mineral
Juncus subnodulosus, and where this predomi- soils, in and around lowland mires irrigated by
nates the vegetation is olive-green in spring and base-rich, highly calcareous, and oligotrophic
reddish brown in winter. Molinia caerulea is also waters. It is often found below springs and seep-
constant. These species form a rough sward about age lines and on flushed margins of valley mires,
50 cm in height with smaller herbs growing in- but also extends into topogenous basins provided
between. Sedges are often important, particularly there is close contact with waters draining from
Carex panicea, C. lepidocarpa and C. flacca. lime-rich substrates. The flushing waters typically
Where the summer water table is close to the have pH between 6.5 and 8. It is widespread but
surface, species such as Equisetum palustre, local throughout lowland England and Wales, but
Pedicularis palustris, Mentha aquatica, Valeriana is restricted by natural scarcity of suitable habitat
dioica and Cardamine pratensis occur, sometimes and its extensive destruction.
with Parnassia palustris, Pinguicula vulgaris and The structure and floristics of this community
Eriophorum latifolium. A variety of orchids are are often influenced by grazing and some stands
found, particularly Epipactis palustris. Taller have been affected by mowing and burning.
herbs can be locally abundant with Succisa Shallow peat-digging has been locally important
pratensis being most common. Phragmites australis in providing a suitable habitat for the community
is also frequent, particularly in ungrazed stands. but more drastic treatment of mires, particularly
On drier areas and particularly tops of Schoenus draining and eutrophication, have reduced its
tussocks less calcicolous plants are found, most extent and eliminated it from some areas.

34
 M13

Juncus subnodulosus and Molinia General floristic and structural features well preserved, with at least
caerulea often very abundant with some of Anagallis tenella, Pedicularis palustris, Angelica sylvestris,
Schoenus nigricans markedly Cirsium palustre, Mentha aquatica, Equisetum palustre and Phragmites
reduced in vigour. Festuca rubra, australis frequent.
Holcus lanatus, Agrostis canina
and A. stolonifera frequent and tall
herbs, orchids and bryophytes Carex hostiana and C. pulicaris very Many smaller runnel herbs sporadic, but
patchy. frequent among an abundant and diverse Caltha palustris and Valeriana dioica
small herb flora in runnels with Briza become common and taller dicotyledons
media, Pinguicula vulgaris, Linum are often prominent with Filipendula
catharticum, and Juncus articulatus com- ulmaria, Eupatorium cannabinum and
mon. Lychnis flos-cuculi frequent.

M13a M13b M13c

Festuca rubra – Juncus acutiflorus Briza media – Pinguicula vulgaris sub- Caltha palustris – Galium uliginosum
sub-community community sub-community
This comprises the more impover- This kind of Schoenetum is strikingly Schoenus nigricans, Juncus subnodulo-
ished stands of M13. Apart from rich. Mixtures of Schoenus nigricans, sus and Molinia caerulea remain of
the reduction or even absence of Juncus subnodulosus and Molinia structural importance but are variously
Schoenus nigricans and presence caerulea usually share dominance but augmented by Carex rostrata, C. diandra,
of the species mentioned above, the small herbs of runnels are especially C. elata, Cladium mariscus and some-
Carex panicea, C. lepidocarpa, and distinctive. Apart from the species men- times Phragmites australis. Runnels are
C. flacca are also important in run- tioned above, Parnassia palustris is fre- well developed but smaller preferentials
nels. Anagallis tenella is totally quent often with mixtures of orchids of M13b are only occasional. The com-
absent and Pedicularis palustris, including Gymnadenia conopsea var. monest species are Carex panicea, C. lep-
Epipactis palustris and other densiflora, Dactylorhiza fuchsii, idocarpa, Mentha aquatica, Hydrocotyle
orchids are very scarce. The com- D. majalis ssp. purpurella and Epipactis vulgaris together with the preferentials
monest herbs are Succisa pratensis palustris. Along with Succisa pratensis Caltha palustris and Valeriana dioica. In
and Hydrocotyle vulgaris. and Serratula tinctoria, taller herbs are addition to Epipactis palustris there is
Bryophytes are generally sparse represented by frequent Angelica often Dactylorhiza incarnata, D. majalis
and low in number with Calliergon sylvestris, Cirsium palustre, Eupatorium ssp. praetermissa and sometimes
cuspidatum the commonest cannabinum and Oenanthe lachenalii. D. traunsteineri. Taller herbs are
species. Bryophytes are quite numerous and common, as listed above, with sprawling
sometimes of high cover. Galium uliginosum and less commonly
This sub-community occurs G. palustre. A pool element is sometimes
through the range of M13. This sub-community occurs in Anglesey present, with Carex rostrata and C. dian-
and East Anglia. dra together with Menyanthes trifoliata,
Equisetum fluviatile and Utricularia
species.
This sub-community is concentrated in
East Anglia.

35
M14 Schoenus nigricans – M15 Scirpus cespitosus –
Narthecium ossifragum mire Erica tetralix wet heath
This is a vegetation type with few constants and a
This mire type includes mildly calcicolous
wide variation in the pattern of dominance and
Schoenus vegetation of south-west England that is
in associated flora. Molinia caerulea, Scirpus
not readily integrated into Schoenus nigricans –
cespitosus, Erica tetralix and Calluna vulgaris are
Juncus subnodulosus mire (M13) and with a less
all of high frequency and it is mixtures of these
varied flora. Schoenus nigricans is usually domi-
species that give the vegetation its general
nant and Molinia caerulea is generally abundant.
character. However, sometimes one or two of them
A mixture of these two species usually cover the
may be missing and their relative proportions are
ground. Juncus subnodulosus is absent in contrast
very diverse. Molinia is the most consistent over-
to M13. Small calcicolous herbs are generally
all and often abundant; in other stands Scirpus is
absent. Narthecium ossifragum and Anagallis
very prominent and both may share dominance
tenella are constants while Drosera rotundifolia,
with Calluna. Molinia may also dominate with
growing on cushions of Sphagnum, is less com-
Scirpus or with Erica tetralix. The shrubby species
mon. Erica tetralix, or occasionally Calluna vul-
Erica cinerea, Vaccinium myrtillus and Myrica
garis, grows on Schoenus or Molinia tussocks.
gale are important in particular sub-communities.
Some stands have a local abundance of Myrica
Other common species are Potentilla erecta,
gale.
and in moister stands, Polygala serpyllifolia,
Bryophytes are variable and also less calcicolous
Narthecium ossifragum and Eriophorum angusti-
in character than in M13. Campylium stellatum and
folium. By contrast E. vaginatum is notably scarce.
Aneura pinguis are frequent and together with
There are few bryophytes common throughout.
Scorpidium scorpioides and, less commonly,
There are usually some Sphagnum spp. but they
Drepanocladus revolvens, can form extensive mats
do not form the luxuriant carpets of the
in runnels. Sphagnum spp. are a consistent feature,
Sphagnetalia mires (M17-M21). The most frequent
particularly on tussocks. Sphagnum subnitens is
species overall are Sphagnum capillifolium and
most common and S. auriculatum is frequent.
S. subnitens. Sphagnum palustre, S. recurvum
Hypnum jutlandicum is preferential and there may
and S. auriculatum can become common in wetter
be patches of hepatics including Kurzia pauciflora
stands. Lichens do not appear consistently but
and Calypogeia species.
Cladonia spp. can be locally prominent.
This community is characteristic of peats and
This wet heath community is characteristic of
mineral soils irrigated by moderately base-rich
moist and generally acid and oligotrophic peats
and calcareous ground waters of a pH range
and peaty mineral soils in the wetter western and
between 5 and 7. It characteristically occurs as iso-
northern parts of Britain. It is associated with thin-
lated flushes among wet heath and moorland veg-
ner or better drained areas of ombrogenous peat
etation, but it is also associated with soligenous
with a surface pH typically between 4 and 5. The
zones within valley mires. The community occurs
community is particularly well represented in the
very locally in Cornwall, east Devon, south-east
west and south-west of Scotland, through Wales
Dorset and the New Forest. It may also be found in
and less extensively in the Lake District, Dartmoor
Wales but it is replaced in comparable situations
and Exmoor.
on north-western blanket bogs by Schoenus-domi-
Grazing and burning have important effects on
nated stands of Scirpus cespitosus – Erica tetralix
the floristics and structure of this community, and
wet heath (M15).
draining and peat-cutting have extended its
The community only occurs very locally. This
coverage to formerly deeper and wetter peats.
is partly because of the natural scarcity of suitable
Without burning or grazing, less damaged stands
habitats, but also because of the reduction in its
may be able to revert to blanket mire. However,
extent by human activities such as drainage and
cessation of burning, especially on peat that is
agricultural improvement. Occasional burning
well aerated or where there has been drainage,
and light grazing are also of common occurrence
may precipitate a vigorous expansion of Molinia.
over the tracts of heath in which this kind of mire
Although progression to woodland is theoretically
usually occurs, although these activities are
possible over most, if not all, of its altitudinal
probably not very damaging. In the absence of
range, widespread deforestation has often re-
grazing or burning it is expected that some stands
moved potential seed-parents, and continued
of this community would progress towards
grazing by livestock and deer and sporadic burn-
wet woodland.
ing may be enough to set back succession contin-
No sub-communities. ually. However, extensive tracts of this kind of
vegetation have been replaced by coniferous forest
after the ground has been drained.

36
 M15

Narthecium ossifragum and Narthecium ossifragum fairly

Eriophorum angustifolium common, but other species
frequent with Sphagnum listed opposite all scarce.
palustre common. Myrica
gale often found.

Usually small stands, often Drosera rotundifolia and Erica cinerea frequent, Erica cinerea rare but
in soakways. Sphagnum small sedges at most occa- sometimes abundant, but Vaccinium myrtillus fre-
carpet extensive with fre- sional. Sphagnum carpet Vaccinium myrtillus rare. quent, commonly with
quent Sphagnum recurvum patchy, but S. papillosum Racomitrium lanuginosum Nardus stricta, Juncus
and S. subnitens. Drosera quite frequent. common and Cladonia spp. squarrosus and Deschampsia
rotundifolia common and often abundant. flexuosa. Racomitrium
scattered small sedges such lanuginosum and
as Carex echinata, C. panicea Cladonia spp. scarce.
and C. nigra.

M15a M15b M15c M15d

Carex panicea sub-community Typical sub-community Cladonia spp. sub-community Vaccinium myrtillus sub-community

This is the richest and most floristically The dominants here are very variable. All four possible dominants have high Mixtures of Molinia caerulea and
distinct sub-community. Molinia Scirpus cespitosus and Calluna vulgaris frequency but Calluna vulgaris usually Calluna vulgaris generally dominate
caerulea and Erica tetralix retain high may share dominance, or Calluna and predominates. Potentilla erecta remains with Scirpus cespitosus and Erica
frequency; Scirpus cespitosus and Molinia caerulea may predominate, constant, but Polygala serpyllifolia and tetralix both rather uneven, often
Calluna vulgaris are more sparse. Molinia and Erica tetralix or Molinia Narthecium ossifragum are less com- with some Vaccinium myrtillus.
Myrica gale sometimes has local alone may be dominant. Myrica gale mon and Eriophorum angustifolium and Commonly there are small tussocks
abundance, but E. cinerea and is quite common but not abundant. Myrica gale very scarce. Sphagnum of Nardus stricta, Juncus squarrosus,
Vaccinium myrtillus almost totally Narthecium ossifragum and Eriophorum spp. are only poorly represented and Deschampsia flexuosa and more
absent. Potentilla erecta and Polygala angustifolium are frequent as in M15a, Hypnum cupressiforme/jutlandicum occasionally some Anthoxanthum
serpyllifolia are very commonly found but small sedges are generally sparse, and Racomitrium lanuginosum become odoratum, Festuca ovina/vivipara,
with Narthecium ossifragum and Erica with only Carex panicea and C. echina- frequent. Cladonia spp. are abundant, F. rubra, Luzula multiflora and
angustifolium. Drosera rotundifolia is ta occasional and fen associates very particularly Cladonia impexa and Carex pilulifera. Sphagnum spp. are
preferential with a variety of species uncommon. Nardus stricta and Juncus C. uncialis together with C. arbuscula, infrequent, their place being taken by
such as Carex panicea, C. echinata, squarrosus may show local promi- C. pyxidata, C. coccifera and C. gracilis. Hypnum cupressiforme/jutlandicum,
C. nigra, C. pulicaris, C. demissa, nence. Eriophorum vaginatum is a Dicranum scoparium, Pleurozium
This sub-community is especially
C. dioica, Selaginella selaginoides, low-cover occasional. Sphagnum papil- schreberi, Plagiothecium undulatum,
common in the drier regions of the
Pinguicula vulgaris, Succisa pratensis, losum is frequent and locally abundant Polytrichum commune and
distribution of M15.
Viola palustris, Juncus bulbosus and Odontoschisma sphagni often Rhytidiadelphus loreus. Racomitrium
and Dactylorhiza maculata ssp. present. In some stands Sphagnum lanuginosum and Cladonia spp.
maculata. The Sphagnum carpet is spp. are sparse and mosses such as are scarce.
also distinctive, as indicated above, Racomitrium lanuginosum, Dicranum
This sub-community is especially
with Sphagnum palustre also often scoparium, Hypnum cupressiforme and
common in the drier regions of the
abundant and with S. capillifolium Campylopus paradoxus provide most
distribution of M15.
patchy. of the cover.
This and the Typical sub-community This and the Carex panicea sub-
are particularly common in the west community are particularly common
of Scotland. in the west of Scotland.

37
M16 Erica tetralix – Sphagnum Sphagnum spp. as well as a number of other
bryophytes, between the dominants. Lichens may
compactum wet heath also be present, especially larger Cladonia species
such as C. impexa and C. uncialis.
This community is characteristically dominated This wet heath community is found on acid
by mixtures of Erica tetralix, Calluna vulgaris and and oligotrophic mineral soils or shallow peats
Molinia caerulea, but their proportions are very that generally have a surface pH of between 3.5
variable, being influenced by differences in the and 4.5 and that are at least seasonally water-
water regime and trophic state of the soils, and logged. It is characteristic of the south of lowland
also by grazing and burning. Erica tetralix is often England, being particularly associated with the
vigorous, particularly on wetter soils, while surrounds of valley mires maintained by a locally
Calluna is often subordinate and weak (although it high water table. It is also found through Wales,
may be abundant in drier stands or where and in northern England and Scotland, where it
controlled burning is carried out). No other sub- extends on to thin ombrogenous peats at higher
shrubs attain a high frequency, although Erica altitudes.
cinerea and Ulex gallii may be abundant in Grazing and burning are important in main-
transitions to drier heaths in south-west England taining the vegetation, and burning is able to
and E. cinerea and U. minor can occur in similar transform the appearance of particular stands over
situations further east. In some situations Molinia short periods of time, producing considerable
may be dominant. structural diversity within a small area. Without
This community may have no other, or only any grazing or burning most stands would proba-
sporadic, vascular associates. The commonest vas- bly progress to woodland, and this has happened
cular associate, where present, is Scirpus cespito- to some stands lying within tracts of heath on
sus. Eriophorum angustifolium and Narthecium commons in south-east England where traditional
ossifragum are quite frequent, as is Drosera rotun- management has fallen into disuse. The combina-
difolia in wetter hollows. Myrica gale occurs occa- tion of frequent burning, draining, and damage
sionally, sometimes with local abundance. due to other operations such as military manoeu-
Most characteristic of the bryophyte layer in vres and mineral extraction, have led to an irre-
drier situations is Sphagnum compactum, a con- trievable loss of this community in many areas
stant and strong preferential for the community. In and its distribution has been considerably frag-
wetter places S. tenellum may be present. These mented with remaining stands closely hemmed in
species may occur as scattered cushions or form a by coniferous plantations or intensive agricultural
continuous carpet, sometimes with several other land.

38
 M16

Molinia caerulea usually Scirpus cespitosus and

dominant with Scirpus Narthecium ossifragum
cespitosus and Narthecium generally frequent but listed
ossifragum less common associates of M16b usually
than usual, scattered plants uncommon.
of Potentilla erecta and
Succisa pratensis frequent, Sub-shrub cover generally Sphagnum carpet often very
with occasional Polygala patchy, but Sphagnum patchy and sometimes absent
serpyllifolia, Carex panicea, carpet quite extensive with and M16c associates hardly
Salix repens, Cirsium frequent Kurzia pauciflora. ever found.
dissectum and Serratula Wetter hollows and runnels
tinctoria. Bryophytes have Drosera rotundifolia
Molinia caerulea reduced in Very variable mixtures
usually sparse. and often D. intermedia,
frequency and abundance of Calluna vulgaris, Erica
Rhynchospora alba and tetralix and Molinia caerulea
and Calluna vulgaris often
locally R. fusca as sub- generally dominate with
exceeding Erica tetralix.
community preferentials. occasional Scirpus cespitosus
Frequent tussocks of Scirpus
cespitosus and Juncus and Juncus squarrosus.
squarrosus. Hypnum Other associates listed
cupressiforme, Dicranum for M16d rare.
M16c scoparium, Racomitrium
M16b
lanuginosum and
M16a
Rhynchospora alba – Drosera Diplophyllum albicans
Succisa pratensis – Carex
intermedia sub-community all common. Cladonia
panicea sub-community Typical sub-community
species often present.
Molinia caerulea tends to Molinia caerulea and Erica tetralix All variations in the proportions of
predominate with Erica tetralix remain very frequent but Calluna vul- Molinia caerulea, Erica tetralix and
garis is less common and the cover of M16d
and Calluna vulgaris having Calluna vulgaris can be found in this
high frequencies. The vascular all three is reduced. In the intervening Juncus squarrosus – Dicranum sub-community so the appearance is
flora is richer than in M16a open areas is an extensive cover of scoparium sub-community very diverse. Where their cover is open,
with Potentilla erecta and Sphagnum compactum and S. tenellum Molinia caerulea rarely has high cover Sphagnum compactum and S. tenellum
Succisa pratensis being con- with leafy hepatics and locally promi- and may be absent and although can be very frequent and often abun-
stant and the species listed nent Cladonia species, particularly Erica tetralix retains constancy, dant. Among other bryophytes Hypnum
above preferential. Myrica C. impexa. There are scattered small Calluna vulgaris is usually dominant. jutlandicum and Kurzia pauciflora are
gale sometimes shows local tussocks of Scirpus cespitosus and Scirpus cespitosus is more frequent the commonest species, while
abundance. Bryophytes and often Narthecium ossifragum and and has higher cover than in other sub- Campylopus brevipilus is very
lichens tend to be less common Eriophorum angustifolium. More communities. Sphagnum compactum consistent in some areas. Lichens,
in this sub-community, distinctive are the preferentials is often abundant and has a high particularly Cladonia impexa, can
but unusually Sphagnum listed above. frequency together with S. tenellum also be frequent.
auriculatum can be frequent This sub-community is concentrated in while S. capillifolium and S. subnitens
This sub-community is found
and locally abundant. the New Forest and Poole Harbour. are relatively scarce. Together with
throughout the south of Britain and as
This sub-community occurs the bryophytes listed above lichens are
more impoverished stands further north.
throughout south-west England often prominent, with Cladonia impexa
and perhaps elsewhere. and C. uncialis especially common.
This is the usual sub-community in the
north and east of Britain.

39
M17 Scirpus cespitosus –
nied by S. tenellum, S. subnitens and other
Eriophorum vaginatum blanket mire species, forming luxuriant carpets. Such carpets
typically have a variety of leafy hepatics including
This community is dominated by mixtures of Odontoschisma sphagni, Mylia anomala, M. tay-
monocotyledons, ericoid sub-shrubs and Sphagnum lori and Pleurozia purpurea. Racomitrium lanugi-
spp. It can occur as extensive, relatively uniform nosum is a common moss throughout, but becomes
tracts, or as hummock and hollow complexes, abundant on hummock tops and in degraded
with this community giving way to bog pool vege- mires. Lichens, particularly larger Cladonia
tation in the hollows. Among the bulkier vascular species, can be prominent and tend to be associat-
species, the most common are Scirpus cespitosus, ed with R. lanuginosum.
Eriophorum vaginatum, Molinia caerulea, Calluna This community is the characteristic blanket
vulgaris and Erica tetralix; mixtures of which form bog vegetation of the more oceanic parts of Britain,
a rather open uneven-topped tier which is 20- occurring extensively on waterlogged ombroge-
30 cm tall. Myrica gale also has occasional local nous peat. The peats show varying humification
abundance in this stratum. but are typically highly acidic, with a surface pH
Eriophorum angustifolium and Narthecium usually not above 4 and often less. It is a commu-
ossifragum are both very frequent and Drosera nity of lower altitudes where extreme humidity is
rotundifolia is very common in wetter areas. combined with a relatively mild winter climate. It
Potentilla erecta is a constant which helps to dis- is largely confined to western Britain from the
tinguish this community from other Sphagnetalia western Highlands of Scotland and the Western
mires (M18-M21). Other species found at low fre- Isles, to south-west Scotland, the Lake District,
quencies throughout are Pedicularis sylvatica, Wales and south-west England.
Huperzia selago, Juncus acutiflorus, Festuca ovina Burning, marginal peat-cutting, and drainage
and Carex echinata. Vaccinium myrtillus, Empetrum have often resulted in surface drying of the peat
nigrum ssp. nigrum and Rubus chamaemorus and hence a modification of the vegetation. It is
are all scarce, in contrast to Calluna vulgaris- also possible that natural climatic change too has
Eriophorum vaginatum blanket mire (M19). played a part in the degeneration of the blanket
Sphagnum spp. are an important component of bogs occupied by this type of community.
the ground layer. Sphagnum capillifolium and However, this community still remains as climax
S. papillosum are constants and may be accompa- vegetation in the more oceanic parts of Britain.

40
 M17

Erica tetralix and Molinia The usual vascular

caerulea usually well dominants are Calluna
represented with Drosera vulgaris and Scirpus
rotundifolia very frequent cespitosus, with Erica
and Myrica gale locally tetralix and Molinia caerulea
plentiful. Extensive and of reduced importance and
varied Sphagnum carpet Myrica gale rare. Drosera
with leafy hepatics often rotundifolia at most
prominent. occasional. Sphagnum
cover rather impoverished
and leafy hepatics infrequent.

M17a Racomitrium lanuginosum very common with several Juncus squarrosus, Nardus stricta and Deschampsia flexu-
Cladonia species. Erica cinerea occasional and locally osa frequent with small amounts of Vaccinium myrtillus
Drosera rotundifolia – Sphagnum abundant. Vaccinium myrtillus and Empetrum nigrum and occasional Empetrum nigrum, but Erica cinerea rare.
spp. sub-community very scarce. Nardus stricta and Juncus squarrosus Racomitrium lanuginosum occasional and Cladonia
occasional. species uncommon.
This has a consistent representation
of all the community constants.
Among vascular dominants mixtures of
Calluna vulgaris and Scirpus cespitosus M17b M17c
or Calluna vulgaris and Molinia
caerulea usually make up the bulk of Juncus squarrosus – Rhytidiadelphus loreus
Cladonia spp. sub-community sub-community
the cover, with Eriophorum vaginatum
sometimes showing local abundance Calluna vulgaris and Scirpus cespitosus are fairly Calluna vulgaris and Scirpus cespitosus are the main
in higher areas and Erica tetralix consistent co-dominants with Molinia caerulea and vascular dominants with Erica tetralix and especially
in wetter parts. Drosera rotundifolia Erica tetralix playing a subordinate role and Eriophorum Molinia caerulea being reduced. Myrica gale is absent and
is strongly preferential. Sphagnum vaginatum distinctly patchy. Myrica gale is scarce and Erica cinerea very scarce. There is a marked increase in
spp. are extensive; Sphagnum Erica cinerea quite frequent and locally prominent. The Juncus squarrosus, Nardus stricta, Deschampsia flexuosa
capillifolium and S. papillosum are Sphagnum carpet is much impoverished with Sphagnum and Carex nigra, and with them Agrostis canina ssp. cani-
most abundant with S. compactum capillifolium as the main species, often rather patchy, na, Anthoxanthum odoratum and Luzula multiflora can be
occasional and S. tenellum and and all other species reduced in frequency. The leafy found. Of the Sphagnum spp., Sphagnum papillosum is
S. subnitens frequent. Common leafy hepatics of M17a are uncommon, with Mylia taylori and usually the most abundant. There is a distinctive contin-
hepatics are Pleurozia purpurea and Diplophyllum albicans preferential at low frequencies. gent of mosses: Hypnum cupressiforme/jutlandicum,
Odontoschisma sphagni. Racomitrium Racomitrium lanuginosum and Cladonia spp. are Rhytidiadelphus loreus and Dicranum scoparium are all
lanuginosum is also frequent but only increased in frequency and abundance, particularly very frequent, while Polytrichum commune, P. alpestre,
as scattered shoots. Cladonia impexa, C. uncialis and C. arbuscula. Plagiothecium undulatum, Aulacomnium palustre,
Ptilidium ciliare, Pohlia nutans and Campylopus
This sub-community occurs throughout
This sub-community along with the Juncus – paradoxus are more occasional but preferential.
the range of M17, but it is particularly
extensive in north-west Scotland. Rhytidiadelphus sub-community occurs in the west but This sub-community along with the Cladonia sub-
they extend the range of the community on to drier peats, community occurs in the west but they extend the range
most notably in south-west and eastern Scotland. of the community on to drier peats, most notably in
south-west and eastern Scotland.

41
M18 Erica tetralix – Sphagnum
S. magellanicum, and a little S. tenellum predom-
papillosum raised and blanket mire inate on the surrounds to wetter depressions.
Other bryophytes are generally subordinate but
This community is generally dominated by can be frequent and locally abundant. The leafy
Sphagnum spp. Ericoid sub-shrubs and mono- hepatics Odontoschisma sphagni and Mylia
cotyledons are often subordinate. It can be found anomala are both common, but Pleurozia pur-
as undulating carpets or can comprise lawn and purea is generally absent. Aulacomnium palustre
hummock components. The bulkier vascular plants and Hypnum cupressiforme/jutlandicum are fre-
typically form a low, patchy canopy with Calluna quent mosses.
vulgaris, Erica tetralix and Eriophorum vaginatum This vegetation is characteristic of waterlogged
being the commonest species and Scirpus cespito- ombrogenous peats, typically at low altitudes in
sus slightly less frequent. Erica tetralix tends to moderately oceanic parts of Britain. It is charac-
predominate on wetter ground where shoots of teristic of raised bogs where it is the main com-
Eriophorum angustifolium can also be abundant. munity of the active plane, but is also found with-
Calluna, Scirpus and E. vaginatum are found more in blanket mires and in some basin mires on acid
typically on the drier areas. peat. The peats it covers are usually deep with a
Sphagnum spp. make up the most important uniformly acid surface with a pH of about 4, and
component of the vegetation. Both Sphagnum oligotrophic. It is widespread but local through
papillosum and S. capillifolium are very common the lowlands of Wales, up to the Scottish Borders
and S. tenellum is also a constant but less abun- and in south-west Scotland. There are also locali-
dant. Sphagnum magellanicum is a preferential ties in southern England and east Scotland.
species and a major peat-builder. S. imbricatum is Erica tetralix – Sphagnum papillosum raised
a distinctive species where present. Over gently- and blanket mire vegetation is a climax of a
undulating surfaces the Sphagnum spp. form an hydroseral succession. However, the typical habi-
irregular patchwork, but with increasing differen- tat of this community has been widely affected by
tiation of hummocks and hollows they show a ver- various treatments, notably peat-cutting, burning
tical stratification. On hummock tops and sides and grazing, and these have often modified the
S. capillifolium is abundant, and S. papillosum, vegetation or reduced it to fragmentary stands.

42
 M18

Sphagnum spp. luxuriant, Sphagnum spp. abundant,

with Sphagnum magellan- with Sphagnum capillifolium
icum frequent and abundant, usually dominant, S. papillo-
along with S. papillosum in sum frequent but usually
wetter lawns. Narthecium subordinate and S. magellan-
ossifragum and Drosera icum only occasional.
rotundifolia common and Narthecium ossifragum and
Vaccinium oxycoccos and Vaccinium oxycoccos occa-
Andromeda polifolia sional and Drosera rotundifo-
especially distinctive. lia and Andromeda polifolia
very scarce. Sub-shrubs often
quite vigorous. Cladonia
species frequent and locally
abundant.

M18a M18b

Sphagnum magellanicum – Empetrum nigrum ssp. nigrum –

Andromeda polifolia sub-community Cladonia spp. sub-community

All the community vascular constants Calluna vulgaris, Scirpus cespitosus

are of high frequency but very often and Eriophorum vaginatum tend to
none is dominant. Sphagnum spp. have higher covers here, Calluna
form an obvious and extensive carpet vulgaris in particular becoming more
in which Sphagnum papillosum, often vigorous and abundant. Empetrum
with abundant S. magellanicum, pre- nigrum is also frequent among
dominates along with the constants them. Among the Sphagnum spp.,
S. tenellum and S. capillifolium. S. capillifolium is strongly dominant,
Scattered through are frequent although S. papillosum is still
individuals of Drosera rotundifolia frequent. S. tenellum is patchy and
and Narthecium ossifragum with the S. magellanicum only occasional.
preferentials Vaccinium oxycoccos and Other mosses become frequent,
Andromeda polifolia. Cladonia spp. with Pleurozium schreberi and
and Pleurozium schreberi are typically Rhytidiadelphus loreus being good
of low frequency. preferentials; a range of hepatics is also
common. There is a marked increase
This sub-community occurs throughout
in Cladonia spp., notably Cladonia
the range of M18.
impexa, C. uncialis and C. arbuscula,
each of which can be locally abundant.
This sub-community occurs throughout
the range of M18.

43
M19 Calluna vulgaris – Eriophorum
comium splendens is common at higher altitudes.
vaginatum blanket mire A variety of leafy hepatics occur in this
community and lichens are frequent. Larger
This vegetation is generally dominated by Cladonia spp. can be abundant on old Eriophorum
mixtures of Eriophorum vaginatum and ericoid hummocks.
sub-shrubs. Sphagnum spp. can be prominent This mire is the typical blanket bog vegetation
over wetter ground but are not as luxuriant or rich of high-altitude ombrogenous peats present in
as in Scirpus cespitosus – Eriophorum vaginatum the wet and cold climate of the uplands of north-
blanket mire (M17) or Erica tetralix – Sphagnum ern Britain. In particular, it occurs on high-level
papillosum raised and blanket mire (M18). The plateaux and broad watersheds, usually above
ground surface is often uneven, but does not show 300 m, and is confined to deeper peats, usually
true hummock and hollow relief. Eriophorum more than 2 m thick, on flat or gently-sloping
vaginatum is abundant and at least co-dominant. ground. The peats are usually well-humified,
Normally this community has very frequent highly acidic with a surface pH often less than 4.
occurrences of Calluna vulgaris, Vaccinium They are not consistently waterlogged and may
myrtillus and Empetrum nigrum ssp. nigrum and, become surface oxidised in summer. Erosion of
at higher altitudes, V. vitis-idaea, V. uliginosum the peat is common. This community is found on
and E. nigrum ssp. hermaphroditum. Overall the higher ground in the Pennines, the central
Calluna is the most common co-dominant along Highlands of Scotland, and Wales.
with Eriophorum vaginatum, but diverse mixtures Treatments such as burning and grazing are
of these sub-shrubs are very frequent. Vascular important in influencing the composition and
associates are few, the commonest being structure of the vegetation throughout the range of
Eriophorum angustifolium, and Rubus chamae- this community, in particular where stands form
morus, a species which is distinctive for this com- part of unenclosed hill grazing or grouse moors.
munity. Deschampsia flexuosa and Juncus squar- A stable diversity of bog vegetation can be
rosus occur occasionally throughout, and at high- maintained by careful burning on a rotation of
er altitudes Carex bigelowii becomes frequent. around 10 years, or by moderate levels of grazing.
In contrast to the vascular plants, the bryophyte However, frequent burning or heavy grazing
flora is rich, often with a cover exceeding 50%. contribute to the conversion of the Calluna vul-
The most frequent Sphagnum species is S. capilli- garis – Eriophorum vaginatum blanket mire to
folium, which forms patches rather than carpets. Eriophorum vaginatum blanket mire (M20). In
Hypnaceous mosses are consistently present and other cases drainage can convert this community
Pleurozium schreberi, Rhytidiadelphus loreus, into heathland or grassland, and this type of blan-
Hypnum cupressiforme/jutlandicum and Plagio- ket mire has been reclaimed for agriculture or
thecium undulatum are all very frequent. Hylo- forestry in many areas.

44
 M19

Eriophorum vaginatum less Eriophorum vaginatum

abundant, Erica tetralix usually abundant and Rubus
more common and Scirpus chamaemorus frequent.
cespitosus frequent. Rubus Scirpus cespitosus rare and
chamaemorus rare. Sphagnum Sphagnum cover patchy.
spp. cover quite extensive.

Empetrum nigrum ssp. Empetrum ssp. hermaphroditum and

hermaphroditum occasional Vaccinium vitis-idaea constant and
at most and often with ssp. often co-dominant with Calluna
nigrum as a replacement. vulgaris, V. myrtillus and Eriophorum
Vaccinium vitis-idaea also vaginatum. Hylocomium splendens,
occasional and V. uliginosum, Racomitrium lanuginosum and
Juncus squarrosus and Carex Polytrichum alpestre common in
bigelowii rare. Bryophytes in the ground layer.
M19c rare.

M19a M19b M19c

Erica tetralix sub-community Empetrum nigrum ssp. nigrum Vaccinium vitis-idaea – Hylocomium splendens
sub-community sub-community
Eriophorum vaginatum can be co-
dominant with the sub-shrubs though This sub-community preserves all This embraces all the high-montane blanket mire.
generally less abundant than in other the features of M19 although it is very It preserves all the general floristic features but is
sub-communities. Calluna vulgaris variable in all its structural elements. distinctive in the frequent and consistent presence
is often the predominant sub-shrub Eriophorum vaginatum is usually of Vaccinium vitis-idaea and Empetrum nigrum
with some Empetrum nigrum ssp. abundant, with sub-shrubs playing a ssp. hermaphroditum and the more restricted pres-
nigrum. Erica tetralix is preferential consistent role and Calluna vulgaris ence of V. uliginosum. These are mixed with other
and sometimes with high cover. being the leading species, though sub-shrubs and usually with abundant Calluna
Scirpus cespitosus is frequent and Vaccinium myrtillus and/or Empetrum vulgaris. Eriophorum angustifolium is less com-
locally abundant. The infrequency of nigrum ssp. nigrum, particularly mon, but Rubus chamaemorus is very frequent.
Rubus chamaemorus is distinctive. the latter, can increase greatly after Juncus squarrosus occurs quite often, as do Carex
Sphagnum spp. tend to be consistently burning. Erica tetralix is very scarce bigelowii and Scirpus cespitosus. Sphagnum spp.
abundant with S. capillifolium being and Scirpus cespitosus and Molinia can be quite prominent, with a range of other
commonly accompanied by caerulea have a low frequency. Rubus bryophytes including the hepatic Ptilidium ciliare.
S. papillosum. chamaemorus however increases great- The lichen flora is usually well developed.
ly. The Sphagnum flora is impover-
This is the usual form found at lower This sub-community extends to the altitudinal
ished and on drier ground hypnaceous
altitudes that have a rather more limit of this kind of blanket bog in central
mosses are particularly abundant.
oceanic climate, such as in Wales Scotland.
and south-west Scotland. This forms the richer type of ‘Pennine
Three variants.
blanket bog’ extending northwards
through Cheviot and the Borders into
eastern Scotland.

45
M20 Eriophorum vaginatum Bulkier species like Cladonia arbuscula,
C. uncialis and C. impexa can sometimes be found
blanket and raised mire or sometimes there is just a patchy cover of peat
encrusters.
Eriophorum vaginatum mire comprises species- This community is characteristic of ombroge-
poor ombrogenous bog vegetation dominated by nous peats on bogs where certain treatments have
E. vaginatum, the tussocks of which form an open greatly affected the vegetation; grazing and burn-
or closed canopy 10-30 cm high. The only other ing have been of greatest significance, but draining
constant plant is E. angustifolium, which is usually and aerial pollution have also played a part. It is
found as scattered shoots. Ericoid sub-shrubs are commonest on blanket mires, where these factors
patchy; Calluna vulgaris, Empetrum nigrum ssp. have contributed both to floristic impoverishment
nigrum and Vaccinium myrtillus can each be and to gross erosion of the peats, but is also found
found quite frequently and the last two may be locally on run-down raised bogs. The Eriophorum
locally abundant. Alternatively these species may mire is present mainly between 500 m and 700 m
be reduced to sparse shoots. Deschampsia flexu- where the climate is cold and wet. The peats are
osa is fairly common while Festuca ovina, Juncus generally dry, often showing surface oxidation and
squarrosus, Scirpus cespitosus and Carex bigel- with a pH frequently as low as 3. This community
owii are all infrequent. can be found locally through northern Britain, and
Bryophytes are sparse and patchy. Sphagnum is especially extensive in the southern Pennines.
spp. are scarce with Sphagnum capillifolium and This community has been seen to revert to the veg-
S. papillosum the most usual species. Hypnaceous etation characteristic of the richer blanket bog
mosses are poorly represented; the only moss of community Calluna vulgaris – Eriophorum vagi-
any frequency being Campylopus paradoxus natum mire (M19) within 25 years of enclosure
which can be accompanied by Dicranum scopari- and freedom from burning, but in many instances,
um. There may be occasional shoots of Ortho- particularly if intensive grazing or frequent burn-
dontium lineare, Pohlia nutans and Drepa- ing have been accompanied by drainage, the
nocladus fluitans. A variety of leafy hepatics may degeneration of the vegetation is perhaps
be present. Lichens are typically few in number. irreversible.

M20

Calluna vulgaris, Vaccinium myr- Sub-shrubs at most occasional and

tillus and Empetrum ssp. nigrum sparse among abundant Eriophorum
occur frequently. Campylopus para- vaginatum with an impoverished
doxus and Dicranum scoparium bryophyte and lichen flora.
also frequent with occasional
patches of peat-encrusting lichens.

M20b M20a

Calluna vulgaris – Cladonia spp. Species-poor sub-community

sub-community
Eriophorum vaginatum forms the
Eriophorum vaginatum not com- main vascular bulk, with only scat-
pletely dominant though, together tered E. angustifolium, occasional
with E. angustifolium, it is the most sub-shrubs and sometimes a little
frequent species. Sub-shrubs scat- Deschampsia flexuosa. Ground pre-
tered, sometimes showing local dominantly bare with scattered
dominance. Bryophytes frequent tufts of Campylopus paradoxus,
but in small amounts. Larger Orthodontium lineare and
Cladonia spp. among shrub stools Calypogeia spp. Lichens
and old Eriophorum tussocks. typically very sparse.

46
M21 Narthecium ossifragum – Narthecium ossifragum are constant and often
abundant. Molinia caerulea is typical, but is often
Sphagnum papillosum valley mire weak and not tussock-forming, and Rhynchospora
alba is characteristic of one sub-community, being
This mire vegetation is dominated by carpets of most prominent around hollows and pools. The
Sphagnum spp. with scattered herbs and sub- sub-shrubs Erica tetralix and Calluna vulgaris
shrubs forming extensive lawns or the drier parts are very frequent, forming an open canopy.
of hummock and hollow complexes. The domi- More restricted but conspicuous when present is
nant Sphagnum is usually S. papillosum. Sphag- Myrica gale. Other plants are generally present as
num auriculatum or S. recurvum (and occasional- scattered individuals, Drosera rotundifolia being
ly both) are frequently encountered and less often the most frequent.
S. cuspidatum. Locally, S. magellanicum and S. This is a community of permanently water-
pulchrum may be present. Sphagnum compactum logged, acid and oligotrophic peats, especially
is almost always absent in contrast with Erica characteristic of valley mires maintained by a
tetralix – Sphagnum compactum wet heath (M16). locally high water table. The peat on which this
Only a few other mosses occur and they are gener- community is found is usually not very deep (20-
ally of low cover, but leafy hepatics are common. 150 cm) with a surface pH mostly in the range of
Odontoschisma sphagni and Kurzia pauciflora 3.5-4.5, and a water table at or very close to the
are most common, but Cladopodiella fluitans, surface. It is found locally in the southern low-
Cephalozia macrostachya, C. connivens, C. bicus- lands of Britain.
pidata and Calypogeia fissa have also been record- Neither burning nor grazing are important in
ed. Lichens are typically sparse, but hummock maintaining this community, as the wetness of the
tops can be a habitat for Cladonia impexa, vegetation gives its some protection from these
C. arbuscula and C. uncialis. treatments, but both can cause damage if com-
The vascular plant species present provide the bined with drainage. With continued autogenic
major distinction between this community and accumulation of peat, it would probably progress
other types of Erico – Sphagnion bogs (M17-M20). naturally to some kind of woodland but this
Eriophorum vaginatum and Scirpus cespitosus process is likely to be very slow in the absence of
are rare, but Eriophorum angustifolium and drainage.

M21

Rhynchospora alba very frequent Rhynchospora alba and Myrica gale

and Myrica gale occasional, rather infrequent, but Vaccinium
sometimes abundant in the oxycoccos patchily present and
Sphagnum carpet, which is Potentilla erecta occasional in a
usually dominated by mixtures of carpet usually dominated by
S. papillosum and S. auriculatum, Sphagnum recurvum with some
with S. recurvum scarce. S. papillosum but little
S. auriculatum.

M21a M21b

Rynchospora alba – Sphagnum Vaccinium oxycoccos – Sphagnum

auriculatum sub-community recurvum sub-community

In this, the most frequently Sphagnum papillosum is often

described M21 community, the abundant but somewhat patchy
Sphagnum carpet is generally and S. recurvum frequently has
dominated by mixtures of an equal cover. S. auriculatum
S. papillosum and S. auriculatum. is much reduced in frequency.
Hepatics are varied and often Odontoschisma sphagni occurs
abundant. All vascular constants sometimes but hepatics are greatly
have a high frequency and in addi- reduced. Among vascular plants
tion Rhynchospora alba is very Rhynchospora alba is scarce and
frequent. Myrica gale frequently Vaccinium oxycoccos reduced in
shows local abundance. frequency.
This sub-community is best This sub-community extends into
represented in central southern the north and west of England and
England. Wales.

47
M22 Juncus subnodulosus – Mentha aquatica, Caltha palustris, Equisetum
palustre, Carex panicea, Valeriana dioica, Hyper-
Cirsium palustre fen-meadow icum tetrapterum, Hydrocotyle vulgaris, Lotus
uliginosus, Lychnis flos-cuculi, Cardamine praten-
This community shows considerable variation in sis, Ranunculus acris, Potentilla erecta, Cerastium
its floristic composition which often reflects a fontanum, and Rumex acetosa, with several
unique and complex history of mowing and scrambling plants including Galium uliginosum,
grazing management. The most prominent struc- G. palustre, Vicia cracca and Lathyrus pratensis.
tural element typically consists of rushes and Bryophytes play a reduced role with only
sedges of moderate stature, appearing as a rank Calliergon cuspidatum and Brachythecium rutab-
sward if it is not grazed. Juncus subnodulosus is ulum being common throughout.
the most important of the bulkier species and the This community brings together secondary
most usual dominant. Juncus inflexus and J. artic- herbaceous vegetation developed over a variety of
ulatus are the commonest accompanying rushes. moist, base-rich and moderately mesotrophic
Of the sedges, the most striking are Carex acuti- peats in southern lowland Britain. It can be found
formis and C. disticha, either of which can be fre- either in, or around, well-developed springs,
quent or occasionally dominant. Much more occa- flushes and mires, or marking out more ill-defined
sionally C. elata or C. paniculata can occur. In areas of influence of surface or ground waters.
summer this rush and sedge layer can be This community marks out soils which are kept
overtopped by flowering dicotyledons. The most moist for most of the year and have a moderate to
frequent of these throughout the community are high base-status, and usually a pH range of 6.5-7.5.
Cirsium palustre, Filipendula ulmaria, Angelica The community has a wide distribution through
sylvestris, Succisa pratensis, Eupatorium canna- the southern British lowlands with particular con-
binum and Scrophularia aquatica. More locally centrations of stands in East Anglia, north
Lythrum salicaria, Lysimachia vulgaris, Valeriana Buckinghamshire and Anglesey.
officinalis, Thalictrum flavum and Symphytum This community is always dependent on vari-
officinale can be found, and, in Broadland, the ous kinds of treatment for its maintenance, as it is
nationally rare Peucedanum palustre. However, derived from other wetland vegetation types by
grazing may keep these species severely in check. mowing or grazing, or both. The reduction in graz-
Among smaller species, grasses are important, ing results in the expansion of bulky dominants
and species found include Holcus lanatus, and ranker grasses and the overwhelming of the
Festuca rubra and, less commonly, Arrhena- smaller herbs. Its overall distribution and the
therum elatius, together with Poa trivialis, extent of the stands has become much less than
Agrostis stolonifera, Anthoxanthum odoratum formerly because of intensive land improvement
and Briza media. Smaller herbs are those charac- and abandonment of traditional agricultural
teristic of moist grasslands, the commonest being practices.

48
 M22

Phragmites australis, Lythrum salicaria Species listed opposite at most

and Hydrocotyle vulgaris frequent, with occasional and hardly ever abundant.
Carex acutiformis or C. elata common,
sometimes abundant.
Juncus subnodulosus usually dominant Juncus subnodulosus often accompanied by
in rather rank and impoverished J. inflexus and J. articulatus; also Carex dis-
vegetation with bulkier grasses ticha and Deschampsia cespitosa patchily
Carex elata constant and sometimes Juncus subnodulosus usually dominant abundant. Molinia caerulea and abundant. Molinia caerulea, Briza media and
abundant among Juncus subnodulosus, with little or no Carex elata, but with dicotyledons listed opposite Anthoxanthum odoratum common at low
with frequent Potentilla palustris, frequent and sometimes abundant infrequent at most. cover with frequent records for Cardamine
Epilobium palustre, Equisetum C. acutiformis. Galium palustre pratensis, Cerastium fontanum, Trifolium
fluviatile and Dactylorhiza incarnata. common but other associates listed repens, T. pratense, Rumex acetosa, Plantago
opposite rare. lanceolata and Epilobium parviflorum.

M22c M22d M22a M22b

Carex elata sub-community Iris pseudacorus sub-community Typical sub-community Briza media – Trifolium spp. sub-community

Carex elata is uncommon in fen- Juncus subnodulosus is dominant, but In this, the most common kind of In comparison to M22a, in this sub-community
meadow vegetation but here it occurs as in M22c, Phragmites australis and fen-meadow, there are no preferential the rush and sedge tier has a lower cover and
with some abundance and occasionally Hydrocotyle vulgaris are often present floristic features and the general density and there is a richer associated flora.
is co-dominant with Juncus subnodulo- and Menyanthes trifoliata, Potentilla impression is of rank structure and Taller dicotyledons such as Cirsium
sus. Phragmites australis is often palustris and Equisetum fluviatile impoverishment. Juncus subnodulosus palustre, Filipendula ulmaria, Angelica
present and there is usually some occasional. Grasses and smaller herbs is frequent and the commonest sylvestris, Eupatorium cannabinum and
Hydrocotyle vulgaris. The plants listed characteristic of M22b are sparse. dominant. The commonest associates Succisa pratensis show an increased
above are preferential. Some stands Carex elata is occasional and tussocks are bulkier grasses such as Holcus frequency and often occur with Centaurea
have Pedicularis palustris, Menyanthes of C. paniculata are sometimes lanatus, Festuca rubra, sometimes nigra and Rumex acetosa. The lower sward is
trifoliata, Ranunculus flammula, prominent. Tall dicotyledons are Molinia caerulea, mat-formers such as particularly rich. There is a range of grasses,
R. lingua and Berula erecta. Grasses prominent in summer; Cirsium Agrostis stolonifera and Poa trivialis, often with Carex panicea and occasionally
and smaller herbs of M22b are palustre, Filipendula ulmaria, Angelica and taller dicotyledons such as Cirsium C. nigra and C. flacca. There is a variety of
particularly sparse. sylvestris and Succisa pratensis are palustre, Filipendula ulmaria, Angelica other herbs; most frequent are Lotus
all common and Iris pseudacorus, sylvestris, Succisa pratensis and uliginosus, Lychnis flos-cuculi, Caltha
This and the Iris sub-community are
Ranunculus flammula, Valeriana Eupatorium cannabinum. There are palustris, Ranunculus acris, Valeriana dioica,
especially well-represented on
officinalis, Lysimachia vulgaris also sprawling plants and a few shorter Potentilla erecta, P. anserina and Hypericum
topogenous mires in East Anglia.
and Thalictrum flavum are all frequent. species such as Mentha aquatica and tetrapterum which are characteristic of M22 as
Equisetum palustre. Within this general a whole. Strongly preferential are the species
This and the Carex sub-community are
framework there is considerable local listed above together with Prunella vulgaris,
especially well-represented on
variation. Ranunculus repens, Triglochin palustre,
topogenous mires in East Anglia.
Rhinanthus minor and Dactylorhiza fuchsii.
This sub-community is the most
Phragmites australis is generally scarce.
frequent type of spring-fed stands.
This sub-community is most frequently
developed around grazed spring-fens and wet
field hollows and occurs through the range
of M22.

49
M23 Juncus effusus/acutiflorus – Hydrocotyle vulgaris, Viola palustris, and Stellaria
alsine.
Galium palustre rush-pasture Bryophytes are variable in their cover. Where
the vegetation is open they may be abundant.
This vegetation is ill-defined and characterised by Calliergon cuspidatum is the most frequent, often
the abundance of either Juncus effusus or J. acuti- occurring with Brachythecium rutabulum and
florus, sometimes both, in a ground of mesophytic Rhytidiadelphus squarrosus.
herbs common in moist agricultural grassland. This rush-pasture occurs over a variety of
The rushes often have a high cover but they may moist, moderately acid to neutral, peaty and min-
also be more sparse. Juncus effusus is more abun- eral soils in the cool and rainy lowlands of west-
dant in the east, while J. acutiflorus has a distinct- ern Britain. It is a community of gently-sloping
ly western distribution. Diversity in dominants is ground around the margins of soligenous flushes,
not great but the associates are quite diverse, mak- as a zone around topogenous mires and wet
ing the bounds of this vegetation type hard to fix. heaths, and especially widespread in ill-drained,
Sometimes J. articulatus may be locally frequent comparatively unimproved or reverted pasture. It
and abundant. Holcus lanatus is the most frequent can be found on a variety of moderately acid to
grass, but Agrostis canina ssp. canina, A. stolon- neutral soils that are kept moist to wet for most of
ifera, Anthoxanthum odoratum and Poa trivialis the year with a pH in the range of 4-6. It is found
are also common in drier stands. Festuca rubra at the opposite climatic and edaphic extreme to
and Agrostis capillaris may also become frequent. the Juncus subnodulosus – Cirsium palustre fen
Molinia caerulea is increasingly common and meadow (M22) with a distinctly oceanic distribu-
abundant towards the west and there may be some tion. The community is wide-spread through the
sedges in the sward. There is a variety of common west of Britain from Devon and Cornwall to Skye
herbs. Among the taller species Cirsium palustre and Caithness.
is the commonest, Rumex acetosa, Angelica This community is maintained mainly by graz-
sylvestris and Epilobium palustre are frequent, ing and more occasionally mowing which pre-
and two sprawling species Galium palustre and vents the succession of the community to wood-
Lotus uliginosus are constant. Frequent smaller land. Draining and other kinds of soil improve-
species are Mentha aquatica, Ranunculus flam- ments such as fertilising and reseeding have
mula, R. repens, R. acris, Cardamine pratensis, reduced its former extent.

50
 M23

Juncus effusus very common, but Juncus effusus constant and usually
exceeded by J. acutiflorus, with dominant, with J. acutiflorus
Molinia caerulea and Holcus scarce. Holcus lanatus common,
lanatus frequent and sometimes but Molinia caerulea and
abundant. Filipendula ulmaria dicotyledons listed opposite
occasional, also some of all scarce.
Ranunculus acris, Potentilla erecta,
Achillea ptarmica and Equisetum
palustre and locally prominent
tall-fen herbs such as Lythrum
salicaria and Iris pseudacorus.

M23a M23b

Juncus acutiflorus sub-community Juncus effusus sub-community

Juncus acutiflorus and J. effusus This sub-community is less

are both constant, J. articulatus well-defined and is essentially a
is locally prominent and J. con- transition between M23a and the
glomeratus is occasional. The com- Holcus lanatus – Juncus effusus
monest grasses are Holcus lanatus rush-pasture (MG10). Other rushes
and, preferential here, Molinia comprise Juncus articulatus and
caerulea. Community herbs such as J. conglomeratus, but J. inflexus,
Cirsium palustre, Rumex acetosa common in MG10, is absent.
and Angelica sylvestris remain fre- Molinia caerulea is infrequent,
quent with Galium palustre and but grasses are important in the
Lotus uliginosus. Filipendula sward. Where the community is
ulmaria is more common than in surrounded by improved pasture,
M23b and there may be an abun- as is common, Cynosurus cristatus
dance of taller herbs. In the lower and Lolium perenne can be present
tier of vegetation, Mentha aquatica, at low cover. Good distinguishing
Cardamine pratensis, Ranunculus features are the high frequencies of
flammula and R. repens are Galium palustre, Cirsium palustre,
frequent with at least some of Ranunculus flammula and Mentha
the herbs listed above. aquatica.
This, the more sharply-defined This is the typical sub-community
sub-community, prevails in of the South-West Peninsula, but
Scotland and is common in Wales. is also found through the range
of M23.

51
M24 Molinia caerulea – Cirsium Mentha aquatica, Prunella vulgaris, Ranunculus
acris, Hydrocotyle vulgaris, and the scramblers
dissectum fen-meadow Vicia cracca and Lathyrus pratensis.
Coarser grasses are often prominent, with
This community includes the bulk of the Molinia Holcus lanatus and Anthoxanthum odoratum
caerulea vegetation in the lowland south-east of most frequent and Festuca rubra, Deschampsia
Britain. Molinia is always the dominant to the cespitosa and Agrostis stolonifera less common,
extent that associates may be reduced to scattered though sometimes abundant. There can also be
individuals of only a few species. Often however some sedges present, the most common being
there are a considerable number of associates. In smaller species such as Carex panicea, which is a
structural terms the most important species are community constant, and less frequent C. hostiana
other monocotyledons. Through most of the and C. pulicaris. Bryophytes are generally few and
central and eastern part of the range where this of low cover.
community is often found in association with This is a community of moist to fairly dry peats
fens, Juncus subnodulosus is the characteristic and peaty mineral soils which are circumneutral,
rush with J. articulatus and J. inflexus sometimes generally having a pH within the range 5-6.5. It
also present. To the south and west where the can be found in association with both soligenous
community often develops among heath vegeta- and topogenous mires, typically marking out the
tion, J. acutiflorus and J. conglomeratus become better-drained fringes of bogs and fens, or the mar-
frequent. gins of wet hollows and flushes. This community
The associated flora of dicotyledons helps to is widespread through the lowland south of
distinguish this community, although it is often Britain but has become increasingly local with
difficult to separate it from Juncus subnodulosus – changes in agricultural practice.
Cirsium palustre fen meadow (M22) and Juncus Although climate and soil together both
effusus/acutiflorus – Galium palustre rush pasture influence the floristics of this community it is
(M23) when these contain Molinia. Cirsium palus- essentially a secondary vegetation type, derived
tre and Angelica sylvestris are both very frequent from a variety of wetland vegetation types and
and Filipendula ulmaria and Centaurea nigra maintained by mowing or grazing. In the
can also be common. More strictly limited are absence of any kind of treatment all the stands of
Valeriana dioica, Succisa pratensis, and Cirsium the community would probably progress to
dissectum, although only the last species is pref- scrub or woodland. It has been reduced in extent
erential. Other common species of wide occur- by agricultural reclamation. Other stands have
rence are Potentilla erecta, Lotus uliginosus, become rank and scrubby with neglect.

52
 M24

Molinia caerulea generally dominant Juncus subnodulosus absent, but

with Juncus subnodulosus common J. acutiflorus and J. conglomeratus
and frequent records for some of common, J. effusus occasional. Species
Valeriana dioica, Galium uliginosum, listed opposite all scarce but Erica
Centaurea nigra, Vicia cracca, tetralix, Calluna vulgaris, Galium
Filipendula ulmaria and palustre and Dactylorhiza maculata
Equisetum palustre. frequent.

Phragmites australis constant, Eupatorium cannabinum and

Cladium mariscus quite common, Phragmites australis only occasional.
both typically subordinate in cover Cladium mariscus and Lythrum
to Molinia caerulea. Eupatorium salicaria usually absent.
cannabinum and Lythrum salicaria
frequent. M24c

M24a M24b Juncus acutiflorus – Erica tetralix

sub-community
Eupatorium cannabinum Typical sub-community
sub-community Rushes are a common feature, most
Molinia caerulea is often found with usually Juncus acutiflorus and J. con-
One of the most conspicuous species smaller amounts of Juncus subnodulo- glomeratus. There is frequently some
here, even when sparse, is Phragmites sus or J. articulatus.Smaller grasses are Erica tetralix, less often Calluna vul-
australis. The best single preferential well represented; Holcus lanatus and garis and Ulex gallii, so that the vegeta-
is Eupatorium cannabinum which Anthoxanthum odoratum are frequent tion looks more like a heath. However
provides continuity with tall-herb fen; and Briza media is strongly preferen- Holcus lanatus, Anthoxanthum odora-
less common are Lythrum salicaria tial. Sedges are common with Carex tum, less often Festuca rubra and
and Lysimachia vulgaris. Cirsium panicea and C. hostiana showing a Agrostis stolonifera are intermixed with
palustre, Angelica sylvestris and peak of occurrence, C. pulicaris espe- Molinia caerulea with frequent Carex
Filipendula ulmaria are also common cially frequent and C. nigra preferential panicea and a range of small herbs.
and, as reed or sedge cover becomes and sometimes in abundance. Succisa Valeriana dioica, Centaurea nigra and
thinner, plants such as Succisa pratensis, Cirsium dissectum, C. palus- Filipendula ulmaria and the tall-fen
pratensis, Cirsium dissectum, tre and Angelica sylvestris are all very herbs of M24a are not represented.
Centaurea nigra and Equisetum common together with the species list- Bryophytes are not conspicuous,
palustre increase in frequency. ed above. Sub-shrubs are typically though a number of species
Bryophytes can be more conspicuous sparse and bryophytes are poorly have been recorded.
in this sub-community, but species are represented. This is the most usual type of M24 in
only few, with Campylium stellatum south-western Britain.
This is the most common sub-commu-
joining Calliergon cuspidatum
nity in central and eastern England.
and Brachythecium rutabulum
as a distinctive preferential.
This sub-community is mainly
confined to East Anglia.

53
M25 Molinia caerulea –
frequency throughout and Holcus lanatus is fairly
Potentilla erecta mire common. Among the dense herbage, bryophytes
are sparse.
Molinia caerulea is the most abundant species This mire is a community of moist, but well-
found in this community The associated flora is aerated, acid to neutral peats and peaty mineral
poor, and most common are rushes and a few soils in the wet and cool western lowlands of
dicotyledons. Among the former, Juncus Britain. It occurs over gently-sloping ground,
acutiflorus and J. effusus are the most frequent. marking out seepage zones and flushed margins
Juncus articulatus and J. subnodulosus are both of sluggish streams, water-tracks and topogenous
occasional, and J. conglomeratus is very scarce. mires, but also extends onto the fringes of
The only constant dicotyledon is Potentilla erecta. ombrogenous mires. Soil and drainage conditions
Lotus uliginosus, Succisa pratensis, Cirsium of this community have similarities to those of
palustre and Angelica sylvestris are sparse and M23 and M24 and geographically this communi-
occasionally there can be some Eupatorium ty can be seen as a northern/western replacement
cannabinum or Filipendula ulmaria. Cirsium dis- of M24. It is especially frequent in south-west
sectum is very rare and its presence separates the England, Wales, and southern Scotland.
Molinia caerulea – Cirsium dissectum mire (M24) Although both climate and soils influence the
from this community. Also, since the soil pH is composition of the vegetation, treatments such as
generally acidic, plants such as Carex hostiana, burning, grazing and drainage are likely to be
C. pulicaris and Briza media, frequent in M24, are largely responsible for the development of this
of very limited occurrence here. Occasionally sub- community over ground that would naturally
shrubs can be quite common, particularly Calluna carry some other kind of mire or wet heath vegeta-
vulgaris and Erica tetralix. Ulex gallii can also be tion. Tracts of this community have been replaced
occasional in Wales and south-west England, and by coniferous plantations, particularly in the
U. europaea occurs in some stands. Myrica gale is upland fringes of the north-west. Elsewhere in the
local but can be quite extensive and dense. lowlands, other tracts of the community together
Grasses are limited in importance but Agrostis with neighbouring vegetation have been lost to
canina and A. stolonifera can be found at low agricultural improvements.

54
 M25

Erica tetralix constant, Calluna Juncus acutiflorus and occasionally Species listed opposite usually all
vulgaris and Myrica gale quite frequent, J. effusus patchily prominent in a occasional but tall herbs prominent
with Eriophorum angustifolium com- grassy community with frequent among Molinia caerulea and rush
mon and occasional Narthecium Holcus lanatus, Festuca rubra, clumps, with frequent Angelica
ossifragum, Drosera rotundifolia and Anthoxanthum odoratum, Agrostis cap- sylvestris and Cirsium palustre.
Vaccinium oxycoccos. illaris, Danthonia decumbens, Luzula Epilobium palustre, Eupatorium
multiflora and L. campestris. Erica cannabinum, Filipendula ulmaria,
tetralix rare but Calluna vulgaris Galium palustre and Mentha aquatica
is occasional. are occasional. Schoenus nigricans
can be locally abundant.

M25a M25b M25c

Erica tetralix sub-community Anthoxanthum odoratum Angelica sylvestris sub-community

sub-community
This is the widely distributed type of This is the most local sub-community,
M25. Molinia caerulea is the dominant Although Molinia caerulea is still but also the most striking sub-
but the associated flora is shifted dominant, the sward is shorter and community, developed on moist ground
towards that of Erica tetralix wet heaths more varied. Apart from the rushes and with freedom from grazing. Molinia
(M15-16). Erica tetralix is strongly pref- grasses mentioned above other associ- caerulea is vigorous, but variegated by
erential, with frequent Calluna vulgaris. ates can be sparse. Calluna vulgaris clumps of Juncus acutiflorus and
Juncus acutiflorus remains common but and Ulex galliiare occasional, as are J. effusus. Taller dicotyledons are
is joined by Eriophorum angustifolium. U. europaeus and Rubus fruticosus. common: Succisa pratensis and
Apart from sparse Anthoxanthum Erica tetralix and Myrica gale are very Lotus uliginosus are notable together
odoratum, Festuca rubra and Agrostis uncommon. Succisa pratensis, Lotus with the preferential species listed
canina, grasses are thin and taller herbs uliginosus and Cirsium palustre are above and also Pulicaria dysenterica,
are poorly represented. Among smaller all more frequent than in M25a and Valeriana officinalis and Centaurea
plants, Viola palustris and Hydrocotyle Serratula tinctoria and Rumex acetosa nigra. Shorter species include Mentha
vulgaris are sometimes present, but are weakly preferential, but all tend to aquatica, Cardamine pratensis and
with occasional Narthecium be grazed to rosettes. Smaller herbs Equisetum palustre with frequent
ossifragum, Drosera rotundifolia and bryophytes are poorly represented. Galium palustre. Bryophytes are again
and Vaccinium oxycoccos. Bryophytes sparse but Calliergon cuspidatum and
This sub-community is scattered
are distinctive, with Aulacomnium C. giganteum form scattered patches.
throughout the range of M25 but is
palustre, Polytrichum commune,
particularly frequent in Wales. This sub-community is found mainly
Hypnum jutlandicum and Calypogeia
in south-west England and south-west
fissa all preferential. Sphagnum spp.
Wales.
are noticeable, forming patches;
Sphagnum recurvum and S. auricula-
tum are the commonest species.
This sub-community can be found
throughout the range of M25.

55
M26 Molinia caerulea –
europaeus. Also common are Sanguisorba offici-
Crepis paludosa mire nalis, Angelica sylvestris, Centaurea nigra, Leon-
todon hispidus, Geum rivale and Lychnis flos-
This community is well-defined by a block of cuculi. Less conspicuous, but also frequent, is
constants and frequent companions but also Equisetum palustre. Potentilla erecta, Ranunculus
shows considerable variation in associated flora. acris, and Anemone nemorosa are common.
Stands range from swamp to those having a rank, Bryophytes are only prominent in exceptional
grassy character. Molinia caerulea is almost cases, with Calliergon cuspidatum most frequent.
always present, being the commonest dominant. This is a very local community of moist,
Carex nigra is also a constant, often as prominent moderately base-rich and calcareous peats and
tufts which can exceed Molinia in cover. Carex peaty mineral soils in the sub-montane northern
panicea can be abundant and C. pulicaris is com- Pennines. It represents a northern and altitudinal
mon. In stands transitional to swamp, C. approp- extreme of the richer kind of Molinia – tall herb
inquata or C. rostrata are present. In the Festuca vegetation. Stands are rare but all occur around
sub-community, by contrast, it is taller rushes and the northern Pennine uplands and the Lake
grasses which, with Molinia, form the bulk of the District between 250 m and 450 m altitude.
cover. Juncus acutiflorus may form dense patches This community is an apparently stable
and J. conglomeratus and J. articulatus both occur component of topogenous sequences around open
occasionally. waters and mires, but where it occurs on flushed
Hemicryptophyte dicotyledons are an impor- slopes, grazing often maintains the community
tant structural element among the Molinia, sedges and prevents progression of the community to
and rushes. Most frequent are Succisa pratensis, scrub or woodland. Drainage and sward improve-
Filipendula ulmaria, Valeriana dioica, Cirsium ment have probably destroyed many smaller
palustre and Caltha palustris together with the stands of this community and contributed to its
northern species Crepis paludosa and Trollius very local distribution.

56
 M26

Molinia caerulea and Carex nigra both Carex nigra often subordinate to
abundant, with locally prominent Molinia caerulea in more grassy or
C. appropinquata or C. rostrata. rushy vegetation, with frequent and
Sanguisorba officinalis, Angelica abundant Festuca rubra, F. ovina,
sylvestris, Serratula tinctoria, Galium Holcus lanatus, Briza media,
palustre and G. uliginosum all frequent; Deschampsia cespitosa, Anthoxanthum
bryophytes patchy. odoratum, Juncus acutiflorus and
J. conglomeratus; associates listed
opposite all occasional.

M26a M26b

Sanguisorba officinalis Festuca rubra sub-community

sub-community
This sub-community appears more
This is generally the less species-rich grassy and is usually developed on
sub-community. Molinia caerulea drier soils. Molinia caerulea is the
and Carex nigra are usually the most usual dominant, forming the bulk of a
abundant plants, with one or both rough sward together with the grasses
dominant in a swamp vegetation with listed above. Sedges are also common;
large sedges as above. Smaller sedges Carex nigra and C. panicea can both
can also occur, Carex panicea and show high cover and C. flacca
C. pulicaris being common. Apart from and C. pulicaris can be frequent.
Agrostis stolonifera, grasses are poorly Rushes are also common; Juncus
represented and among small herbs acutiflorus and J. conglomeratus
only Potentilla erecta, Ranunculus are preferential and J. articulatus
acris and Anemone nemorosa occur also occurs. Taller dicotyledons
with any frequency. The most common remain frequent, although Sanguisorba
tall herbs are listed above. Bryophytes officinalis, Angelica sylvestris and
are patchy but better-developed than Serratula tinctoria are all scarce. Geum
in M26b. Calliergon cuspidatum, rivale, Centaurea nigra and Leontodon
Lophocolea bidentata s.l., Thuidium hispidus are more common than in M26a.
tamariscinum and Campylium stella- In shorter swards, Prunella vulgaris,
tum are frequent, with Ctenidium Plantago lanceolata and Trifolium
molluscum, Plagiochila asplenioides, repens can be found. Bryophytes
Campylium elodes and Aulacomnium are often poorly represented, but
palustre being preferential. Calliergon cuspidatum, Pseudo-
scleropodium purum and Lophocolea
This sub-community is found most
bidentata remain frequent.
extensively at Sunbiggin and
Malham Tarns. This sub-community has a scattered
distribution through the dales along
the upland fringes.

57
M27 Filipendula ulmaria – with Juncus effusus the most common. Molinia
caerulea can also be found occasionally. Among
Angelica sylvestris mire smaller dicotyledons there can be Ranunculus
repens, Mentha aquatica, Lotus uliginosus, and
Although Filipendula ulmaria is frequent and Caltha palustris with more occasional Ranun-
locally abundant in a variety of vegetation types, culus acris, Cardamine pratensis, C. flexuosa,
in this community it forms the overwhelming Potentilla anserina, and Polygonum hydropiper.
dominant and the only constant. The dominants of Bryophytes are few in number and of low cover.
other communities in which it occurs, tall helo- This community is typically found where moist,
phytes, bulky sedges, rushes and rank grasses are, reasonably rich, circumneutral soils occur in
if present, all subordinate in this community. In situations protected from grazing. It can be found in
the deep shade cast by Filipendula only scattered both topogenous and soligenous mires and is espe-
individuals or dispersed clumps of other species cially typical of silted margins of slow-moving
are found. The commonest accompanying tall streams and soakways, the edges of flushes and
herbs are Angelica sylvestris, Valeriana officinalis damp hollows, and also of artificial habitats such as
and Rumex acetosa. In the Valeriana – Rumex along dykes and roadside ditches and around
sub-community they are often accompanied by ponds. This community occurs throughout lowland
Lychnis flos-cuculi, Succisa pratensis, Geum Britain.
rivale and sprawling Galium palustre. In the Both draining and grazing have reduced the
Urtica – Vicia sub-community these species are extent of this community to small remnants in
more scarce and Urtica dioica is very common many places. The community cannot tolerate
with Cirsium arvense, Epilobium hirsutum, any other than very light or sporadic grazing
Eupatorium cannabinum and Vicia cracca. At low and so stands often only persist outside enclo-
frequency throughout there can be scattered sures, and around unreclaimed mires and flush-
Lythrum salicaria, Rumex crispus, R. sangui- es. For example, this community can be found
neus, Epilobium palustre, Equisetum palustre, in wet field bottoms and edges that have been
E. arvense and E. fluviatile. fenced off, and alongside streams and ditches
There are few bulky monocotyledons; between pasture and boundaries. Progression to
Phragmites australis can be common and Phalaris woodland, even in the absence of treatments
arundinacea is found occasionally. Rushes are few such as grazing or mowing, appears to be slow.

58
 M27

Angelica sylvestris, Valeriana Associates listed for M27a at most Associates listed opposite at most
officinalis, Rumex acetosa, Lychnis occasional, but sparse Phragmites aus- occasional but Juncus effusus and
flos-cuculi, Succisa pratensis and tralis common, often with prominent Holcus lanatus are constant, and
Geum rivale are common among taller clumps of Urtica dioica, Eupatorium Juncus acutiflorus and Molinia caerulea
associates, with Caltha palustris, cannabinum and Epilobium hirsutum. occasional in ranker swards with
Ranunculus flammula, R. repens, The sprawling herbs Galium aparine Anthoxanthum odoratum, Agrostis
R. acris, Cardamine flexuosa and and Vicia cracca are common. stolonifera, Mentha aquatica and
C. pratensis occasional to frequent Lotus uliginosus quite common.
below and Galium palustre, G. uligi-
nosum and Lathyrus pratensis climbing
or sprawling.

M27a M27b M27c

Valeriana officinalis – Rumex acetosa Urtica dioica – Vicia cracca Juncus effusus – Holcus lanatus
sub-community sub-community sub-community

Filipendula ulmaria is abundant Filipendula ulmaria and a variety Filipendula ulmaria is still the most
and dominant. The vegetation may be of tall herbs again provide the main abundant species, but other tall herbs
species-poor, but overall it is charac- structural element. Angelica sylvestris, such as Angelica sylvestris, Valeriana
terised by a variety of associates. Most Cirsium palustre and Lythrum salicaria officinalis, Cirsium palustre and
common are Angelica sylvestris and are occasional, but others present in Rumex acetosa occur occasionally.
Valeriana officinalis, both sometimes M27a, e.g. Valeriana officinalis and Of greater importance, rushes and
locally abundant, with Rumex acetosa, Rumex acetosa, become scarce or grasses may have moderate abundance.
Lychnis flos-cuculi, Succisa pratensis absent. Urtica dioica, however, is very Juncus effusus and Holcus lanatus are
and Geum rivale are less common. common and is found with occasional both constant and J. acutiflorus and
Among smaller herbs can be the species Eupatorium cannabinum and Molinia caerulea occasional, with a
listed above with Stellaria alsine and Epilobium hirsutum forming patches. range of other grasses. In some stands
Ajuga reptans and Galium palustre as Scattered throughout can be Cirsium Mentha aquatica and Lotus uliginosus
the most common sprawling plant. arvense and Centaurea nigra. are frequent. In others Iris pseudacorus
Apart from Poa trivialis, preferential Phragmites australis can be common; and/or Oenanthe crocata can be
here, grasses and rushes are infrequent. alternatively there may be tussocks prominent.
Carex rostrata is quite common and can of Arrhenatherum elatius and some
The sub-community is western
occur with Menyanthes trifoliata and Holcus lanatus or scattered clumps of
in distribution.
Potentilla palustris. Bryophytes rushes. Smaller herbs and bryophytes
are better developed in this sub- are few and sparse.
community, with Brachythecium
This sub-community is found in
rutabulum the most common species.
central, southern and eastern Britain.
This sub-community is the usual form
in northern England and in southern
and eastern Scotland.

59
M28 Iris pseudacorus –
marsh transitions Atriplex prostrata and Matri-
Filipendula ulmaria mire caria maritima may be frequent with Samolus
valerandi, Oenanthe lachenalii and halophytic
In its typical form this is a luxuriant and herbs. Bryophytes are few, with Eurhynchium
species-rich community with Iris pseudacorus praelongum being the commonest throughout.
more abundant than Oenanthe crocata, although This community is confined to moist, more
both are constants, except in the far north of nutrient-rich soils along the oceanic seaboard of
Scotland where O. crocata is not found. Other tall Britain. It is especially characteristic of the fresh-
herbs are nearly always present, though only water seepage zone along the upper edge of salt-
Lycopus europaeus, Rumex crispus and Scut- marshes in the sheltered sea-lochs of western
ellaria galericulata are frequent throughout the Scotland. Other situations in which it occurs are
community. Other species such as Rumex acetosa, over stabilised shingle down the west coast and
Lychnis flos-cuculi, Angelica sylvestris, Valeriana in wetter hollows and flushes on raised beach
officinalis, Cirsium palustre, C. arvense and Urtica platforms and gentle cliff slopes. The community
dioica are often common and conspicuous but is the oceanic counterpart of the Filipendula
preferential to particular sub-communities. ulmaria – Angelica sylvestris mire (M27) and is
Rushes and grasses are frequently important. largely confined to the west coast of Britain. In
Juncus effusus and J. acutiflorus are common, as particular it is found in west Scotland from
are Poa trivialis and Agrostis stolonifera. There are Orkney and Shetland southwards, with scattered
a variety of smaller dicotyledons. Some typically stands in south-west England and west Wales.
occur as scattered plants, for example Ranunculus The community was probably once much more
acris, Caltha palustris, Stellaria alsine, Mentha widespread in south-west England and west Wales
aquatica and Hydrocotyle vulgaris, while Ranun- but it may have been largely destroyed in its salt-
culus repens and Potentilla anserina form mats marsh habitat by human interference with the tran-
with high local cover. On patches of wet and open sitional upper zones. The community, where it
ground, annuals may be prolific, such as Poly- does occur, is often not heavily grazed and it
gonum hydropiper, Montia fontana, and on cattle- appears to be a relatively stable vegetation type
poached mud, Ranunculus sceleratus. On salt- with only a slow progression to scrub or woodland.

60
 M28

Juncus effusus and/or J. acutiflorus Species listed opposite occasional at Groups of species opposite, and even
constant and patchily abundant with most, but Urtica dioica and Cirsium Filipendula ulmaria, infrequent in
frequent Rumex acetosa, Cirsium arvense constant with Galium aparine rather open vegetation with Atriplex
palustre, Epilobium palustre, Lychnis and occasional to frequent Elymus prostrata and Samolus valerandi com-
flos-cuculi, Ranunculus acris, Caltha repens, Stellaria media, Arrhenatherum mon and sporadic records for maritime
palustris, Lotus uliginosus and elatius and Dactylis glomerata. plants.
Galium palustre.

M28a M28b M28c

Juncus spp. sub-community Urtica dioica – Galium aparine Atriplex prostrata – Samolus
sub-community valerandi sub-community
This is the richest sub-community in
which other dicotyledons, rushes and The vegetation here, although as Although Iris pseudacorus and
grasses form a consistent associated tall and luxuriant as in M28a, is less Oenanthe crocata can be abundant,
flora. Iris pseudacorus is generally a species-rich. Iris pseudacorus is still taller associates are generally lacking;
clear dominant although both Oenanthe dominant and both Oenanthe crocata even Filipendula ulmaria is scarce and
crocata and Filipendula ulmaria can be and Filipendula ulmaria remain fre- Lycopus europaeus and Rumex crispus
patchily abundant with the above quent, but apart from Lycopus euro- are only present as scattered plants.
Juncus spp. Among the taller herbs, paeus and Scutellaria galericulata, the Among smaller plants the commonest
Lycopus europaeus, Rumex crispus, only other common taller dicotyledons grasses are Agrostis stolonifera and
Scutellaria galericulata, Angelica are Urtica dioica and Cirsium arvense Festuca rubra. The most common pref-
sylvestris and Valeriana officinalis, which can be abundant. Other taller erentials are plants tolerant of saline
with the species listed above, all occur species of M28a are occasional or habitats, including Atriplex prostrata,
frequently to occasionally. Common scarce and Galium aparine replaces the most common, with Samolus
grasses are Festuca rubra, Holcus lana- G. palustre. Grasses are often conspicu- valerandi, Oenanthe lachenalii,
tus, Anthoxanthum odoratum, Poa ous with Poa trivialis and Agrostis Matricaria maritima, Triglochin
pratensis and Elymus repens forming stolonifera very common as patchy maritima and Glaux maritima.
scattered tussocks, and with Carex carpets interspersed with the other
otrubae and trailing Galium palustre. species listed above.

61
M29 Hypericum elodes – M30 Related vegetation of
Potamogeton polygonifolius seasonally-inundated habitats
soakway
Other vegetation of the same type as Hypericum
elodes – Potamogeton polygonifolius soakway
This community has a very distinctive appear- (M29), and characteristic of similar, seasonally
ance, typically consisting of low creeping or float- inundated habitats, with rather base-poor and
ing mats of Hypericum elodes and Potamogeton only moderately enriched waters, has only been
polygonifolius. Very often, unless the ground has poorly sampled. Some examples, lacking
been badly trampled by grazing animals, these two Hypericum elodes but otherwise the same as
constants are set in a carpet of submerged M29, may be regarded as impoverished stands of
Sphagnum auriculatum, sometimes with S. cuspi- that community, though it must be noted that
datum, S. palustre or S. recurvum. Other bryo- species such as Potamogeton polygonifolius,
phytes are sparse but Polytrichum commune or Eriophorum angustifolium, Juncus bulbosus/
Aulacomnium palustre, and Drepanocladus exan- kochii and Sphagnum auriculatum also occur
nulatus, D. revolvens, and Calliergon cuspidatum, with some frequency in bog-pool and poor-fen
may be locally abundant. vegetation.
Other vascular plants are scattered. Juncus There are also stands in which Eleocharis
bulbosus and Ranunculus flammula are the only multicaulis is strongly dominant with little or no
constants but Hydrocotyle vulgaris, Anagallis Hypericum elodes or Potamogeton polygonifolius;
tenella, Drosera rotundifolia, Narthecium ossi- these look similar to the Eleocharitetum multi-
fragum, and Galium palustre can all be caulis recorded from Eire and elsewhere in west-
moderately frequent, along with sedges such as ern Europe. Deschampsia setacea is listed as a
Carex demissa, C. echinata, C. panicea and characteristic species of such vegetation, and in
C. nigra. There can also be sparse shoots or small Britain this rare species is typical of this type of
patches of Molinia caerulea, Agrostis canina ssp. habitat. Scirpus fluitans can also be found domi-
canina, Juncus articulatus, J. effusus, J. acuti- nating in swards which lack some of the
florus, Eleocharis multicaulis, Eriophorum angus- typical plants of M29, and in the New Forest and
tifolium and Rhynchospora alba. Carex rostrata Cornwall, Baldellia ranunculoides is a frequent
can also be found in some stands. Two rare species and conspicuous component of low-growing
associated with this community are Galium vegetation in seasonally wet pools.
debile, in the New Forest, and the fern Pilularia All these vegetation types, along with M29,
globulifera. have been grouped in the Hydrocotylo-Baldellion
This community is characteristic of shallow alliance, comprising assemblages of mesotrophic
soakways and pools in peats and peaty mineral to oligotrophic, and periodically fluctuating
soils with fluctuating water levels, such as waters.
seepages and runnels around mires and in
heathland pools, at moderate altitudes. The water No sub-communities.
is typically clear, still or gently-flowing, moder-
ately acid to neutral, with a pH between 4 and 5.5,
and probably quite oligotrophic. This vegetation is
confined to the warm oceanic parts of Britain and
extends in a well-defined zone from west Surrey
through the New Forest to the South-West
Peninsula and north through Wales to Galloway. It
may well be found further north, following the
distribution of Hypericum elodes.
This soakway appears to be a stable vegetation
type in the absence of nutrient enrichment. In
situations where there is some nutrient enrich-
ment, grazing and trampling may help continually
set back any tendency to succession. Trampling by
grazing animals can also play a part in keeping the
vegetation open and varied, although heavy
poaching can be deleterious to the Sphagnum
carpet.

No sub-communities.

62
M31 Anthelia julacea –
Saxifraga stellaris. Less frequent are Eriophorum
Sphagnum auriculatum spring angustifolium, Carex bigelowii, C. nigra, Festuca
vivipara, Agrostis canina, A. stolonifera, Juncus
In this community Anthelia julacea forms bulbosus, Thalictrum alpinum, and Viola palustris.
cushions up to a metre or more in thickness and This is a montane community typical of often
several square metres in extent. Its associated flora skeletal mineral and organic soils kept more or
is species-poor and vascular plants are sparse. less permanently wet by the trickling of acid
Among other bryophytes, Sphagnum auriculatum, and oligotrophic waters, of pH 4.5-5.0, frequently
Marsupella marginata and Scapania undulata are derived at higher altitudes from snow-melt. It
all constant, the first of which can form prominent occurs at moderate to high altitudes, from about
patches. Racomitrium lanuginosum and Philonotis 400 m to over 1000 m, marking out areas of less
fontana are also frequent with occasional vigorous seepage among tracts of montane grass-
Calliergon sarmentosum, Campylopus atrovirens, lands and grass-heaths or among snow-bed vegeta-
Polytrichum commune and Racomitrium fascicu- tion. It is a local but widespread community found
lare. Rare bryophytes which have been recorded through much of Scotland, the Lake District
in this community include Anthelia juratzkana, and Snowdonia.
which almost totally replaces A. julacea in some Anthelia julacea – Sphagnum auriculatum
stands, and Pohlia ludwigii. spring is an essentially stable community in the
The commonest vascular plant is Deschampsia harsh environment in which it characteristically
cespitosa. Scattered plants of Nardus stricta occur occurs.
quite often with occasional Narthecium ossi-
fragum, Pinguicula vulgaris, Carex demissa, and No sub-communities.

63
M32 Philonotis fontana – This is a community of springs and rills at
moderate to high altitudes, above 450 m to over
Saxifraga stellaris spring 1000 m, where there is irrigation with circum-
neutral and oligotrophic waters with a pH of
These bryophyte-dominated springs, flushes and around 4.5-6. This is one of the most common
rills are striking in appearance. Philonotis fontana is and widespread types of spring vegetation in the
usually dominant and obvious by its fresh-green uplands of north-west Britain and is dependent
colour. The golden-green Dicranella palustris and on sustained and vigorous irrigation by ground-
Scapania undulata are often also abundant. These waters, common in the wetter parts of the coun-
species together often form cushions or plush mats. try. It marks out permanent springs of a well-
Less consistent but sometimes prominent are defined character, also diffuse flushes and seep-
Sphagnum auriculatum, Scapania uliginosa, Calli- age lines, rills and small streams and occasional-
ergon sarmentosum, Drepanocladus exannulatus, D. ly steep, dripping ground. In some places snow-
fluitans and the characteristic, but not very frequent, melt may be an important water source. It is
Jungermannia exsertifolia. Bryum pseudotriquetrum found on a variety of waterlogged soils from frag-
is also common, but not as consistent as in more mentary accumulation of silt among stones to
base-rich springs, and Cratoneuron commutatum flushed peats and gleys. The community is
and C. filicinum are scarce. common through the Scottish Highlands, the
The associated vascular flora is rather varied in Southern Uplands, the Lake District and north
composition and cover and typically species-poor. Wales and over the non-calcareous parts of the
The only constant is Saxifraga stellaris, but Pennines. It can also be found as fragmentary
Deschampsia cespitosa (with ssp. alpina at higher stands at lower altitudes without the montane
altitudes) is often found in small quantities with element in the vegetation, particularly at the
scattered Stellaria alsine. More occasional are southern limit of the range of this community.
Festuca rubra, Anthoxanthum odoratum, Agrostis The harsh montane environment has a striking
stolonifera and A. canina, all usually at low cover, influence on the composition of the community
with sparse Viola palustris, Nardus stricta, Carex and though stands can be grazed and trampled,
bigelowii and C. panicea. In stands which are climatic and soil conditions probably play the
perhaps less base-poor, Montia fontana and major part in maintaining the vegetation as an
Chrysosplenium oppositifolium become frequent effective climax. At lower altitudes, it could
and abundant, along with Caltha palustris, perhaps show some successional development in
Cardamine pratensis and a number of other ungrazed situations.
associated species.

64
 M32

Sphagnum auriculatum often abun- Sphagnum auriculatum and other

dant, with Calliergon sarmentosum bryophytes listed opposite very scarce
and Scapania uliginosa occasional and but Bryum pseudotriquetrum frequent.
locally prominent. Montia fontana Montia fontana and Chrysosplenium
and Chrysosplenium oppositifolium oppositifolium constant and sometimes
very occasional. abundant.

M32a M32b
Sphagnum auriculatum Montia fontana – Chrysosplenium
sub-community oppositifolium sub-community
In this species-poor sub-community Vascular plants are more numerous
the bryophyte mat consists of mixtures and varied, although bryophytes still
of Philonotis fontana, Scapania generally have dominance. Philonotis
undulata and Dicranella palustris with fontana, Dicranella palustris and
Sphagnum auriculatum strongly prefer- Scapania undulata are all very com-
ential and often abundant. Calliergon mon and each, especially the first, can
sarmentosum and Scapania uliginosa be abundant. Bryum pseudotriquetrum
are occasional and Polytrichum com- is frequent and Jungermannia exsertifo-
mune and Hygrohypnum ochraceum lia occasional. Among the vascular
occur at low frequencies. Among the plants, Montia fontana and
vascular plants only Saxifraga stellaris Chrysosplenium oppositifolium have
and Deschampsia cespitosa are con- high frequencies with Saxifraga stel-
stant but other grasses such as Agrostis laris, and are sometimes abundant.
stolonifera, A. capillaris, Anthoxanthum Along with Stellaria alsine there are
odoratum and Festuca rubra are quite often very small plants of Caltha palus-
frequent as scattered tufts. tris (ssp. minor) and Cardamine praten-
sis. Epilobium palustre can sometimes
This sub-community occurs mainly on
be found but more distinctive are the
the harder acidic quartzites and sand-
frequent occurrence of E. alsinifolium
stones of the north-west Highlands.
and E. anagallidifolium. Grasses can be
quite common; Deschampsia cespitosa
is often joined by Anthoxanthum odor-
atum, Agrostis canina and several other
grasses together with some sedges.
This community is associated with a
range of substrates slightly more base-
rich than those of M32a.

65
M33 Pohlia wahlenbergii var. M34 Carex demissa – Koenigia
glacialis spring islandica flush
In this community Pohlia wahlenbergii var. This is an open vegetation type with a
glacialis dominates in spongy carpets of a bright bryophyte-dominated carpet broken by areas of
apple-green colour, often of small extent, but wet, silty and stony ground. Scapania undulata,
exceptionally up to 200 m2. Few other bryophytes Calliergon sarmentosum and Blindia acuta are
occur with any frequency although P. ludwigii is a all common and each can be abundant, with occa-
constant. Philonotis fontana can be prominent, sional patches of Dicranella palustris, Philonotis
although not with the high cover found in fontana, Drepanocladus revolvens, Marsupella
Philonotis fontana – Saxifraga stellaris spring aquatica and Sphagnum auriculatum. Scattered
(M32). Other bryophytes recorded occasionally through this and over the rills are plants of Carex
are Hygrohypnum luridum, Bryum weigelii, Calli- demissa, Koenigia islandica, Deschampsia cespi-
ergon stramineum, Scapania undulata, S. uligi- tosa, Saxifraga stellaris, Juncus triglumis, J. bulbo-
nosa, Dicranella palustris and Marchantia sus and the rare J. biglumis and Sagina saginoides.
alpestris. All these are generally of low cover, though many
In this carpet there are only a few vascular plants. can show a measure of abundance and Koenigia,
Deschampsia cespitosa (presumably ssp. alpina) and although individual plants are small, can cover
Saxifraga stellaris are constant, but Cerastium ceras- quite a large ground area.
toides is quite often found and there can be Stellaria This community occurs on ground which is
alsine, Chrysosplenium oppositifolium, Epilobium kept periodically moist by circumneutral and
anagallidifolium, Veronica serpyllifolia var. humi- oligotrophic waters. Typically it is found in open
fusa, and Rumex acetosa. Other rare plants found silty or stony flushes fed by vigorous seepage from
occasionally are Epilobium alsinifolium, Alopecurus springs issuing at moderately high altitudes,
alpinus and Phleum alpinum. over 500 m, from basalt. In their base status,
This community is strictly confined to with pH values around 6.0, and their low cation
spring-heads associated with late snow-beds content, the waters are similar to those which feed
where there is vigorous irrigation by cold waters. the Philonotis fontana – Saxifraga stellaris spring
The flushing waters, and often sloppy, ill-struc- community (M32), which often occupies the
tured mixtures of mineral and organic matter spring-heads above the flushes. The community is
beneath the moss carpet, are base-poor and confined to Skye, where it occurs scattered along
oligotrophic. Although Pohlia wahlenbergii var. the Trotternish Ridge extending several kilometres
glacialis occurs over quite a range of altitudes north of the Storr. It forms one of several
through the uplands of Wales, Cumbria and communities in which Koenigia can be
Scotland, it is only found with the kind of domi- found.
nance characteristic here within the high montane
zone at altitudes generally above 850 m. Within No sub-communities.
this area, which includes the central and north-
western Highlands of Scotland, the community is
further restricted to situations where the snow
lies longest, especially on north- and east-facing
slopes.
The general climatic and edaphic features
determine the overall character of this community
with its cold-tolerant plants and montane species.

No sub-communities.

66
M35 Ranunculus omiophyllus – M36 Lowland springs and
Montia fontana rill streambanks of shaded situations
These rills typically have a rather crowded, though There is a clear contrast, among the Cardamino –
not always continuous, cover of vascular plants and Montion springs and flushes of acid to circum-
bryophytes. Much of the growth is often submerged neutral habitats, between the upland communities
in the shallow waters, with a floating or shortly which have been described, where Montia
emergent canopy. Ranunculus omiophyllus is often fontana, Saxifraga stellaris and Philonotis fontana
abundant, frequently with Montia fontana. Floating are conspicuous, and the vegetation of lowland
leaves of Potamogeton polygonifolius are common- and often shaded situations. In these, Chryso-
ly prominent and there can be local patches of splenium oppositifolium occurs with bryophytes
Agrostis stolonifera, Glyceria fluitans, Juncus bul- such as Hookeria lucens, Rhizomnium puncta-
bosus, J. articulatus and Callitriche stagnalis, with tum, Trichocolea tomentella, Pellia epiphylla and
scattered plants of Ranunculus flammula, a con- Conocephalum conicum. This type of vegetation
stant, Myosotis secunda and Stellaria alsine. More has not been separately sampled here but it figures
occasional are Ranunculus repens, Equisetum in the field and ground layers of various wet
palustre, Hydrocotyle vulgaris, Galium palustre woodlands, notably the Alnus – Carex, Alnus –
and Lotus uliginosus. Juncus bufonius and Scirpus Urtica and Alnus – Fraxinus – Lysimachia types,
setaceus can sometimes be seen on open mud. where it is distinctive of seepage lines and damp
Bryophytes can contribute substantially to the stream banks, quite often with Cardamine flexu-
cover although there are only a few frequent osa, C. amara and Chrysosplenium alternifolium.
species. Sphagnum auriculatum is a constant and Similar mixtures of plants can be found widely
often grows semi-submerged with patches of through lowland Britain, especially in the wetter
Philonotis fontana but, apart from occasional west and around the upland fringes, along
Polytrichum commune, other species are sparse, stream-sides and wet banks, probably once
with only occasional records of Calliergon wooded, but where shade is now provided by tall
cuspidatum, C. stramineum, Drepanocladus exan- herbs or by the aspect of the site. These need
nulatus, D. vernicosus, Scapania irrigua and further sampling.
Rhytidiadelphus squarrosus.
This community is typical of spring-heads and No sub-communities.
rills at moderate altitudes in south-western
Britain, where there is irrigation by circumneutral
and probably quite oligotrophic waters. These are
typically rather base- and nutrient-poor with pH
values ranging from 4.5 to 6.5 over acidic rocks. It
has been recorded only from south-western
England, Wales, and around the Lake District. It
may occur throughout the range of R. omiophyllus
in Britain.

No sub-communities.

67
M37 Cratoneuron commutatum – Equisetum palustre, Chrysosplenium oppositifoli-
um, Poa trivialis, Carex panicea, C. nigra and
Festuca rubra spring C. dioica.
This is a community of ground kept permanent-
Cratoneuron commutatum occurs frequently in a ly moist by irrigation with base-rich, calcareous and
variety of calcareous mires, but here it is generally oligotrophic waters. It is dependent on
consistently dominant in large masses, often sustained irrigation common in areas of higher rain-
forming prominent mounds or banks. In some fall. Here it can be found marking out spring-heads,
stands of the same general floristic composition, seepage lines and drip zones in areas of lime-rich
C. filicinum accompanies or totally replaces it. bedrocks, where waters emerge along bedding
Other bryophytes can make a contribution, but planes or at junctions with impervious substrates.
typically a minor one. However, the constant Provided the ground is permanently wet, the com-
Bryum pseudotriquetrum is very common. munity can even occur on vertical surfaces and bare
Occasional species include Philonotis fontana, P. rock, forming curtain-like masses. The community
calcarea, Aneura pinguis, Pellia endiviifolia, can be found throughout the north-western uplands
Drepanocladus revolvens, Gymnostomum recurvi- of Britain with its more Arctic-Alpine element best
rostrum, G. aeruginosum, Brachythecium rivulare developed in the Scottish Highlands, with outliers
and Dicranella palustris. Very typically there is in the Lake District and Upper Teesdale. Springs
some tufa deposition allowing the mat to build dominated by Cratoneuron species also occur
into mounds. The vascular element is typically widely, but locally, in the British lowlands, and fur-
species-poor and of low total cover. There may be ther sampling of these is needed.
considerable variation in associated flora and, par- In most circumstances it is a permanent
ticularly where stands are developed on gently- community maintained by edaphic and climatic
sloping ground, a richer and more extensive layer conditions of the environment. On gentle slopes,
can be found, coming close to the Cratoneuron trampling by grazing stock or deer often plays an
commutatum – Carex nigra spring (M38). Often, important part in maintaining the characteristically
however the only species present are Festuca open conditions of flushed soils, but trampling
rubra, Cardamine pratensis and Saxifraga and grazing can have an adverse effect on the
aizoides, the last of which is absent from southern bryophyte carpet.
Scotland and Wales. Occasional herbs include
Agrostis stolonifera, Deschampsia cespitosa, No sub-communities.

68
M38 Cratoneuron commutatum – Juncus triglumis, J. articulatus, J. bulbosus,
Cera-stium fontanum, Prunella vulgaris, Caltha
Carex nigra spring palustris, Galium palustre, Equisetum palustre,
Achillea ptarmica, Cochlearia officinalis (often
This type of spring preserves the same pattern of ssp. alpina), Triglochin palustris, Ranunculus
dominance by Cratoneuron commutatum (again acris, Anthoxanthum odoratum, Festuca ovina,
occasionally supplemented or replaced by Epilobium anagallidifolium and, in north England,
C. filicinum) as in Cratoneuron commutatum – the introduced E. nerteroides. In Teesdale this
Festuca rubra spring (M37), but the associated community is the locus for Saxifraga hirculus.
flora is much richer. This is partly seen among This vegetation is confined to montane springs
the bryophytes. Bryum pseudotriquetrum and and flushes strongly irrigated by base-rich,
Philonotis fontana are the commonest and can calcareous and oligotrophic waters. As in M37,
have moderately high cover, and there are many sites of sustained irrigation with waters draining
others which can occur locally as prominent from lime-rich bedrock are marked out and tufa
patches. These include calcicolous species such as encrustation is often seen. It is very local around
Aneura pinguis, Fissidens adianthoides, Philo- Upper Teesdale in the north Pennines and in the
notis calcarea, Ctenidium molluscum, Cinclidium central Highlands of Scotland, mostly above
stygium, Drepanocladus revolvens and Campy- 650 m altitude.
lium stellatum. Although the harsh climatic and edaphic
The increased richness is most seen among the conditions exert a strong influence on the
vascular plants. Small sedges are noticeable. structure and composition of the vegetation, heavy
Carex demissa, C. nigra and C. panicea are grazing plays a major role in maintaining the
constant and often abundant, and C. pulicaris, distinctive richness of the community, and it is
C. flacca and C. dioica are common. There are this trampling and cropping by sheep and deer
frequent scattered plants of Cardamine pratensis, which is responsible for the most obvious floristic
Selaginella selaginoides, Leontodon autumnalis, differences between this community and M37.
Polygonum viviparum, Trifolium repens, Cirsium
palustre, Ranunculus flammula, Sagina nodosa, No sub-communities.

69
70
A KEY TO HEATHS

1. Calluna vulgaris 1a. Calluna vulgaris

may be present but is not constant and usually
constant or prominent in dominant in the
the vegetation. vegetation.

2. Ulex spp. constant in 2a. Ulex spp. absent or,

the sward and usually if present, then not con-
prominent. stant and subordinate in
the vegetation.

3. Sphagnum capillifolium 3a. Sphagnum capillifolium

constant within a luxuri- absent or, if present,
ant moss-carpet, usually sparse and infrequent.
over steep slopes with a
thin mineral soil.

4. At least two of 5. Racomitrium lanugi-

Racomitrium lanugi- nosum and the above
nosum, Cladonia species not constant.
arbuscula, C. uncialis, At least one of: Erica
heath communities

Huperzia selago, cinerea, Galium saxatile,

4 Dendrogram keys to

Empetrum nigrum Vaccinium myrtillus,

hermaphroditum and Festuca ovina,
Cetraria islandica Deschampsia flexuosa
constant in the vegeta- and/or Hypnum spp.
tion. Sub-montane and constant in the vegeta-
montane heaths. tion. Lowland and sub-
montane heaths.

See Fig 1. See Fig 2. See Fig 3. See Fig 4. See Fig 5.
 Fig 1. Key to heaths in which Calluna vulgaris is not constant or prominent but may be present at low frequency.

1. Erica vagans present 1a. Erica vagans absent.

and usually co-dominant Vaccinium myrtillus
with other species. and/or Empetrum
Heaths Figure 1

nigrum (either sub-

species) constant and
prominent. Montane
heaths in northern
Britain.
3a. Carex bigelowii
absent or infrequent,
occasionally locally
abundant. Vegetation
usually dominated by
Vaccinium myrtillus
2. Erica vagans co- 2a. Erica vagans co- 3. Carex bigelowii
with constant Galium
dominant with Schoenus dominant with Ulex constant and frequent.
saxatile, Deschampsia
nigricans, Molinia europaeus. Ulex gallii
flexuosa, Dicranum
caerulea, Erica cinerea and Erica cinerea
scoparium and
and E. tetralix; Ulex constant and frequent
Pleurozium schreberi.
gallii and Genista anglica often with Carex flacca,
sometimes frequent. Potentilla erecta and
Polygala vulgaris. Erica 4. Cladonia arbuscula 4a. Cladonia arbuscula
tetralix never prominent. constant and abundant. not constant but may be
Racomitrium lanugi- frequent. Empetrum
nosum generally subor- nigrum hermaphroditum
dinate. Empetrum and mats of bulky
nigrum hermaphroditum pleurocarpous mosses:
and bulky pleurocarpous Pleurozium schreberi,
mosses may be frequent Rhytidiadelphus loreus
but never abundant. Racomitrium
lanuginosum and
Hylocomium splendens
constant and abundant.

H5 Erica vagans – H6 Erica vagans – Ulex H19 Vaccinium myrtillus H20 Vaccinium myrtillus H18 Vaccinium myrtillus
Schoenus nigricans europaeus heath – Cladonia arbuscula – Racomitrium – Deschampsia flexuosa
heath heath lanuginosum heath heath

Typical of the east and Typical of the north-west

central Highlands Scottish Highlands
These communities are almost between the upper limit between the upper limit
entirely restricted to the Lizard of ericoid dominance of ericoid dominance
Peninsula in Cornwall. and summit moss-heaths. and summit moss-heaths

71
Heaths Figure 2

Fig 2. Key to heaths with constant Calluna vulgaris and in which Ulex spp. are constant and prominent

1. Ulex minor con- 1a. Ulex minor gener-

stant and usually ally absent. Ulex gallii
prominent. constant and promi-
nent.

2. Agrostis curtisii, 2a. Agrostis curtisii 3. Agrostis curtisii, 3a. Agrostis curtisii
Molinia caerulea, absent or present only Molinia caerulea, generally absent;
Erica cinerea and E. at low frequency. Erica cinerea and E. Erica tetralix never
tetralix Erica tetralix never tetralix frequent.
constant. frequent. constant. Ulex europaeus may
be abundant.
H3 Ulex minor – H2 Calluna vulgaris – H4 Ulex gallii –
Agrostis curtisii heath Ulex minor heath Agrostis curtisii heath
H8 Calluna vulgaris –
Ulex gallii heath

Restricted in its Found mainly in

occurrence to Dorset and south-west Britain.
Hampshire.

72
Heaths Figure 3

Fig 3. Key to heaths with constant Calluna vulgaris and Sphagnum capillifolium over a luxuriant bryophyte-rich carpet.
Constants include: Vaccinium myrtillus, Empetrum nigrum nigrum, Deschampsia flexuosa, Rhytidiadelphus loreus,
Pleurozium schreberi, Dicranum scoparium and Hylocomium splendens.

1. Vaccinium vitis- 1a. Vaccinium

idaea and Cladonia vitis-idaea and
arbuscula constant Cladonia arbuscula
in the vegetation. absent or at very low
Rubus chamaemorus frequency. Constant
frequent and may be Potentilla erecta. The
prominent. bryophyte carpet
includes Dicranum
majus and
Plagiothecium
undulatum.
H22 Vaccinium
myrtillus-Rubus
chamaemorus heath H21 Calluna vulgaris
– Vaccinium myr-
tillus – Sphagnum
capillifolium heath

Almost entirely Usually located on

confined in its steep, north-east to
distribution to the north-west facing
central and north- slopes on thin miner-
west Highlands. al soils, often with
rock outcrops.

73
74
Fig 4. Key to sub-montane and montane heaths with constant Calluna vulgaris and at least two constants from: Racomitrium
lanuginosum, Cladonia arbuscula, C. uncialis, Huperzia selago, Empetrum nigrum hermaphroditum and Cetraria islandica.

1. Juniperus communis 1a. Juniperus communis

ssp. nana or and Arctostaphylos alpi-
Arctostaphylos alpinus nus absent or at very low
Heaths Figure 4

constant, usually with frequency. Racomitrium

some Erica cinerea. lanuginosum and/or
Cladonia spp.
prominent.

3a Racomitrium
lanuginosum can be
frequent but is rarely
abundant. Cladonia
2. Vegetation dominated 2a. Juniperus communis 3. Racomitrium
spp. usually prominent
by Juniperus communis ssp nana usually absent. lanuginosum dominant,
including constant
ssp. nana with constant Arctostaphylos alpinus usually with constant
C. arbuscula, C. uncialis
Scirpus cespitosus, constant in a low open Cladonia uncialis.
and C. rangiferina.
Deschampsia flexuosa, mat. Also constant: Empetrum nigrum
Empetrum nigrum
Potentilla erecta, Erica Huperzia selago, (either sub-species),
hermaphroditum also
cinerea, Racomitrium Deschampsia flexuosa, Deschampsia flexuosa
constant. Deschampsia
lanuginosum, Cladonia Racomitrium lanugi- and Hypnum cupressi-
flexuosa and Hypnum
uncialis, Pleurozia nosum, Cladonia arbus- forme can be frequent.
cupressiforme occasional
purpurea. cula and C. uncialis. 4a. Racomitrium and sparse.
lanuginosum abundant
but not forming a dense
mat. Cladonia arbuscula
4. Racomitrium
absent. Erica cinerea
lanuginosum forms
constant.
a dense woolly mat
5-10 cm thick. Cladonia
arbuscula constant, Erica
cinerea may be frequent
but not constant.

H15 Calluna vulgaris – H17 Calluna vulgaris – H14 Calluna vulgaris – H10b Calluna vulgaris – H13 Calluna vulgaris –
Juniperus communis Arctostaphylos alpinus Racomitrium lanugi- Erica cinerea heath, Cladonia arbuscula
ssp. nana heath heath nosum heath Racomitrium lanugi- heath
nosum sub-community
 Fig 5. Key to lowland and sub-montane heaths with constant, usually dominant Calluna vulgaris and at least one of Erica cinerea, Vaccinium myrtillus,
Empetrum nigrum nigrum, Festuca ovina, Deschampsia flexuosa, Galium saxatile and Hypnum cupressiforme constant in the vegetation.

1. Scilla verna and 1a. Scilla verna and

Plantago maritima Plantago maritima
constant in short heaths absent.
around maritime cliffs.
Heaths Figure 5

2. Carex arenaria 2a. Carex arenaria and

constant but not often Ammophila arenaria
prominent. Ammophila absent.
arenaria frequent but
sparse. Sandy dunes
and coastal plains.

3. Arctostaphylos uva- 3a. Arctostaphylos

ursi and the following uva-ursi absent or rare.
species constant: Erica
cinerea, V. vitis-idaea,
Deschampsia flexuosa,
Cladonia impexa and 4. Erica cinerea constant 4a. Erica cinerea not
pleurocarpous mosses. and usually prominent. constant and present
only at low frequency.

5. Deschampsia flexuosa 5a. Deschampsia

constant. Vaccinium flexuosa absent or
myrtillus usually present only at
frequent. low frequency.
Calluna vulgaris
usually the only
6a. Calluna vulgaris abundant sub-
always dominant but shrub in a
6. Vaccinium myrtillus Vaccinium myrtillus, species-poor
constant. Constant and V. vitis idaea or heath. Other con-
frequent Dicranum Empetrum nigrum stants include
scoparium, Pleurozium nigrum may be locally Festuca ovina,
schreberi, Hypnum abundant. Pohlia Hypnum
cupressiforme and/or nutans is constant cupressiforme and
Hylocomium splendens. but other Dicranum scopar-
pleurocarpous mosses ium.
are only occasional.

H7 Calluna vulgaris – H11 Calluna vulgaris – H16 Calluna vulgaris – H10 Calluna vulgaris – H12 Calluna vulgaris – H9 Calluna vulgaris H1 Calluna vul-
Scilla verna heath Carex arenaria heath Arctostaphylos uva-ursi Erica cinerea heath Vaccinium myrtillus – Deschampsia flex- garis – Festuca
heath heath uosa heath ovina heath

75
5 Heath community
descriptions and
sub-community keys
H1 Calluna vulgaris – Festuca both pioneer and degenerate Calluna. In such sit-
uations Hylocomium splendens, Pleurozium
ovina heath schreberi, Ptilidium ciliare and Dicranella
heteromalla are occasional. Lichens may exceed
This is a heather-dominated community which is mosses in cover. Cladonia species are prominent
very poor in vascular associates, although some- with encrusting species such as Cladonia pyxida-
times showing a modest diversity among the ta, C. squamosa and C. fimbriata on bare ground.
bryophytes and, more especially, the lichens. Species like C. impexa, C. furcata and C. arbuscu-
Calluna vulgaris is usually the only woody species la are especially abundant on old Calluna togeth-
and the most abundant. The height and cover of er with Hypogymnia physodes.
the canopy are very variable depending on the age This community is confined to acid, base-poor
of the heather and the consequent phase of and oligotrophic sandy soils in the more
development, and also on grazing intensity. Erica continental lowlands of eastern England. The
cinerea, Ulex minor and U. gallii, important in dry profiles under the community are usually brown
heaths further south and west, are largely exclud- sands which are free to excessive-draining and
ed as is Erica tetralix. Ulex europaeus is uncom- have a low surface pH. In some localities, such as
mon, except where there has been disturbance. in Lincolnshire and around the Weald, the
Typically there are no grassy areas but Festuca impoverished soils are derived from arenaceous
ovina is very common throughout, usually as bedrock, but they have mostly developed from
scattered tussocks, often with less than 30% total sandy glacio-fluvial drift, sometimes supplement-
cover. Other grasses are few. Agrostis capillaris is ed by aeolian sand. The community occurs
occasional and there may be a little Deschampsia through the eastern lowlands of England, although
flexuosa. Associated dicotyledons are also few it is now very local.
and patchy. Occasionally Senecio jacobaea, This heath has been traditionally managed
Galium saxatile, Cerastium fontanum, Camp- with burning and grazing (both domestic livestock
anula rotundifolia and Luzula campestris may be and wild herbivores such as rabbits and deer).
present. Two species locally important in particu- However in many areas the abandonment of this
lar situations are Pteridium aquilinum and Carex traditional management has been followed by
arenaria. In bare areas Rumex acetosella together agricultural improvement or afforestation which
with ephemerals such as Aphanes arvensis, has reduced and fragmented tracts of this
Teesdalia nudicaulis, Myosotis ramosissima and community. In other areas the lack of grazing and
the annual Aira praecox may be found. burning has often permitted seral progression to
Only a few bryophyte species occur throughout scrub and woodland. The most common woody
the community. Hypnum cupressiforme and invaders are Betula pendula and Pinus spp., and
Dicranum scoparium are both constant. These two more occasionally, Quercus robur if mature trees
usually form the bulk of the bryophyte cover in are fairly close by.

76
 H1

Cover of Calluna vulgaris rather Lichens poorly represented.

open with some of Cladonia uncialis,
C. fimbriata, C. pyxidata, C. impexa,
C. squamosa, Cornicularia aculeata
and Hypogymnia physodes locally
abundant on mor and bare ground.

Hypnum cupressiforme very Bryophytes rather patchy but Teucrium scorodonia frequent Carex arenaria constant, Very impoverished rank
common among collapsed lichens often extensive with with Senecio jacobaea, Agrostis sometimes abundant. canopies of Calluna vulgaris.
Calluna vulgaris and in Cladonia furcata and capillaris and Galium saxatile
grassy patches. C. macilenta common. occasional.

H1a H1b H1c H1d H1e

Hypnum cupressiforme Hypogymnia physodes – Teucrium scorodonia Carex arenaria Species-poor sub-community
sub-community Cladonia impexa sub-community sub-community
sub-community
Cover of Calluna vulgaris Degenerate bushes are scarce Unbroken canopies of dense
is often less than complete and The heather cover is the same and heather cover is generally The heather is often tall and and often tall heather, up
only moderately tall. The most as in H1a but degenerate bushes vigorous and extensive. somewhat open with scattered to 50 cm or more high, are
prominent associates are predominate. The contribution Within more open areas plants or denser patches of characteristic here, among
Festuca ovina and the cryp- of Festuca ovina, Hypnum scattered plants of Teucrium Carex arenaria, sometimes which virtually no associates
togams which often exceed cupressiforme and Dicranum scorodonia or Senecio jacobaea co-dominant and growing on can survive, apart from
50% cover. Among the mosses scoparium is more uneven, can be found or small stretches sandy soil profiles. Festuca occasional sparse plants
Hypnum cupressiforme and the mosses occurring as small of turf with Festuca ovina, ovina occurs sparsely of Hypnum cupressiforme.
Dicranum scoparium are very patches among collapsed stems. Agrostis capillaris, Deschampsia sometimes with a little
This sub-community is
frequent. Lichens are also Lichens by contrast are abun- flexuosa and scattered Rumex Rumex acetosella, but other
typically of building and
abundant, Cladonia pyxidata, dant in these areas, encrusting acetosella, Galium saxatile and herbs are scarce. Among the
mature heather.
C. squamosa, C. fimbriata and species being joined or exceed- Cerastium fontanum. Lichens cryptogams only Hypnum
C. gracilis usually predominate, ed by Cladonia impexa, are very infrequent, but cupressiforme is frequent.
bulkier species like C. impexa, C. furcata and C. macilenta. Hypnum cupressiforme and
This sub-community is found
C. uncialis and C. arbuscula Hypogymnia physodes is most Dicranum scoparium remain
on coastal and inland dune
being less frequent and frequent here. common.
systems.
Hypogymnia physodes
occasional.

77
H2 Calluna vulgaris – saxatile may also be found in more open areas. In
some stands invading seedling and sapling trees
Ulex minor heath may occur, particularly Quercus spp., Betula spp.
and Pinus sylvestris.
This community is generally dominated by In the absence of burning, mosses and lichens
Calluna vulgaris, but with both Erica cinerea and can become common. Dicranum scoparium and
Ulex minor playing a very frequent and sometimes Hypnum jutlandicum are the most frequent
prominent role in the sub-shrub layer. The bryophytes, with peat-encrusting Cladonia species
constancy of the latter two species provides the and larger species such as Cladonia furcata and
most obvious floristic distinction between this C. arbuscula. Hypogymnia physodes can some-
community and the Calluna vulgaris – Festuca times be found on old heather stems.
ovina heath (H1). The canopy is very variable in This community is characteristic of impoverished
height, from 10 cm to 80 cm or more, and the acid soils, predominantly free-draining in south-east
structure depends greatly on the growth phase of and central southern England. It is characteristic of
the Calluna and whether the individuals are of free-draining profiles developed from pervious arena-
even or uneven age. Where burning occurs (for ceous or pebbly parent materials. Typically, these par-
example, in the New Forest) a characteristic ent materials have given rise to some kind of podzolic
patchwork of swales is formed. After fire E. cin- profile under this community which is highly acidic,
erea often increases in frequency because of its often with a superficial pH between 3.5 and 4.5, and
prolific seeding. Ulex minor normally plays a sub- generally impoverished. The community occurs from
sidiary role, forming a patchy understorey below the Poole Harbour area in the west through the New
the Calluna. No other sub-shrubs are found Forest, where stands are particularly numerous and
throughout the community. Ulex europaeus is extensive, to Surrey and the High Weald in the east,
occasional, but may be locally abundant after where it occurs as more local and fragmented tracts of
disturbance. Erica tetralix and Vaccinium myr- heathland.
tillus are found in particular sub-communities. The vegetation takes much of its structural, and
Other consistent associates are very few. some of its floristic, character from traditional
Deschampsia flexuosa is constant but patchy, grazing and burning treatments. However, when
though it may be more common in grazed areas. released from these treatments a progression to
On wetter soils it tends to be replaced by Molinia woodland can be expected. Agricultural improve-
caerulea. Pteridium aquilinum is occasional ments and forestry have caused fragmentation and
overall and preferential for one sub-community. isolation of small remnants of this community in
Scattered plants of Potentilla erecta or Galium many places.

78
 H2

Vaccinium myrtillus constant Pteridium aquilinum can be

and locally abundant, Pteridium quite common, but Vaccinium
aquilinum frequent and with myrtillus and young trees very
scattered tree seedlings and scarce.
saplings.

Molinia caerulea very common, Molinia caerulea occasional at

often almost totally displacing most with Deschampsia flexuosa
Deschampsia flexuosa. Erica very common. Dicranum sco-
tetralix frequent and sometimes parium and Hypnum ericeto-
exceeding E. cinerea. rum with Cladonia spp. patchi-
ly prominent in more open
areas.

H2b H2c H2a

Vaccinium myrtillus Molinia caerulea Typical sub-community

sub-community sub-community
Calluna vulgaris is generally
The sub-shrub canopy usually Calluna vulgaris is abundant, strongly dominant with sub-
consists of mixtures of heather usually with smaller amounts sidiary amounts of Ulex minor
with sometimes substantial of Ulex minor and particularly and Erica cinerea, the latter
amounts of Ulex minor, Erica of Erica cinerea which may be very variable and sometimes
cinerea and, strongly preferen- joined with or replaced by absent. Neither Vaccinium myr-
tial, Vaccinium myrtillus. Erica tetralix. More obviously tillus nor Erica tetralix occur
Pteridium aquilinum is more preferential is Molinia caerulea. and Molinia caerulea is scarce.
frequent, and in open areas Pteridium aquilinum is uncom- Deschampsia flexuosa is com-
there is usually some mon and young trees are rarely mon and sometimes with or
Deschampsia flexuosa and found. The ground layer is replaced by Festuca rubra.
occasionally a little Molinia also sparse. Pteridium aquilinum and tree
caerulea. Young trees are also seedlings are infrequent.
This sub community is
strongly preferential, with oak Bryophytes and lichens can
predominantly found on soils
and birch seedlings and be conspicuous among older
with impeded drainage.
saplings frequent, and small heather or after burning,
pines locally prominent. with Cladonia fimbriata,
Hypnum jutlandicum and C. coccifera, C. chlorophaea
Dicranum scoparium are occa- and C. arbuscula all slightly
sional but lichens are sparse. preferential.
This sub-community tends to be
found at the higher altitudes in
the range of H2.

79
H3 Ulex minor – Agrostis
curtisii heath
bryophytes Campylopus brevipilus is most distinc-
This community contains nearly all the sub-shrub tive and can be accompanied by C. paradoxus,
vegetation in which Ulex minor and Agrostis Polytrichum juniperinum, Dicranum scoparium,
curtisii occur together as important components, Hypnum jutlandicum and Leucobryum glaucum.
although A. curtisii can also occur occasionally in Cladonia impexa is one of the most common and
Calluna vulgaris – Ulex minor heath (H2). The conspicuous lichens with peat-encrusting species
canopy is usually fairly low, 20-30 cm high, and such as C. floerkeana, C. coccifera and C. pyxidata.
Calluna frequently dominates, especially when it Hypogymnia physodes often colonises old heather
has not been burned for some time. Compared with stems.
less oceanic heaths the most unusual feature of the This community is the characteristic sub-shrub
woody cover is the occurrence together of Erica community of impoverished acid soils which are
cinerea and E. tetralix, both of which are constants. protected against parching by a measure of
Both can grow vigorously, although E. cinerea is drainage impedance and a moderately oceanic
likely to be more prominent especially after burn- climate. It occupies a distinct position on soils
ing. Erica tetralix can have high cover locally, espe- that are too dry for the Erica tetralix – Sphagnum
cially on more strongly gleyed soils. Ulex minor compactum wet heath (M16) and too moist for the
maintains its high frequency throughout, although Calluna vulgaris – Ulex minor heath (H2). It is
its abundance is very variable. In stands which largely confined to south Dorset and Hampshire.
have not been burnt for some time A. curtisii and The combination of drainage impedance and
Molinia caerulea, the two characteristic and con- climate is the major influence on the floristics
stant grasses, are generally scattered, but after of this community, although grazing and burn-
burning A. curtisii and to a lesser degree Molinia ing still often exert an important measure of
can become prominent. In contrast to H2 control on its composition and structure. The
Deschampsia flexuosa is very scarce. Pteridium general effect of the combination of these treat-
aquilinum occurs occasionally and other herbs are ments is to curtail the mature and degenerate
found as scattered individuals. Potentilla erecta, phase of Calluna and to set back repeatedly any
Polygala serpyllifolia, Carex pilulifera and the par- invasion of trees and seral progression to wood-
asitic Cuscuta epithymum can all be found occa- land. The abandonment of traditional land use
sionally. In disturbed or burned situations Viola and soil improvement for agriculture in many
lactea is very characteristic. stands of this type of vegetation has meant that
Burning has the effect of opening the canopy, and surviving tracts can be fragmented, and are
mosses and lichens become prominent. Among the often sharply delineated from their surrounds.

80
 H3

Calluna vulgaris and Erica Agrostis curtisii very frequent,

tetralix reduced in frequency but not extensive, and Ulex
and cover and Molinia caerulea europaeus occasional at most
and Ulex minor somewhat among mixed or Calluna vul-
patchy. Agrostis curtisii very garis-dominated canopies.
abundant with Ulex europaeus
common and Viola lactea often
persistent.

H3c Polygala serpyllifolia common Species opposite very sparse

with a patchy cover of bryo- among usually dense sub-shrub
Agrostis curtisii sub-community phytes and lichens on more canopies.
open areas. Species present
Agrostis curtisii is very abun- include Campylopus brevipilus,
dant and Erica cinerea often co- Polytrichum juniperinum,
dominant, but Ulex minor and Cladonia impexa, C. floerk-
Molinia caerulea are patchy eana and C. coccifera.
and Calluna vulgaris and Erica H3a
tetralix much reduced in
frequency and cover. Ulex Typical sub-community
europaeus is strongly preferen-
tial in this sub-community The sub-shrubs typically form
where disturbance, often by an extensive canopy, often with
burning, is characteristic. The H3b Calluna vulgaris as the main
early stages of development dominant, although, sometimes
often allow colonisation by Cladonia spp. sub-community more mixed. Grasses are usual-
Viola lactea, which then per- ly subordinate. Ulex europaeus
sists as the dominants expand. The sub-shrub canopy is is occasional and Erica ciliaris
somewhat open, and although is found in south Dorset vegeta-
Calluna vulgaris is the leading tion. Other species are few,
species, dominance is often although Potentilla erecta is
shared between woody plants preferential at low frequency,
and Polygala serpyllifolia, a and there is sometimes a little
strong preferential here. There Pteridium aquilinum, Carex
is a noticeable cover of mosses pilulifera, Polygala serpyllifolia
and lichens, the species listed or Cuscuta epithymum.
above being most frequent. Bryophytes and lichens are
sparse.

81
H4 Ulex gallii – Agrostis heaths, Festuca ovina and Danthonia decumbens
are particularly important, with the sedges Carex
curtisii heath binervis and C. pilulifera also characteristic. On
cooler, moister slopes Scirpus cespitosus can be
This community is very similar to Ulex minor – prominent. The only dicotyledonous herb which
Agrostis curtisii heath (H3), with the replacement is a constant of this community is Potentilla erec-
of one gorse by another. The western limit of ta, which occurs as scattered individuals. Other
U. minor in east Dorset forms the boundary bet- occasional herbs are Polygala serpyllifolia and
ween these two heath types. Apart from this dif- Pedicularis sylvatica, with Viola lactea in dis-
ference they share five constants, namely, Calluna turbed situations. There are a variety of bryo-
vulgaris, Erica cinerea, E. tetralix, Molinia caer- phytes and lichens, but none occur with any
ulea and Agrostis curtisii, and these species, frequency.
together with U. gallii, generally account for the This community is confined to the warm
bulk of the vascular cover. Their proportions and oceanic parts of south-west Britain where it occurs
structure, however, vary considerably so that the on a variety of moist, acid soils. Like its eastern
appearance of stands differs markedly. The vegeta- counterpart, H3, this is a vegetation type of acid
tion can vary from a short mixed canopy of grass- soils that are too moist for dry heath but not so
es and sub-shrubs no more than 10 cm high (grass consistently waterlogged as to be able to sustain
heath), to a canopy of woody plants 50 cm or more wet heath. The community is confined to south-
high. There may also be quite extensive areas of west Britain, beyond a line from mid-Dorset to the
barer ground. Calluna and U. gallii are the most Quantocks, and including parts of the south Wales
common species and are often abundant. Calluna seaboard up to altitudes of 500 m.
often dominates. The frequent occurrence of Both climatic and edaphic conditions combine
E. cinerea and E. tetralix together distinguishes to influence the general character of this vegetation.
this vegetation from the corresponding dry heath However, in most situations burning and grazing
Calluna vulgaris – Ulex gallii heath (H8). Four have a marked effect on the floristics and physiog-
other sub-shrubs are more restricted. Vaccinium nomy of the vegetation and, with the exception of
myrtillus is commoner at higher altitudes with situations such as the Lizard where exposure to
increased rainfall. The others, Salix repens, Erica high and frequent winds is combined with a scarci-
ciliaris and E. vagans (a species restricted to ty of seed parents, these treatments are important
the Lizard in Cornwall) are found in the wetter for maintaining the community against succession
E. tetralix sub-community. to woodland. As with many lowland heath com-
Two grasses are constant, Agrostis curtisii and munities intensive improvement for agriculture
Molinia caerulea, which always make some and afforestation has reduced and fragmented its
contribution to the cover. Among the grassier extent.

82
 H4

Erica tetralix very infrequent in vegeta- Erica tetralix a frequent

tion usually dominated by Agrostis cur- component of the sub-shrub
tisii or E. cinerea. community.

Erica cinerea rather patchy but Erica cinerea and E. tetralix

Vaccinium myrtillus common in both very common, but associ-
a grassy heath with Festuca ates listed opposite occasional
ovina, Danthonia decumbens, at most in a less grassy heath.
Agrostis capillaris, Galium sax-
atile and occasional Carex
H4a pilulifera and C. binervis.

Scirpus cespitosus constant in Scirpus cespitosus absent from

Agrostis curtisii – Erica cinerea vegetation usually dominated vegetation variously dominated
sub-community by Calluna vulgaris, but with by one or more sub-shrubs and
H4b
other sub-shrubs and grasses grasses and with Erica vagans
Unusually there is an almost total locally abundant. or E. ciliaris locally abundant.
absence of Erica tetralix. Otherwise all Festuca ovina sub-community
the constants remain frequent except
for Agrostis curtisii which is often dom- Agrostis curtisii is often the most abundant H4d H4c
inant, forming a virtually pure and tus- species, most commonly with a fairly rich
socky sward. Other grasses do not gen- mixture of sub-shrubs and herbs forming a Scirpus cespitosus Erica tetralix sub-community
erally increase in cover. Other vascular grass heath. Both Ulex gallii and Calluna sub-community
and cryptogam associates are few in vulgaris have high frequencies; Erica Both grasses and sub-shrubs
number. cinerea is more patchy and Erica tetralix- Scirpus cespitosus is constant have very high frequency,
common but not constant. Vaccinium myr- and strongly preferential with including Erica tetralix,
This sub-community occurs throughout tillus is most common in this sub-commu-
the range of H4 and is often present as very frequent Calluna vulgaris, and each can be abundant.
nity, often as sparse shoots. The associates Molinia caerulea and Erica Potentilla erecta is frequent
regenerating vegetation after burning. listed above are more abundant and prefer- tetralix and common E. cinerea and Danthonia decumbens,
ential. Potentilla erecta is very common, and Vaccinium myrtillus. Ulex Polygala serpyllifolia and
often with Galium saxatile.Bryophytes and gallii and Agrostis curtisii, both Carex panicea occasional. This
lichens are very sparse in ungrazed stands constants, distinguish this sub- vegetation is best known on the
but occasional in grazed vegetation. community from wet heath. Lizard where Erica ciliaris or
This sub-community, along with the Dicranum scoparium and E. vagans occur in this type of
Scirpus sub-community, can be found at Leucobryum glaucum are fre- heath. Bryophytes and lichens
higher altitudes on Dartmoor and Exmoor quent and Cladonia impexa show a varying representation.
and also throughout the range on and C. uncialis occasional.
This sub-community is espe-
free-draining soils with grazing. This sub-community, along cially characteristic of lower
with the Festuca sub-communi- altitudes and is well represent-
ty, can be found at higher alti- ed on the Devon Pebble-Bed
tudes on Dartmoor and Exmoor. commons, the lower fringes of
Dartmoor and Bodmin Moor
and on the Lizard.

83
H5 Erica vagans – Schoenus There are a number of occasional associates. On
wetter ground Phragmites australis can be present
nigricans heath as conspicuous but scattered shoots.
Bryophytes vary considerably among stands
This heath is one of two sub-shrub communities but Campylium stellatum is constant and
in which the nationally-rare Erica vagans makes very frequent in runnels and may be abundant,
a constant and prominent contribution. Schoe- often with Riccardia multifida, R. sinuata and,
nus nigricans is also constant and usually abun- over gabbro, Scorpidium scorpioides. After wet
dant as strongly-developed tussocks. Molinia weather runnels often have swollen gelatinous
caerulea and Erica tetralix are also constant, globules of blue-green algae.
often with high cover, and together these four This community is confined to wet, base-rich
species dominate in mixtures. Between these but calcium-poor mineral soils and shallow
species there is a well-defined system of runnels peats on the Lizard in Cornwall. Here the distinc-
giving a distinct microhabitat. Among other sub- tive parent materials of serpentine and gabbro
shrubs only Ulex gallii occurs with any frequen- found in this area have given rise to soils that have
cy and may be co-dominant. Calluna is only a superficial pH of between 5.5 and 7.5 but in
occasional and E. cinerea scarce. Genista anglica, which magnesium predominates over calcium.
however, can occur frequently and is preferential The community makes the major proportion of the
to this community. open and enclosed heaths of the hinterland of
In undisturbed stands which have not been the peninsula.
burnt or grazed recently, Schoenus and E. vagans The floristics of this community are influenced
tend to be dominant and the vegetation is choked both by the mild oceanic climate and underlying
with their litter. Then, even common associates bedrocks of serpentine and gabbro, but the com-
like Potentilla erecta and Festuca ovina can be position and physiognomy of particular stands are
crowded out. After burning or grazing, or both, the affected by burning and sometimes also by graz-
associated flora is much richer. In the wetter run- ing. Other past treatments like the cutting of turf
nels sedges are often important with Carex puli- have also probably influenced the appearance and
caris constant, C. panicea and C. flacca frequent distribution of this community. There have
and, on gabbro, C. hostiana. Anagallis tenella, also been losses of this vegetation type to modern
constant, may form extensive mats. Among taller techniques of land improvement and much of the
herbs Serratula tinctoria and Succisa pratensis are remaining extent has statutory or voluntary pro-
constant and Sanguisorba officinalis frequent. tection.

84
 H5

Frequent Eleocharis multicaulis, Vegetation variable in composition

Eriophorum angustifolium, Drosera and structure, but species listed
rotundifolia, Pinguicula lusitanica, and opposite rare.
Dactylorhiza incarnata incarnata grow-
ing in runnels that are usually flooded
for much of the year. In ungrazed
stands, Phragmites australis may
be locally abundant.

H5b H5a

Eleocharis multicaulis sub-community Typical sub-community

Schoenus nigricans dominant with This vegetation has all the general
Molinia caerulea and Erica vagans. features of the community with no
Erica tetralix is somewhat less abun- additional preferential species. The
dant. Calluna vulgaris and Erica tussock/runnel structure is often well-
tetralix usually absent and Carex defined, but species-richness depends
panicea is also typically missing. greatly on treatment and especially
time since burning.

85
H6 Erica vagans – Ulex Thymus praecox, Lotus corniculatus, Galium
verum, Jasione montana, Danthonia decumbens
europaeus heath and Brachypodium sylvaticum. Immediately after
burning, diversity is increased, with ephemerals
This community is a distinctive type of sub-shrub including Aira caryophyllea and Centaurium
vegetation, but rather variable in floristics and erythraea. Continued burning and the dense shade
structure. The most obvious feature is a mixed and litter of older stands inhibit bryophytes and
canopy of sub-shrubs in which Erica vagans and lichens, which as a result are uncommon.
Ulex europaeus are the usual co-dominants. The This community is confined to the Lizard in
canopy is generally 30-60 cm high but in exposed Cornwall where it is characteristic of free-draining
situations may be not more than 10 cm high. brown earths that are usually quite base-rich but
Two other constant sub-shrubs, Ulex gallii and calcium-poor and fairly oligotrophic. It is found
E. cinerea, can also be abundant although the for- on soils similar to that of H5 with a pH of general-
mer may be suppressed in dense stands. Calluna ly between 5 and 7, but which are more free-drain-
vulgaris is not frequent and has generally ing. Therefore it is typically found on the steeper,
low cover. shedding slopes around coves and on the cliff tops
In contrast to the Erica vagans – Schoenus of the headlands. Although it is mainly coastal in
nigricans heath (H5) community, E. tetralix is only distribution it is not strictly speaking a maritime
occasional and confined to wetter soils (see heath and is replaced on slopes which are
sub-community H6d) with several preferential exposed to salt spray by Calluna vulgaris –
associates. The only herbaceous associates Scilla verna heath (H7).
common throughout are Carex flacca, Potentilla Edaphic variation and local differences in the
erecta and Polygala vulgaris. The most common warm oceanic climate strongly influence floristic
and distinctive herbs of this community are Viola diversity, but treatments, especially burning, and
riviniana, Filipendula vulgaris, Stachys betonica, to a lesser extent grazing, also have a marked effect
Hypochoeris radicata, Agrostis canina ssp. on composition and physiognomy of the vegeta-
montana, Dactylis glomerata and Scilla verna. tion. However, the progression to scrub and wood-
Most of these species are found in recently burned land in the absence of these treatments would
stands but become more scattered and reduced in probably be slow due to the lack of seed parents
number as the vegetation and litter increase. and the poor quality of the soil. Preferential culti-
On shallower soils, especially when grazed, a vation of the more fertile soils developed over gab-
rich short herb layer is maintained with several bro and schists means that the community sur-
additional species including Festuca ovina, vives most extensively over serpentine.

86
 H6

Molinia caerulea and Potentilla erecta Filipendula vulgaris very frequent but
constant with Serratula tinctoria fre- Molinia caerulea absent and Potentilla
quent, but Filipendula vulgaris rather erecta and Serratula tinctoria only
uncommon. occasional.

Agrostis curtisii constant and Agrostis curtisii and associates Sub-shrub canopy usually low Sub-shrub canopy usually
often abundant, particularly listed opposite absent, but Erica and open with a rich flora extensive with occasional
after burning, with Calluna vul- tetralix constant in small between the bushes including Rubus fruticosus agg., Prunus
garis scarce. Carex panicea, amounts and Sanguisorba Danthonia decumbens, Koeleria spinosa and Pteridium aquil-
Hypericum pulchrum, Viola officinalis and Schoenus macrantha, Aira caryophyllea, inum, and scattered Teucrium
lactea and Polygala serpyllifolia nigricans frequent. Galium verum and Leontodon scorodonia and Geranium san-
frequent. taraxacoides as constants. guineum, but associates listed
opposite very sparse in more
recently regenerating stands.

H6c H6d H6b H6a

Agrostis curtisii Molinia caerulea Festuca ovina sub-community Typical sub-community

sub-community sub-community
Here the abundance and height Typical heath with a complete
Apart from occasional occur- Erica vagans and Ulex of the sub-shrub canopy is less range of floristic and structural
rences in H6a, Agrostis curtisii europaeus retain high frequen- than in the typical form with a vegetation related to burning.
is largely confined to this sub- cy and abundance with smaller total cover of often less than A few years into the cycle there
community where it can be amounts of Ulex gallii. Both 50%. Apart from Potentilla is a well-developed sub-shrub
very abundant especially after Erica cinerea and Calluna vul- erecta the common herbs are all canopy with most of the com-
burning. Molinia caerulea is garis are reduced in frequency well represented in this sub- munity constants. Richer stands
also frequent and the two grass- but Erica tetralix is constant in community. In the more open have occasional Teucrium
es sometimes dominate under small amounts. In stands not conditions there are more cryp- scorodonia and Geranium san-
an open canopy of Erica recently burned Molinia togams than usual, Hypnum guineum, the only preferential
vagans, E. cinerea, Ulex gallii caerulea is distinctive. Its litter cupressiforme s.l. is frequent dicotyledons here. With
and U. europaeus. Among the depresses herbs and generally and various Cladonia spp. increasing age the constant
herbs Viola riviniana is absent only Carex flacca, Viola rivini- common. herbs begin to thin out leaving
and Filipendula vulgaris scarce. ana, Potentilla erecta and a dense woody cover.
Danthonia decumbens and Stachys betonica occur with
Potentilla erecta are very fre- any frequency, with small
quent and Stachys betonica and amounts of the preferentials
Serratula tinctoria are common, listed above.
as are the preferential species
listed above.

87
H7 Calluna vulgaris –
corniculatus, Thymus praecox and Hypochoeris
Scilla verna heath radicata, the latter of which tends to favour drier
soils. Anthyllis vulneraria also favours drier soils
In this heath, sub-shrubs are a consistent feature, as do Euphrasia species. Other species are more
though they are not always obvious. The canopy is characteristic of particular sub-communities.
typically very short, rarely over 20 cm, and some- In contrast cryptogams are few and never
times when grazed forming a mat only 2-3 cm show high cover. Among the mosses only Hypnum
high. The cover of woody plants is rarely continu- cupressiforme s.l. is moderately frequent and
ous. Even where sub-shrubs are more extensive, Frullania tamarisci, the commonest hepatic, is
they are commonly penetrated by herbs. Calluna infrequent. Several Cladonia species are occasional.
vulgaris is the most frequent sub-shrub and the This community occurs over a wide variety of
commonest dominant, though on dry soils it is moderately base-poor soils on the less exposed
accompanied by Erica cinerea. On wetter soils the parts of maritime cliffs all around the coast of
latter is much reduced and E. tetralix and/or Britain except to the east and south between
Empetrum nigrum ssp. nigrum are the usual asso- Durham and Dorset. The single most distinctive
ciates. No other woody species occurs frequently difference between the habitat of this kind of
throughout, although Ulex gallii is occasional. heath and the habitats of other sub-shrub commu-
Among herbaceous associates grasses are often nities is the input of salt spray generated by break-
important. Festuca ovina is the most frequent ing waves and carried inland by the wind.
grass species, though F. rubra is also common. The floristic and structural variation in this
Also common and a constant is Holcus lanatus, community is influenced by the climatic and
often with Dactylis glomerata on drier soils or edaphic differences both throughout the consider-
Danthonia decumbens on moister ground. In able geographic range of the community and over
wetter, northern heaths Agrostis capillaris and particular stretches of cliff. Grazing also affects the
Anthoxanthum odoratum can become very composition and appearance of the vegetation and
common, but Molinia caerulea is infrequent. probably contributes to maintaining it against suc-
There are a variety of other herbs. Most distinctive cessional change. However, over much of its range
among the constants are Plantago maritima and this vegetation can be considered a climatic cli-
Scilla verna. Other common and constant species max as exposure to even small amounts of salt
are Plantago lanceolata, Potentilla erecta, Lotus spray hinders the invasion of woody invaders.

88
 H7

Erica cinerea and Hypochoeris radicata reduced in Agrostis capillaris and Anthoxanthum
frequency but Erica tetralix and Empetrum nigrum odoratum occasional but Erica cinerea
common. Plantago maritima and P. lanceolata often and Hypochoeris radicata remain very
very conspicuous. Anthoxanthum odoratum and frequent and E. tetralix, Empetrum
Agrostis capillaris frequent and Carex panicea and nigrum, Carex panicea and C. nigra
Carex nigra occasional. are rare.

Erica tetralix constant with Empetrum nigrum constant, Armeria maritima and Sedum Dactylis glomerata and
occasional Empetrum nigrum with Erica tetralix, Danthonia anglicum constant and often Anthyllis vulneraria occasional
and frequent Danthonia decum- decumbens and Succisa praten- abundant with frequent Dactylis but other listed associates rare.
bens and Succisa pratensis. sis only very occasional but glomerata, Anthyllis vulneraria
Molinia caerulea, Nardus Trifolium repens and Luzula and Jasione montana, and occa-
stricta and Salix repens multiflora quite common. sional Plantago coronopus and
locally prominent. Silene vulgaris maritima.

H7c H7d H7a Viola riviniana, Polygala Viola riviniana occasional

vulgaris, Carex flacca and but other listed associates rare
Erica tetralix sub-community Empetrum nigrum ssp. nigrum Armeria maritima C. caryophyllea frequent. among an often impoverished
sub-community sub-community Achillea millefolium, Leontodon Calluna vulgaris-dominated
Two features distinguish this taraxacoides, Galium verum, cover.
type of heath. First, among This type of heath shares Here, where salt-spray deposi- Stachys betonica and Serratula
the sub-shrubs Erica cinerea is several features with H7c; Erica tion is high, the canopy of sub- tinctoria occasional and Erica H7e
much reduced and replaced by cinerea is seldom found but is shrubs is generally less exten- vagans and Ulex europaeus
Erica tetralix as the usual com- replaced here by Empetrum sive, either with a mosaic of locally prominent. Calluna vulgaris
panion to Calluna vulgaris. The nigrum which is often co-domi- open areas or reduced to dis- sub-community
canopy is typically extensive, nant with Calluna vulgaris. crete patches of bushes. The H7b
but short because of grazing. Erica tetralix and Ulex species only common sub-shrubs are In general floristics this heath
Secondly, although grasses are are rare. Festuca ovina, F. rubra, Calluna vulgaris and Erica Viola riviniana sub-community type resembles impoverished
prominent Festuca ovina is Agrostis capillaris and cinerea, although the latter is versions of the Armeria and
often replaced by Festuca rubra Anthoxanthum odoratum are reduced in exposed situations. The sub-shrub canopy is more Viola sub-communities though
as the most abundant species frequent and Carex panicea Growing among and between extensive than in H7a with both usually with a taller canopy.
and Agrostis capillaris, common. Among dicotyledons the bushes is Festuca ovina as Calluna vulgaris and Erica Calluna vulgarisis the usual
Anthoxanthum odoratum and Plantago maritima and P. lance- the dominant, sometimes with cinerea very frequent and often dominant though Erica cinerea
Danthonia decumbens are very olata are very frequent and F. rubra, frequently with Holcus co-dominant. Erica tetralix and is common; all other sub-shrubs
common and sometimes co- Thymus praecox and Lotus lanatus and Dactylis glomerata. Empetrum nigrum are typically are scarce. Festuca ovina is
dominant. Plantago maritima is corniculatus common, while Apart from Potentilla erecta, very scarce. The herbaceous the most frequent grass
often abundant with P. lanceo- Hypochoeris radicata is very which is scarce, all other com- plants are most distinctive. All with F. rubra less common.
lata common. rare on the moist soils. munity constants and the asso- the community constants are Hypochoeris radicata is quite
ciates listed above are well rep- well represented together with frequent but rosette herbs are
This sub-community is most This sub-community is found the species listed above. poorly represented in the rank
resented.
prominent on the north-west predominantly in northern herbage. Thymus praecox and
coast of Britain, particularly in Britain. This sub-community, along This sub-community, along
Lotus corniculatus, although
the Hebrides and Sutherland, with the Viola sub-community, with the Armeria sub-communi-
common are often not
with scattered occurrences occurs throughout the range of ty, occurs throughout the range
abundant.
down to Anglesey. H7 but is better developed to of H7 but is better developed
the south of Galloway with only south of Galloway with only This sub-community is found
local stations beyond this. local stations north of this. throughout the range of H7 but
is rarer to the north, although it
is well-represented in Shetland.

89
H8 Calluna vulgaris – Rhytidiadelphus squarrosus and Pleurozium
schreberi are more occasional. In more open
Ulex gallii heath situations, or on burned or disturbed bare ground,
mosses such as Campylopus paradoxus, Poly-
Floristically this is a diverse community with only trichum piliferum or P. juniperinum can become
three constants overall, namely Calluna vulgaris, abundant along with lichens species such as
Erica cinerea and Ulex gallii. Erica tetralix, Cladonia impexa and C. squamosa.
Molinia caerulea and Agrostis curtisii are typical- This community is found on free-draining,
ly lacking from this community. Often the three generally acid to circumneutral soils, in the warm
constant sub-shrubs are co-dominant, but propor- oceanic regions of lowland Britain. It can be found
tions are variable and where E. cinerea is reduced over a wide range of arenaceous sedimentaries and
Vaccinium myrtillus can appear. On disturbed acid igneous and metamorphic rocks as well as on
ground U. europaeus may be abundant and both silty and sandy superficials like loess and aeolian
Pteridium aquilinum and Rubus fruticosus agg. sands. The superficial pH underneath this com-
may appear in the heath. munity is usually from 3.5 to 4.5. It occurs
Typically sub-shrub cover is high and herbs are throughout south-western England and Wales, on
sparse, but often the bushes are separated by the Isle of Man and, more sporadically, in the
grassy runnels, a feature accentuated by grazing. southern Pennine fringes and the East Anglian
The most frequent grasses are Agrostis capillaris coast.
and Festuca ovina with A. canina ssp. montana, Local climatic and edaphic conditions influence
F. rubra, Anthoxanthum odoratum and Danthonia floristic variation; grazing by rabbits, sheep or
decumbens occasional to frequent. Deschampsia cattle, and sometimes burning (which is normally
flexuosa and Nardus stricta are much more patchy an accidental occurrence), affect physiognomy and
in their occurrence. There is often some Potentilla composition. The community is maintained against
erecta and Galium saxatile, and much more succession to woodland in most situations by graz-
occasionally Teucrium scorodonia and Polygala ing and burning, although in some situations expo-
serpyllifolia. Additional herbs are characteristic of sure to the wind prevents the establishment of
particular sub-communities. woody invaders such as Betula spp. and Quercus
In general bryophytes and lichens are not spp. Much former heath has been improved for
numerous or diverse. There may be some Hypnum agriculture and it now often survives as patches on
cupressiforme and Dicranum scoparium, and marginal grazing land.

90
 H8

Danthonia decumbens, Anthoxanthum Potentilla erecta can be quite common,

odoratum, Festuca rubra and Potentilla but other species listed opposite are
erecta frequent. scarce.

Sanguisorba minor, Carex flac- Combinations of species listed Scilla verna, Plantago maritima, Species listed opposite usually
ca, Helianthemum nummulari- opposite not present. Hypochoeris radicata and very scarce.
um and Plantago lanceolata fre- Thymus praecox frequent.
quent, with occasional Galium Erica cinerea reduced in fre- Erica cinerea remains common
verum, Lotus corniculatus, quency but Vaccinium myrtillus but Vaccinium myrtillus and
Carex caryophyllea, Linum constant in small amounts with other species listed opposite
catharticum, Brachypodium syl- Deschampsia flexuosa frequent scarce.
vaticum, Hypericum pulchrum and Nardus stricta, Pleurozium
and Stachys betonica. schreberi and Rhytidiadelphus
squarrosus occasional.

H8c H8b H8d H8e H8a

Sanguisorba minor Danthonia decumbens Scilla verna sub-community Vaccinium myrtillus Species-poor sub-community
sub-community sub-community sub-community
The three constant sub-shrubs There is an extensive and dense
This is the most striking of the The canopy is well-developed are typically extensive as co- Erica cinerea is present with sub-shrub canopy. The canopy
sub-communities, developing here with Ulex gallii and Erica dominants. There is a rather reduced frequency, though it can be quite tall, thicker stands
on more calcareous soils. The cinerea especially abundant, species-poor herbaceous ele- can remain locally abundant being less penetrable to live-
sub-shrubs and structural varia- but not as dense as in H8a and ment with Festuca ovina the in this upland sub-community. stock. The associated flora is
tion are similar to those of H8b. typically there is a system of most frequent, and often the The most prominent sub-shrub very sparse with grasses such
With them, however, is a group grassy runnels. In these Agrostis only, grass. Potentilla erecta is usually Ulex gallii with as Agrostis capillaris, Festuca
of strongly preferential herbs, capillaris and Festuca ovina are remains common while smaller amounts of Calluna ovina and F. rubra reduced to
listed above. Where numbers quite frequent but more charac- Hypochoeris radicata is more vulgaris and, strongly diagnostic, small tufts with occasional
of these are present in grazed teristic is Danthonia decumbens distinctive and preferential Vaccinium myrtillus. Grasses and weak Potentilla erecta
stands the vegetation can look and less commonly Anthox- with Scilla verna. Plantago can have a high cover, with and Galium saxatile. Where
like Mesobromion grassland anthum odoratum, Festuca maritima can occur frequently Agrostis capillaris, Festuca the canopy is opened by
growing among islands of the rubra and Agrostis canina mon- and may be abundant in a ovina and Anthoxanthum disturbance or burning Ulex
heath sub-shrubs. tana. Mixtures of these species grazed sward. odoratum all frequent, but europaeus or Pteridium aquil-
typically form the bulk of a Deschampsia flexuosa is prefer- inum may expand. Cryptogams
This sub-community is found in This sub-community is con-
rough cover between the bush- ential. Nardus stricta can also are generally infrequent with
areas with calcareous bedrock fined to the coastal fringe in
es. Carex pilulifera is also typi- occur with local abundance but a few patches of Hypnum
mantled with drift. western Britain.
cal but patchy, though particu- dicotyledonous associates are cupressiforme.
larly abundant in open places few. Galium saxatile and
This sub-community occurs
after burning, and Viola lactea Digitalis purpurea are occasion-
throughout the range of H8.
can also become prominent in al. In more open situations
such places. bryophytes can form a lush
but patchy cover.
This sub-community occurs
throughout the range of H8 but This sub-community is found
is better represented in the west. mainly in the upland fringes.

91
H9 Calluna vulgaris – dontium lineare may be frequent. On exposed soil
there can be locally abundant Polytrichum juniper-
Deschampsia flexuosa heath inum, P. piliferum and P. commune. Among leafy
hepatics Gymnocolea inflata is particularly charac-
Calluna vulgaris is almost always the most teristic. The commonest lichens are Cladonia
abundant plant in this community, often forming a chlorophaea, C. floerkeana, C. squamosa, C. coni-
fairly low and open canopy. Where burning is ocraea and C. fimbriata.
frequent, the individuals are immature and stands This heath is the characteristic sub-shrub
are uniform in age. No other sub-shrubs are vegetation of acid and impoverished soils at low
consistently frequent throughout, although some to moderate altitudes through the Midlands and
can be quite common and locally abundant. northern England. It is normally found on very
Vaccinium myrtillus is the most important, base-poor soils with a surface acidity generally of
particularly at higher altitudes. More locally pH 3-4, highly oligotrophic and at least moderate-
V. vitis-idaea and Empetrum nigrum ssp. nigrum ly free-draining, often excessively so, which have
can be found. Erica cinerea, E. tetralix and Ulex been derived from a wide variety of parent materi-
gallii by contrast are very scarce. als. It is found mainly in the southern Pennines
The only other vascular constant is and North York Moors with more local occur-
Deschampsia flexuosa, although even in open rences scattered through the Midland plain.
heather it often occurs only as sparse tufts, and The cool and wet climate has some influence
under dense canopies it can almost disappear. on the floristics of this community, but much of its
Molinia caerulea can become frequent on moister character derives from a combination of frequent
ground, but Agrostis capillaris, Holcus lanatus, burning and grazing. Also the heavy atmospheric
H. mollis and Festuca rubra only occur occasion- pollution in the areas in which this heath occurs
ally. Other herbs are also few and are of low cover. is thought to inhibit bryophyte and lichen diversi-
Galium saxatile and Potentilla erecta are frequent ty of the community. The community has been
in grazed stands and Juncus squarrosus and reduced considerably in extent. In the lowlands
Pteridium aquilinum occasionally occur. Seed- large tracts of heath have been reclaimed for agri-
lings of Quercus spp., Betula spp. and Pinus culture whilst other areas have been lost to inva-
sylvestris may be seen but rarely survive to the sion by trees after the neglect of traditional treat-
sapling stage due to frequent burning and grazing. ments. Furthermore, both in the lowlands and
The bryophyte and lichen flora is characteristic, around the upland fringes, the community has
although poor in species. Hypnum cupressiforme been replaced with coniferous plantations, or
s.l. is restricted, but Pohlia nutans is constant and land use changes have led to the spread of
very common with occasional Campylopus U20 Pteridium aquilinum – Galium saxatile
paradoxus and Dicranum scoparium. Ortho- community.

92
 H9

Vaccinium myrtillus or, more Vaccinium myrtillus and Campylopus

locally, V. vitis-idaea, V. inter- paradoxus at most occasional, and
medium or Empetrum nigrum other species listed opposite rare.
common or locally abundant
with occasional to frequent
Campylopus paradoxus,
Molinia caerulea constant at Molinia caerulea absent.
Gymnocolea inflata,
low cover.
Barbilophozia floerkii,
Deschampsia flexuosa often Deschampsia flexuosa may be
Cladonia chlorophaea,
especially abundant with occa- abundant but not with associ-
C. floerkeana and C. squamosa.
sional Holcus mollis and ates listed opposite.
Festuca rubra; Galium saxatile
and Potentilla erecta frequent Hypnum cupressiforme and Calluna vulgaris and Deschampsia
with occasional Rumex ace- Dicranum scoparium common flexuosa often the only plants,
tosella. and sometimes abundant. with occasional Pohlia nutans.

H9b H9e H9d H9a H9c

Vaccinium myrtillus – Molinia caerulea Galium saxatile sub-community Hypnum cupressiforme Species-poor sub-community
Cladonia spp. sub-community sub-community sub-community
Calluna vulgaris remains In this, the most impoverished
This is the richest sub-commu- Calluna is generally very abun- constant but is often rivalled Calluna is typically strongly form, Calluna vulgaris and
nity characterised by younger dant, but Deschampsia flexuosa in cover by Deschampsia dominant but exceptionally the Deschampsia flexuosa are the
canopies of heather, often is frequently accompanied flexuosa,and stands are locally bushes tend to be large and only constants, and in frequent-
recovering from burning. There by small amounts of Molinia enriched by a little Holcus mol- mature or even degenerate. ly burned heather even the
are frequently one or more of caerulea. The ground layer, lis or Festuca rubra. Commonly Vaccinium myrtillus and latter can almost disappear.
the sub-shrubs listed above. however, is poorly developed there are scattered plants or Pteridium aquilinum are both Vaccinium myrtillus is
Often Deschampsia flexuosa with just very sparse Pohlia prominent patches of Galium occasional and there are fre- occasional. Pohlia nutans,
has a rather low cover. Among nutans and Campylopus para- saxatile and scattered Potentilla quent, even dense tufts of Campylopus paradoxus and
the sub-shrubs, bryophytes are doxus. erecta with Rumex acetosella Deschampsia flexuosa. The Orthodontium lineare all show
more varied than in any other on bare areas. Lichens and mosses are most distinctive; reduced frequencies compared
This sub-community is mainly
type of this heath. Pohlia hepatics are sparse and among Hypnum cupressiforme s.l. is with H9b.
found on wetter soils, and along
nutans, Campylopus paradoxus the mosses only Pohlia nutans unusually common and abun-
with the Galium and Hypnum This and the Vaccinium sub-
and Orthodontium lineare all and Hypnum cupressiforme dant with Dicranum scoparium
sub-communities, is primarily community are the usual forms
occur frequently and the leafy s.l. occur more than very also preferential and frequently
found on lowland sites where in the southern Pennines and
hepatics and lichens listed occasionally. rivalling Pohlia nutans in its
burning is no longer practised. the North York Moors, and are
above are occasional to fre- cover. Apart from occasional
This sub-community is mainly widespread and sometimes
quent. Hypogymnia physodes growing
found on wetter soils, and along extensive over heathlands that
on older Calluna vulgaris,
This and the species-poor sub- with the Molinia and Hypnum are still frequently burned.
lichens are very few.
community are the usual forms sub-communities, is primarily
in the southern Pennines and found on lowland sites where This sub-community is mainly
the North York Moors and are burning is no longer practised. found on wetter soils, and along
widespread and sometimes with the Galium and Molinia
extensive over heathlands that sub-communities, is primarily
are still frequently burned. found on lowland sites where
burning is no longer practised.

93
H10 Calluna vulgaris – Erica species can develop. In exposed stands there is
often a patchy carpet of Racomitrium lanuginosum
cinerea heath and fruticose lichens. However, more important
than these species in the community as a whole are
This community is typically dominated by bulky pleurocarpous mosses such as Hypnum
Calluna vulgaris, but the cover, height and cupressiforme s.l., Pleurozium schreberi and
structure of the sub-shrub canopy vary markedly Hylocomium splendens, with Rhytidiadelphus tri-
depending on the intensity and timing of burning quetrus and R. loreus also occurring occasionally.
and grazing. Erica cinerea, a constant, is frequent These species, with Dicranum scoparium, become
but generally subordinate to heather and persists abundant with the maturing and opening up of the
below taller Calluna canopies. Vaccinium Calluna bushes.
myrtillus, by contrast, is at most occasional and This heath is characteristic of acid to circum-
V. vitis-idaea is scarce. Empetrum nigrum ssp. neutral and generally free-draining soils in the
nigrum can occur, but mainly in sub-community cool oceanic lowlands and upland fringes of
H10b. The restricted occurrence of these northern and western Britain. The soils on which
sub-shrubs is a contrast with Calluna vulgaris – this community is found can be quite moist as a
Vaccinium myrtillus heath (H12). result of the climate and the superficial pH
Apart from the abundance of the two constant beneath the community can be anywhere between
sub-shrubs there are two other distinctive floristic 3.5 and 6. It occurs widely through the more
features of this type of heath. These are firstly the oceanic parts of Scotland, with outlying stands in
high frequency of grasses and to a lesser extent Wales, western England and around the east-
sedges and dicotyledons, and secondly the central Highlands.
striking contribution that the ground layer makes In more exposed situations it may be considered
to this community. Deschampsia flexuosa is the as an edaphic or climatic climax, but often
most consistent grass throughout, with Agrostis burning and grazing are important in controlling its
canina and Nardus stricta occasional to frequent. composition and structure. Steady grazing pressure
In certain sub-communities Festuca ovina, pushes the vegetation towards the Festuca ovina –
Anthoxanthum odoratum, Agrostis capillaris and Agrostis capillaris – Galium saxatile grassland (U4)
Molinia caerulea become very common. Carex or, over more base-rich soils, the Festuca ovina –
binervis and C. pilulifera are very characteristic of Agrostis capillaris – Thymus praecox grassland
this community. After burning, mixtures of these (CG10). After fire, heavy grazing can precipitate a
plants can become patchily abundant and run-down of the heath to swards in which Nardus
Deschampsia flexuosa and C. pilulifera temporar- stricta or Juncus squarrosus play an important part
ily dominant. There are typically only a few or permit the spread of Pteridium aquilinum.
dicotyledons, but Potentilla erecta is a Release from grazing and burning, in all but the
constant and Galium saxatile is fairly common. most exposed sites, would theoretically permit pro-
After burning, a local abundance of Polytrichum gression to scrub and woodland, although in many
piliferum, P. juniperinum and encrusting Cladonia areas natural seed parents are now scarce.

94
 H10

Sub-shrub canopy often short in a grassy Combinations of such species rare.

heath with frequent Festuca ovina, F. rubra,
Agrostis capillaris, Anthoxanthum odora-
tum and Galium saxatile and occasional
Campanula rotundifolia, Succisa pratensis
and Hypericum pulchrum. Dicranum sco-
parium, Pleurozium schreberi and Empetrum nigrum ssp. nigrum Empetrum nigrum ssp. nigrum
Hylocomium splendens patchy. quite common and Scirpus and Scirpus cespitosus occa-
cespitosus patchily prominent sional at most and Molinia
with frequent Carex panicea caerulea very common at low
and C. pilulifera and occasional covers with frequent Carex
Danthonia decumbens very Combinations of such species rare. Huperzia selago. Racomitrium binervis and occasional Juncus
common with occasional to lanuginosum common and squarrosus. Campylopus para-
frequent Carex pulicaris, Viola often abundant among degener- doxus, Sphagnum capillifolium
riviniana, Linum catharticum, ating bushes with patches of and Diplophyllum albicans
Prunella vulgaris and Primula Cladonia uncialis and occasional to frequent.
vulgaris. C. impexa.

H10d H10c H10b H10a

Thymus praecox – Carex pulicaris Festuca ovina – Anthoxanthum Racomitrium lanuginosum Typical sub-community
sub-community odoratum sub-community sub-community
In this, the most species-poor sub-com-
This heath is found on relatively base- Calluna vulgaris is still abundant but This is found on exposed sites where munity, Calluna is typically dominant
rich brown earth soils and is very sim- Erica cinerea may often be co-domi- the sub-shrub canopy is more open and abundant in pioneer and building
ilar to H10c with Calluna vulgaris and nant. The sub-shrubs are usually short, and Calluna vulgaris is the usual regrowth after burning. Erica cinerea is
Erica cinerea both able to show promi- commonly forming a mosaic with a dominant. Erica cinerea is frequent, very frequent and can be prominent.
nence and with herbs and bryophytes grassy turf. Most frequent here are the Vaccinium myrtillus occasional Vaccinium myrtillus is occasional and
both being of structural importance. grasses and other species listed above. and Empetrum nigrum spp. nigrum Empetrum nigrum nigrum and Erica
Here there are additional preferentials, Carex binervis and C. pilulifera are is preferential and quite common. tetralix scarce. Monocotyledons are
making this the most species-rich sub- also common. Dicotyledonous herbs Grasses are sparse with scattered tufts few with Deschampsia flexuosa very
community. The species listed above are more numerous than in 10a and of Deschampsia flexuosa other grasses frequent and sometimes prominent.
are most frequent together with Carex 10b. Potentilla erecta and Galium very occasional. Carex binervis is rare, Molinia caerulea is preferential and
panicea and Thymus praecox. Among saxatile are both very common with its place being taken by Carex pilulif- patchily abundant with occasional
the bryophytes Dicranum scoparium occasional records for several species . era and C. panicea. Scirpus cespitosus Scirpus cespitosus and Juncus squarro-
and the pleurocarps remain very com- Bulky pleurocarpous mosses are con- is also frequent. Potentilla erecta sus. Carex binervis is well represented.
mon; additionally Rhytidiadelphus sistent and distinctive here with fre- is the only frequent dicotyledon Apart from Potentilla erecta and
triquetrus and Breutelia chrysocoma quent Hypnum cupressiforme s.l., Huperzia selago is preferentially com- Galium saxatile, dicotyledons are
are frequent. Pleurozium schreberi, Hylocomium mon. There are substantial areas of the very sparse. The ground layer is also
splendens and also Dicranum ground layer with Racomitrium lanugi- poor in species and of low cover.
This sub-community is local in
scoparium. nosum the most abundant moss, and
occurrence but can be found on Skye, This sub-community is found through-
frequent Hypnum cupressiforme s.l.
Rum and Uist and scattered localities This sub-community is common in out the range of H10.
Lichens are well represented.
through the Highlands and Southern south-west Scotland.
Uplands. This sub-community is typical of the
Western Isles and Shetland.

95
H11 Calluna vulgaris – There may be hypnoid mosses such as Hypnum
cupressiforme s.l., Pleurozium schreberi, Hylo-
Carex arenaria heath comium splendens and Rhytidiadelphus tri-
quetrus in the turf. On areas of bare ground acro-
Calluna vulgaris is the only constant sub-shrub carps such as Polytrichum juniperinum, P. pilifer-
found in this community and is often abundant, um and Ceratodon purpureus may be patchily
although cover may be discontinuous and patchy abundant.
in younger or grazed stands. Other frequent This is the characteristic sub-shrub vegetation
sub-shrubs are Erica cinerea and Empetrum of stabilised, base-poor sands on dunes and plains
nigrum ssp. nigrum, and each can be locally around the coasts of Britain. The heath is largely
abundant, to the exclusion of Calluna itself. confined to sands with a pH of less than 5 and can
Sometimes Rosa pimpinellifolia is plentiful, and only establish on sediments with surface stability
together with Erica tetralix and Salix repens is such as found on older dunes and on consolidated
found in transitions to wetter heath. sand plains. It is very local along the coasts of
Carex arenaria is constant, but no more than western England and Wales, becoming commoner
moderately abundant and often senile, except in Scotland.
where the sand is locally mobile. Ammophila The community develops in primary succession
arenaria is also frequent throughout, though by colonising fixed dune grasslands on acid sands
usually sparse. In more species-poor stands these or where more lime-rich sands have become
may be the only species, but often there is some leached. Relief from grazing is probably important
Festuca rubra (or F. ovina) with Agrostis capillaris for its establishment but once established predation
and Anthoxanthum odoratum and less commonly by herbivores, along with variation in regional cli-
Poa pratensis. Variation among dicotyledons is mate and substrate, influences its composition and
modest, but Galium verum, Lotus corniculatus, structure, and ultimately, grazing maintains the
Viola riviniana and Thymus praecox all occur community against reversion to grassland or pro-
quite frequently with several other herbs. gression to scrub and woodland.

96
 H11

Erica cinerea constant and Aira Erica cinerea and Aira praecox only The associates listed opposite are
praecox frequent but Empetrum occasional at most, but Empetrum all scarce in impoverished mixtures
nigrum nigrum, Agrostis capillaris nigrum nigrum, Agrostis capillaris of Calluna vulgaris and Carex
and Galium saxatile are rare. and Galium saxatile very frequent. arenaria.
Dicranum scoparium common Cladonia arbuscula and C. impexa
and some of Cladonia furcata, can be patchily prominent but
C. floerkiana, C. pyxidata, stretches of lichen-rich turf are
C. gracilis and C. foliacea in an not characteristic.
often extensive carpet of lichens.

H11a H11b H11c

Erica cinerea sub-community Empetrum nigrum ssp. nigrum Species-poor sub-community

sub-community
This sub-community is prominent Calluna vulgaris is overwhelmingly
in southern areas where rainfall is Calluna vulgaris and Empetrum dominant with only occasional or
relatively low. Calluna vulgaris and nigrum, which forms large patches, even no bushes of other species.
Erica cinerea are generally co-domi- are the co-dominants here, especial- Carex arenaria remains constant
nant with Erica cinerea sometimes ly in the cooler wetter north of but grasses may only be represented
colonising first. Rosa pimpinellifolia Britain. Festuca rubra/ovina by a few tufts of Anthoxanthum
is a distinctive invader. Festuca remains common among frequent odoratum or Deschampsia
rubra/ovina is very common with Carex arenaria and Luzula flexuosa. Other herbs are also
scattered shoots of Carex arenaria campestris with other species as sparse. Bryophytes may include
and Luzula campestris. Other herbs listed above. Hypnoid mosses are hypnoid species and Dicranum
are occasional. The distinctive more prominent with Hypnum scoparium.
element is the cryptogams, often cupressiforme, Pleurozium schre-
This sub-community can be found
occupying the bulk of the ground. beri, Rhytidiadelphus triquetrus
throughout the range of H11.
Hypnoid mosses are scarce and most and Hylocomium splendens and
obvious are the lichens, particularly less commonly Ptilidium ciliare.
Cladonia spp., including those listed Lichens are more patchy.
above. Cornicularia aculeata and
This sub-community is found local-
Hypogymnia physodes are also com-
ly in north and east Scotland.
mon, sometimes with Peltigera cani-
na and low-growing Usnea species.
This is the most widespread sub-
community but it is replaced local-
ly by the Empetrum sub-community
in north and east Scotland.

97
H12 Calluna vulgaris – lichens and Polytrichum species can be abundant
in the years following burning.
Vaccinium myrtillus heath This community is the typical sub-shrub
community of acidic to circumneutral, free-draining
This heath is generally dominated by Calluna mineral soils throughout the cold and wet sub-mon-
vulgaris and includes most of the Calluneta from tane zone generally between 200 m and 600 m. The
less oceanic sub-montane areas where burning is soils on which it occurs are widespread throughout
commonly practised, including many grouse this zone, developing from a variety of siliceous par-
moors. Here a predominance of building-phase ent materials, intrusive igneous rock or coarse glacio-
Calluna is found, but a more open cover of degen- fluvial gravels. Despite being free-draining the soils
erate Calluna can often also be present. Vaccinium are normally moist for the majority of the year
myrtillus is constant though it is usually subordi- because of the climate and the superficial pH is usu-
nate to Calluna and is most vigorous out of reach ally between 3.5 and 4.5. It is extensive in the east-
of grazing animals. Vaccinium vitis-idaea is also central Highlands but also important in south-east
found, sometimes with local prominence, and Scotland, the Lake District, parts of Wales and the
Erica cinerea may also be present on drier slopes. South-West Peninsula and the North York Moors. In
Empetrum nigrum ssp. nigrum is frequent, form- places like the southern Pennines, where air pollu-
ing mats after burning, but then becomes reduced tion is severe, it is largely replaced by Calluna vul-
after the Calluna has regrown. garis – Deschampsia flexuosa heath (H9).
In many stands herbs are rare. Only Burning and grazing are the major influences
Deschampsia flexuosa is frequent throughout. on floristics and structure, although climatic and
When grazing is regular there may be additional edaphic difference play some part in determining
herbs including Festuca ovina, Agrostis capillaris, variation within the community. Successional
A. canina, Nardus stricta, Potentilla erecta and developments are usually held in check by
Galium saxatile. burning and grazing and without these most
The ground-layer is often prominent with stands would probably progress to scrub and
bulky mosses characteristic, such as Dicranum woodland. Continuous heavy grazing favours the
scoparium, Pleurozium schreberi, Hypnum loss of sub-shrub vegetation to grassland and in
cupressiforme s.l. and Hylocomium splendens, some instances, particularly after burning, may
together with larger Cladonia species. Encrusting result in the spread of Pteridium aquilinum.

98
 H12

Vaccinium vitis-idaea and Empetrum Vaccinium vitis-idaea and Empetrum

nigrum nigrum frequent, with occasional nigrum nigrum both scarce in rather
Juncus squarrosus and Blechnum spicant. species-poor heath, usually over-
whelmingly dominated by Calluna
vulgaris.
Calluna vulgaris not so abundant as Sub-shrub cover extensive and varied
usual, and other sub-shrubs of generally with Vaccinium vitis-idaea especially
moderate cover in a grassy heath, with frequent and species listed opposite
frequent Festuca ovina, Agrostis capil- typically scarce. Cryptogam flora var-
laris, Nardus stricta, Galium saxatile ied and often abundant, with bulky
and Potentilla erecta and occasional pleurocarps prominent, Hylocomium
Carex pilulifera, Campanula rotundifo- splendens often joining Pleurozium
lia and Polygala serpyllifolia. Lichens schreberi and Hypnum jutlandicum.
not extensive, often patchy. Cladonia impexa, C. uncialis and C.
pyxidata common.

H12c H12b H12a

Galium saxatile – Festuca ovina Vaccinium vitis-idaea – Cladonia Calluna vulgaris sub-community
sub-community impexa sub-community
Vegetation is typically species-poor
This sub-community is found on better This includes most of the richer stands with Calluna vulgaris overwhelmingly
soils and after burning, often followed of this heath, which develop a number dominant and other sub-shrubs of low
by grazing. Calluna vulgaris is less of years after burning. Although cover. Vaccinium myrtillus is very
dominant and with other sub-shrubs Calluna vulgaris is still the general frequent and Erica cinerea common,
forms an open growth within a grassy dominant it is frequently accompanied but both only as scattered shoots.
sward. Deschampsia flexuosa is joined by Vaccinium myrtillus, V. vitis-idaea Other vascular associates are few.
by a variety of herbs including those and Empetrum nigrum nigrum and Deschampsia flexuosa is frequent as
listed above. Where the soils are less occasionally with Erica cinerea. Herbs scattered shoots and sparse plants of
base-poor, species such as Lotus are generally sparse with only scat- Potentilla erecta and Pteridium aquil-
corniculatus, Lathyrus montanus, tered plants of Deschampsia flexuosa, inum are quite common. The ground
Succisa pratensis, Viola riviniana and and occasional Potentilla erecta, cover is not extensive and only
Anemone nemorosa can be locally Juncus squarrosus and Blechnum spi- Dicranum scoparium, Hypnum jut-
abundant. Bryophytes remain quite cant. Bryophytes and lichens are more landicum and Pleurozium schreberi
varied, but lichens are few and of low numerous including the species listed occur frequently as scattered shoots.
cover. above.

99
H13 Calluna vulgaris – Cladonia
Cornicularia aculeata. Among these, bryophytes
arbuscula heath are generally few and rarely of any abundance.
Racomitrium lanuginosum is constant and can
This heath has a dwarfed mat of sub-shrubs with form locally conspicuous patches.
few vascular associates, but with a prominent This heath is the characteristic sub-shrub
lichen flora. Calluna vulgaris is the most frequent vegetation of base-poor soils, over exposed ridges
species, generally prostrate and forming a carpet and summits of mountains, in parts of Britain with
or in wave-like bands or on solifluction terraces. a cold continental climate. It is found on soils
Among other sub-shrubs Empetrum nigrum is with a superficial pH of between 4 and 5, and
most important, usually as ssp. hermaphroditum, frequently a humic surface above pervious acidic
but with ssp. nigrum at lower altitudes. It may be bedrocks and superficials. It is most widespread
intermixed in the mat or forming clumps. through the east-central Highlands of Scotland,
Loiseleuria procumbens is quite frequent and thinning out westwards into the central Grampians
abundant, but Arctostaphylos uva-ursi is at most and north-west Highlands where it is progressive-
occasional. Both Vaccinium myrtillus and ly replaced by its oceanic counterpart Calluna vul-
V. vitis-idaea are common, but always subordinate garis – Racomitrium lanuginosum heath (H14).
in cover. There are a few fragmentary localities in northern
The other vascular associates are few and England and Wales.
sparse. Deschampsia flexuosa and Carex bigelowii It is a vegetation type of unsheltered slopes
are most frequent with species such as Scirpus generally between 600 m and 900 m where there
cespitosus, Agrostis canina and Molinia caerulea are almost constant strong winds which frequent-
at lower altitudes and Juncus trifidus becoming ly clear the ground of snow and subject the vege-
occasional at higher levels. Huperzia selago is also tation not only to reduced precipitation but also to
frequent in higher altitude stands. the effects of frequent and severe frosts and subse-
Lichens are important structurally. Cladonia quent thaws. Burning and grazing may have cur-
arbuscula is especially common and, where there tailed its range in suitable localities in the more
is some shelter, may be abundant. It is usually southerly uplands, but in the eastern Highlands
mixed with C. rangiferina which locally may be the vegetation seems to be largely unaffected by
co-dominant. Also constant are C. uncialis, treatments and the community can be considered
Cetraria islandica, Alectoria nigricans and a climax.

100
 H13

Vaccinium species common and Vaccinium species less fre-

Loiseleuria procumbens frequent in a quent and Loiseleuria
nearly prostrate mat, with Cladonia procumbens and listed
crispata, C. coccifera, Ochrolechia lichens occasional at most.
frigida and Thamnolia vermicularis
often found.

H13c Deschampsia flexuosa very com- Deschampsia flexuosa only

mon with occasional Diphasium occasional and Carex bigelowii
Cladonia crispata – Loiseleuria alpinum and Juncus trifidus. scarce but Erica cinerea and E.
procumbens sub-community Cetraria nivalis frequent and tetralix occasional among the
Cladonia gracilis and sub-shrubs, sometimes
Calluna vulgaris is again the usual C. bellidiflora occasional. with Scirpus cespitosus and
dominant, but one or both subspecies Molinia caerulea. Cetraria
of Empetrum nigrum are common and nivalis and Cladonia gracilis
can be abundant. In the flattened sub- uncommon, but C. arbuscula
shrub mat there is a mixed carpet of and C. rangiferina especially
lichens including the species listed abundant.
above. The herbs comprise scattered H13b
shoots of Carex bigelowii, Huperzia
selago and Deschampsia flexuosa. Empetrum nigrum ssp.
hermaphroditum – Cetraria
nivalis sub-community H13a

On bleak exposed sites at higher Cladonia arbuscula – Cladonia

altitudes lichens remain abun- rangiferina sub-community
dant, but Calluna vulgaris or occa-
sionally Empetrum nigrum her- This sub-community is found at
maphroditum is dominant. Both lower altitudes or more shel-
Vaccinium myrtillus and V. vitis- tered sites where larger Cladonia
idaea are more frequent than in species, as above, are especially
H13a but usually have low cover. abundant, often exceeding the
Loiseleuria procumbens is occa- sub-shrubs in cover. Cladonia
sional. Cladonia arbuscula is still impexa can also be found, but
the most frequent lichen but is C. gracilis and C. crispata are
commonly joined by the species scarce. The most common sub-
listed above. Mosses only make a shrub is Calluna vulgaris but
minor contribution and the few Empetrum nigrum is
vascular associates are present as very common, often as ssp.
scattered individuals. nigrum. Vaccinium species are
only occasional.

101
H14 Calluna vulgaris – out. Cladonia impexa is frequent at lower alti-
tudes and Cladonia gracilis, C. bellidiflora,
Racomitrium lanuginosum heath Cetraria islandica and Ochrolechia frigida occur
often at higher altitudes.
This heath consists essentially of a dwarfed This community is the typical sub-shrub
sub-shrub mat with Calluna vulgaris usually community of base-poor soils on windswept
predominant, together with Racomitrium lanugi- plateaux and ridges at moderate to fairly high
nosum. Other sub-shrubs play a subordinate role, altitudes in the cool oceanic climate of the moun-
but may be common. Most frequent is Empetrum tains of north-west Scotland. It can be found up to
nigrum, with the two subspecies characterising 750 m, although this can extend up to 1000 m in
opposite ends of the altitudinal range (ssp. nigrum the east; to the west and north, on islands like
preferentially common towards lower levels and Skye, Orkney and Shetland, it can extend down to
ssp. hermaphroditum largely confined to higher below 250 m. The community is found on the
altitudes). Erica cinerea is also frequent. base-poor rankers and podzolic soils which are
Other vascular associates are few and usually widespread in this region, with a superficial pH
scattered. Deschampsia flexuosa, Huperzia between 4 and 5 and a humic surface. It is very
selago, Carex pilulifera, Potentilla erecta and much a community of the north-west Highlands
Scirpus cespitosus are all frequent, and Carex with scattered occurrences in the central
bigelowii becomes common at higher altitudes. Grampians.
The extensive woolly carpet of Racomitrium Like its eastern counterpart Calluna vulgaris –
lanuginosum which can be up to 5-10 cm thick is Cladonia arbuscula heath (H13) it is found over
the most noticeable feature of this community. gentle to moderately steep slopes which are
Hypnum cupressiforme s.l. is also very frequent in exposed to fairly constant strong winds that clear
some stands, often with several other mosses and the snow which might otherwise provide shelter
occasional hepatics. Lichens are common and var- in the coldest months. Although it is sometimes
ied but not abundant, and species like Cetraria grazed by sheep and deer, it is unlikely that this
nivalis and Alectoria ochroleuca are absent. factor is important in maintaining the characteris-
Cladonia arbuscula and C. uncialis are the most tic composition and physiognomy, and this vege-
frequent, and Sphaerophorus globosus and tation can be regarded as the natural climax in
Cornicularia aculeata are also common through- such exposed situations in its range.

102
 H14

Arctostaphylos uva-ursi constant with Empetrum nigrum nigrum and

frequent A. alpinus, Empetrum nigrum Erica cinerea at most occasional.
nigrum and Erica cinerea. Scirpus Arctostaphylos species and the herbs
cespitosus common with Molinia and bryophytes listed opposite very
caerulea and Carex binervis scarce.
occasional. Dicranum scoparium,
Diplophyllum albicans and
Pleurozium schreberi frequent.
Empetrum nigrum hermaphroditum Empetrum nigrum hermaphroditum
constant with frequent Nardus stricta and herbs and lichens listed opposite
and Diphasium alpinum and some- occasional at most. Festuca ovina and
times with an extensive lichen cover, Agrostis canina very common, some-
including Cetraria islandica, Cladonia times with Antennaria dioica, Carex
gracilis and Ochrolechia frigida. panicea, Thymus praecox and
Euphrasia micrantha.

H14c

Arctostaphylos uva-ursi
sub-community
H14b H14a
Calluna vulgaris and Racomitrium
Empetrum nigrum ssp. Festuca ovina sub-community
lanuginosum retain the representation
hermaphroditum sub-community
of H14b but the sub-shrubs are
Calluna vulgaris or Racomitrium
more varied with Erica cinerea and
Calluna vulgaris generally abundant lanuginosum or mixtures of the two
Empetrum nigrum ssp. nigrum at
with Racomitrium lanuginosum some- dominate the vegetation mat with
their peak of frequency, and locally
times co-dominant but often subordi- other species playing only a minor
abundant Arctostaphylos uva-ursi and
nate. The variety of herbs characteris- role. Empetrum nigrum ssp. nigrum
A. alpinus. Among the vascular associ-
tic of H14a is not found although and hermaphroditum and Erica
ates Carex bigelowii is very scarce. In
Deschampsia flexuosa and Carex cinerea are only occasional and in
the bryophyte mat Dicranum scopari-
bigelowii are very frequent and some stands Loiseleuria procumbens
um and larger pleurocarps such as
Potentilla erecta, Huperzia selago can be prominent. More striking is
Hypnum cupressiforme s.l. and
and Carex pilulifera remain common. the variety of herbaceous associates.
Pleurozium schreberi make their
Most distinctive are the cryptogams Deschampsia flexuosa is less common
biggest contribution. In the lichen flora
with several pleurocarpous mosses than usual, but Carex bigelowii is fre-
Cladonia impexa is common together
and the typical lichen flora of the quent with C. pilulifera, Huperzia sela-
with the community species.
community with additional species, go and Potentilla erecta. More prefer-
This and the Festuca sub-community including those listed above. ential are the species listed above.
are found at the lowest altitudes and Lichen cover is comparatively low
This is the typical form of this sub-
most sheltered sites occupied by the but Cladonia uncialis is very common.
community and is found at the highest
community.
altitudes of the range of H14. This and the Arctostaphylos sub-com-
munity are found at the lowest alti-
tudes and most sheltered sites occu-
pied by the community.

103
H15 Calluna vulgaris – Juniperus H16 Calluna vulgaris –
communis ssp. nana heath Arctostaphylos uva-ursi heath
Prostrate juniper, referable to Juniperus communis Although Arctostaphylos uva-ursi is found as an
ssp. nana, is occasional in a wide variety of sub- occasional in a variety of heath types, it is most
shrub heaths. Here, however, it is consistently often found in this community, which has a dis-
dominant in the sub-shrub mat, accompanied by a tinct boreal character. Calluna vulgaris is always
small but distinctive group of oceanic hepatics. present and is the most usual dominant, forming a
The mat is generally less than 10 cm high, fairly canopy 20-40 cm high and having a substantial
continuous in the best stands, but it may form a total cover. Arctostaphylos uva-ursi is constant
mosaic with islands of vegetation on tracts of bare and can become modestly abundant in gaps with-
rock and debris. Several other sub-shrubs are well in the heather cover. Erica cinerea is also very
represented: Calluna vulgaris and Erica cinerea common but of low cover. In many stands there is
are especially frequent and the former often fairly some Vaccinium myrtillus and V. vitis-idaea.
abundant. Arctostaphylos uva-ursi and A. alpinus Quite commonly there are small amounts of
are less common, and Empetrum nigrum ssp. her- Genista anglica, but herbaceous associates are few
maphroditum is occasional. except in the Pyrola media – Lathyrus montanus
Vascular associates are typically few and are sub-community. The only constant grass is
usually scattered in the mat. Deschampsia flexuosa, Deschampsia flexuosa and this can be joined by
Scirpus cespitosus and Potentilla erecta are constant, Luzula multiflora and L. pilosa.
with Huperzia selago, Solidago virgaurea, Dactyl- Bryophytes are variable, with the bulkier moss-
orhiza maculata, Polygala serpyllifolia, Succisa es often strongly associated with particular stages
pratensis, and Antennaria dioica more occasional. in the heather regeneration cycle. Hypnum
In some stands the cryptogam flora is similar to jutlandicum, Pleurozium schreberi and Dicranum
other kinds of dwarfed sub-shrub heath. In typical scoparium, however, are very common overall and
examples of this community, however, the species Hylocomium splendens is also a constant through
Racomitrium lanuginosum, Cladonia uncialis, C. imp- much of the community.
exa, Sphaerophorus globosus and Cornicularia Lichens also differ in their representation, with
aculeata, which are common in all these other only Cladonia impexa constant and, in many
kinds of heath, are joined by Pleurozia purpurea, stands, of low cover. Fruticose species such as
Frullania tamarisci and Diplophyllum albicans which C. arbuscula and C. rangiferina tend to follow the
are not. Where the sub-shrub canopy is well-devel- larger pleurocarps in developing among the more
oped the total cover of the cryptogams is much less shady and humid conditions of older heather
than in the typical moss-heaths of the region. canopies. Hypogymnia physodes and, less com-
This heath is confined to humic rankers at monly, Cetraria glauca can be seen on decaying
moderate altitudes in the cool oceanic climate woody stems.
along the western seaboard of the north-west This heath is characteristic of base-poor to cir-
Highlands and some of the Western Isles. Soil cumneutral soils at moderate altitudes, generally
development under this community is typically between 250 m and 600 m altitude, in the cold
rudimentary with just shallow accumulations of continental climate of the east-central Highlands
decaying juniper and bryophyte litter on Cambrian of Scotland. It is found on a variety of acid
quartzite screes. Although perhaps once more soils developed from lime-poor parent material. It
widespread throughout the north-west Highlands, occurs widely but fairly locally through the
the community is now of rather patchy occurrence east-central Highlands with especially good
along the western side of the more northerly moun- representation in Speyside.
tains with especially good stands on Beinn Eighe The community forms an important part of
and Foinaven. The community is replaced in the grouse-moor in the central Highlands and although
continental climate of the east-central Highlands by edaphic differences play some part in determining
the Juniperus communis ssp. communis – Oxalis floristic variation in the community, their effects
acetosella woodland (W19). are often overlain and modified by the influence
It is confined to the lower portion of the altitu- of burning which ultimately maintains this vege-
dinal ranges of the other dwarf sub-shrub heaths tation as a plagioclimax. Stretches of moorland
and, although the vegetation mat is typically blown including stands of the community are often open
clear of snow, is not usually found in the kind of to livestock but there is little information on the
severely exposed situations of which the other impact of grazing on this vegetation.
communities are so characteristic. This communi-
ty is given some protection against the effects of
grazing by the rocky ground on which it is typical-
ly found, but it is readily damaged by burning.

No sub-communities.

104
 H16

Arctostaphylos uva-ursi often exten- Calluna vulgaris usually a strong domi-

sive with Calluna vulgaris, Erica nant with Arctostaphylos uva-ursi subor-
cinerea and Vaccinium species. dinate and associates listed opposite
Festuca ovina common with occasional at most.
occasional Agrostis capillaris and
Anthoxanthum odoratum. The rich
herb flora includes frequent Potentilla
erecta, Pyrola media, Viola riviniana, Vaccinium myrtillus commonly Vaccinium myrtillus absent, V. vitis-
Lathyrus montanus, Hypericum pul- accompanies V. vitis-idaea in a sparse idaea and Empetrum nigrum nigrum
chrum, Anemone nemorosa, Trientalis understorey with Empetrum nigrum scarce, and hypnoid mosses patchy.
europaea, Galium saxatile and Lotus nigrum intermingled. Hypnoid mosses Scirpus cespitosus and Carex pilulifera
corniculatus. often extensive with Cladonia arbus- common and lichens extensive on
cula and C. rangiferina also frequent. areas of bare ground with frequent
Cladonia uncialis, C. impexa, C. flo-
erkeana, C. coccifera and C. squamosa.

H16a H16b H16c

Pyrola media – Lathyrus montanus Vaccinium myrtillus – Vaccinium Cladonia spp. sub-community
sub-community vitis-idaea sub-community
Although Arctostaphylos uva-ursi is
Arctostaphylos uva-ursi is most promi- Arctostaphylos uva-ursi remains con- sometimes quite abundant, Calluna
nent in this sub-community, on less stant here but Calluna vulgaris is usu- vulgaris is more often overwhelmingly
acidic brown earth soils, forming a ally dominant and with the sub-shrubs dominant. Both Erica cinerea and
patchwork with Calluna vulgaris. mentioned above. In contrast with Genista anglica occur frequently.
Erica cinerea can have a high cover H16a, herbs are scarce though Festuca The herbs of sub-community H16a
and the two Vaccinium species are ovina, Carex pilulifera, Potentilla are hardly ever found. Among the
common. Genista anglica is fairly erecta, Luzula multiflora, L. pilosa bryophytes the characteristic mosses
frequent. The most striking feature is and Listera cordata are occasional. Hylocomium splendens and
the associated herb flora developed Cryptogams, however are more Pleurozium schreberi are very
in more mesotrophic conditions. diverse and extensive with the bulky patchy, but Hypnum jutlandicum and
Grasses are more common with pleurocarps Hypnum jutlandicum, Dicranum scoparium remain frequent
Festuca ovina frequently joining Pleurozium schreberi and Hylocomium often with a little Pohlia nutans. Peat-
Deschampsia flexuosa together with splendens often abundant. Larger encrusting lichens are most noticeable,
the herbs listed above. The mosses lichens are also more apparent particularly the Cladonia species
characteristic of H16 are common, but including several Cladonia species. listed above.
not abundant, including the preferential
Rhytidiadelphus triquetrus.

105
H17 Calluna vulgaris – aculeata, Alectoria nigricans and Sphaerophorus
globosus. Mosses are not abundant. Racomitrium
Arctostaphylos alpinus heath lanuginosum is constant, though in small
amounts, and Hypnum jutlandicum becomes
Arctostaphylos alpinus occurs with some frequent at lower altitudes.
frequency in various kinds of dwarfed sub-shrub This heath is the typical climax sub-shrub
heath, but is most typical of this community vegetation of rather base-poor moder soils over
where it is a constant, although usually a subordi- very exposed ridges and crests at moderate to
nate one, in the woody mat which is normally less fairly high altitudes in the cold and humid climate
than 10 cm tall. It is usually dominated by stunted of the mountains of north-west Scotland. It is
bushes of Calluna vulgaris with stretches of bare found at higher altitudes than the Calluna
stones between. Empetrum nigrum ssp. hermaph- vulgaris – Racomitrium lanuginosum heath (H14)
roditum is strongly preferential to higher altitudes which has a similar distribution, and its normal
and ssp. nigrum is largely confined to lower situa- range is between 500 m and 750 m, although it can
tions. Loiseleuria procumbens is characteristically exceptionally be found up to 900 m, and down to
found with E. nigrum ssp. hermaphroditum, and 250 m along the north Scottish coast and on
Erica cinerea with E. nigrum ssp. nigrum. Orkney. It is typically found on humic poor
Vaccinium myrtillus is common throughout but rankers and more occasionally mature podzolised
other Vaccinium species are scarce. soil that have been derived from a variety of
There are few herbs. Huperzia selago is the lime-poor parent materials. This community is
commonest and a constant, and is often accompa- confined to the north-west Highlands, the north
nied, at higher altitudes, by Diphasium alpinum, Scottish coast and Orkney.
Carex bigelowii and Antennaria dioica. Desch- The community may be lightly grazed by
ampsia flexuosa is also frequent throughout, sheep and deer but this probably has little effect
though more so at lower altitudes where Potentilla on its floristics or physiognomy. The inhospitable
erecta, Scirpus cespitosus and Carex pilulifera environment and the harsh conditions maintain
occur most commonly. the vegetation as a climax. Burning is very delete-
More conspicuous are the lichens which form a rious and may cause damage from which recovery
patchy mosaic. Cladonia arbuscula and C. unc- is extremely slow if not impossible. Burning may
ialis are constant. Preferential to higher altitudes have eliminated this community from many sites
are Cetraria glauca, C. islandica, Cornicularia throughout its range.

106
 H17

Empetrum nigrum ssp. hermaphrodi- Loiseleuria procumbens occasional,

tum and Loiseleuria procumbens but Empetrum nigrum ssp. hermaphro-
constant with frequent Carex bigelowii, ditum replaced by ssp. nigrum and
Diphasium alpinum, Antennaria with Erica cinerea becoming common.
dioica, Cladonia uncialis, C. arbuscula, Potentilla erecta very frequent with
C. gracilis, C. pyxidata, C. bellidiflora, Scirpus cespitosus and stricta often
Cetraria glauca, C. islandica, Alectoria present. Cladonia uncialis and
nigricans and Sphaerophorus globosus. C. arbuscula remain common but
lichen flora not so varied or abundant.

H17a H17b

Loiseleuria procumbens – Empetrum nigrum ssp. nigrum

Cetraria glauca sub-community sub-community

In this distinctive sub-community Calluna vulgaris dominates the

mixtures of Calluna vulgaris sub-shrub mat with Arctostaphylos
with subordinate Arctostaphylos alpinus constant and Empetrum
alpinus, Loiseleuria procumbens nigrum ssp. nigrum and Erica cinerea
and Empetrum nigrum ssp. as common associates. Vaccinium
hermaphroditum make up the bulk myrtillus is again sparse with
of the mat with scattered Vaccinium occasional Arctostaphylos uva-ursi.
myrtillus and occasional V. vitis-idaea, Carex bigelowii is less common and
Arctostaphylos uva-ursi, Juniperus Deschampsia flexuosa more frequent
communis nana and Salix herbacea. than in H17a, with the other associates
Deschampsia flexuosa occurs sparsely listed above. Among bryophytes
with the above herbs and other more Hypnum jutlandicum frequently joins
occasional species. Small patches Racomitrium lanuginosum. Lichens
of Racomitrium lanuginosum are are not so varied or abundant as in
frequent and there is a rich and H17a, but Cladonia uncialis and
extensive lichen flora including C. arbuscula remain very common.
the species listed above.

107
H18 Vaccinium myrtillus – rosus, Plagiothecium undulatum, Dicranum
majus and Racomitrium lanuginosum. Some
Deschampsia flexuosa heath lichens occur frequently although an extensive
carpet is never found. The most common species
This community includes a variety of moss-rich are Cladonia arbuscula, C. impexa and C. uncialis.
and grassy sub-shrub vegetation, in which This community is typical of moist but
Vaccinium myrtillus is the most frequent ericoid, free-draining, base-poor to circumneutral soils
with Calluna vulgaris only occasional and often over steeper slopes at moderate to high altitudes
lacking in vigour. Other sub-shrubs can make a through the uplands of northern Britain. It is large-
sizeable contribution to the canopy; in particular ly confined to altitudes above 400 m and can
Empetrum nigrum, usually ssp. hermaphroditum, extend up to 800 m. It occurs over a wide variety
is most frequent, often forming patches. of bedrocks on a variety of soil profiles which
Vaccinium vitis-idaea is also common. have a superficial pH of 3.5-5.5. Typically, howev-
Among vascular associates Deschampsia flexuosa er, the soils have a well-developed humic layer.
and Galium saxatile are constant throughout, with This community is widespread through the
Nardus stricta, Agrostis canina ssp. montana and uplands of Britain but is particular common in
Potentilla erecta all very frequent. In some stands northern Scotland, where the heart of its range
these species account for virtually all the occurs in the central and eastern Highlands, with
herbaceous cover. The grasses Festuca ovina, Agrostis more sporadic occurrences to the north-west.
capillaris and Anthoxanthum odoratum occur at At higher levels this vegetation is probably a
least occasionally and increase in frequency and natural climax with the floristics and distribution
abundance in some sub-communities. of the community being influenced by snow-lie,
The other element, which is usually promi- but towards the sub-montane zone quite extensive
nent, comprises bulky mosses. Dicranum scopari- stands of the vegetation have been biotically
um, Pleurozium schreberi and Hypnum cupressi- derived as a result of woodland clearance and
forme s.l. are very common throughout, Hylo- pasturing. In other places treatments such as burn-
comium splendens is also conspicuous and there ing and grazing have precipitated its spread on to
can be frequent Rhytidiadelphus loreus, R. squar- blanket peats.

108
 H18

Sub-shrubs often co-dominant with Sub-shrubs generally strongly domi-

Alchemilla alpina and/or grasses nant with the listed herbs opposite
among which Festuca ovina, Agrostis occasional at most.
capillaris and Anthoxanthum odora-
tum are especially common. Luzula
campestris, Carex pilulifera and
C. binervis also common with
Potentilla erecta and occasional Calluna vulgaris quite common among Calluna vulgaris sparse, and Blechnum
Campanula rotundifolia, Viola the sub-shrubs with Blechnum spicant spicant absent and pleurocarps rather
riviniana and Ranunculus acris. frequent. Mosses forming an exten- patchy. Racomitrium lanuginosum
sive ground carpet with Hylocomium common and lichens quite conspicu-
splendens and Rhytidiadelphus loreus ous, Cladonia arbuscula being locally
joining Pleurozium schreberi and abundant and C. impexa and
Dicranum scoparium. C. uncialis occasional.

H18b H18a H18c

Alchemilla alpina – Carex pilulifera Hylocomium splendens – Racomitrium lanuginosum – Cladonia

sub-community Rhytidiadelphus loreus spp. sub-community
sub-community
This sub-community is found at lower This is sometimes developed where
altitudes and favours sunnier aspects This sub-community is generally peaty soils have been burned or on
with less humic and sometimes less found in and around snow-beds on degraded blanket peats. Vaccinium
base-poor soils. Grazing often plays acidic soils. Sub-shrubs are generally myrtillus or mixtures of this with
an important part in its development. dominant with Vaccinium myrtillus Empetrum nigrum are usually domi-
Vaccinium myrtillus can be quite usually most abundant, but Empetrum nant, with sparse plants of Calluna
abundant but Empetrum nigrumis nigrum is occasionally co-dominant vulgaris and Vaccinium vitis-idaea.
scarcer than in H18a. Ericoids as a and in some stands with Vaccinium Deschampsia flexuosa and Festuca
whole are often co-dominant with vitis-idaea or V. uliginosum. ovina are very frequent and can be
Alchemilla alpina in the canopy Deschampsia flexuosa is very common abundant, with other grasses only
and/or with grasses which are best and sometimes abundant, but grasses occasional. Carex bigelowii is infre-
represented here. There is a variety are not as abundant as in H18b. A high quent but locally abundant. Apart
of dicotyledonous herbs including the frequency of Blechnum spicant is diag- from Galium saxatile herbaceous
species listed above. Bulkier mosses nostic. Bryophytes attain their most dicotyledons are uncommon. The only
can be quite frequent but not with the diverse and extensive cover here, common pleurocarps are Pleurozium
same abundance and variety as in particularly the species listed above. schreberi and Hypnum cupressiforme
H18a. Lichens are sparse. Lichens are generally much less s.l. Lichens, however, are best repre-
obvious, the only frequent species sented in this sub-community,
This sub-community is especially
being Cladonia arbuscula. particularly the species listed above.
characteristic of the Breadalbane-
Clova region. This sub-community is found through- This sub-community is found through-
out the range of H18. out the range of H18.

109
H19 Vaccinium myrtillus –
frequent or there may only be sparse shoots of
Cladonia arbuscula heath Polytrichum alpestre and P. piliferum.
Much more important are the lichens, particular-
This community consists essentially of a very low ly larger fruticose species such as Cladonia arbuscu-
mat, 5-10 cm high, of sub-shrubs with an abun- la and C. uncialis, both constants, and less common-
dance of lichens, often marking stands with a yel- ly C. rangiferina and C. gracilis, mixtures of which
lowish tinge. Lichens are more extensive and can exceed the sub-shrubs in total cover. Cetraria
dominant than in the Calluna vulgaris – Cladonia islandica and Cornicularia aculeata are also very
arbuscula heath (H13). Calluna vulgaris is uncom- common, often with a variety of other species.
mon overall and Vaccinium myrtillus is the most This heath is typical of base-poor soils on
consistent sub-shrub, being co-dominant in more moderately sheltered and snow-bound slopes at
sheltered situations, although sparse in exposed high altitudes, particularly in the more continen-
sites. Vaccinium vitis-idaea is less common, but tal mountains of northern Britain. The vegetation
constant, and V. uliginosum scarce overall. is strongly montane, being found mainly above
Empetrum nigrum, almost always ssp. hermaph- 650 m up to 1000 m or even beyond, and usually
roditum, is frequent and can exceed Vaccinium stands are found in sites with some shelter so that
species in its cover. there is winter protection from lying snow. It is
Vascular associates are few but Carex bigelowii, characteristic of acid soils with a superficial pH of
a constant, is frequent and often abundant and about 4. It has a similar range to that of H13, being
may be co-dominant with the ericoids and lichens. strongly concentrated in the central and eastern
The other common and constant plant is Highlands of Scotland, particularly the Grampians,
Deschampsia flexuosa. Festuca ovina is also but also in the mountains of the north-west and
fairly frequent together with several herbs, such as the Southern Uplands and with scattered locali-
Galium saxatile, in sub-community H19a. ties in northern England.
Bryophytes are generally not important and Floristic variation reflects differences in expo-
Racomitrium lanuginosum is only abundant in sure and soil type, but overall the vegetation is a
one sub-community. Dicranum fuscescens is quite climatic climax.

110
 H19

Vaccinium myrtillus and Carex Festuca ovina and Galium saxatile can
bigelowii usually co-dominant among occur occasionally but not in any abun-
the vascular plants, with Festuca dance.
ovina/vivipara. Galium saxatile fre-
quent and Potentilla erecta occasional.
Racomitrium lanuginosum sometimes Empetrum nigrum hermaphroditum
quite abundant in a rather patchy very common with lichens very abun-
lichen cover. Alchemilla alpina fre- dant in the mat. Cladonia rangiferina,
quent in small amounts, with occa- C. gracilis and Ochrolechia frigida are
sional Salix herbacea and Juncus preferentially frequent and C. bellidi-
trifidus. flora, C. pyxidata and Cetraria nivalis
more occasional. Racomitrium lanugi-
nosum frequent but not abundant.

H19a H19b H19c

Festuca ovina – Galium saxatile Racomitrium lanuginosum Empetrum nigrum ssp. hermaphrodi-
sub-community sub-community tum – Cladonia spp. sub-community

Vaccinium myrtillus is often abundant The mat is dominated by various Mixed mats of Vaccinium myrtillus
and co-dominant with Carex bigelowii. mixtures of Vaccinium myrtillus, and Empetrum nigrum hermaphrodi-
Empetrum nigrum is sometimes found Carex bigelowii, lichens and tum provide the bulk of the vascular
and Vaccinium vitis-idaea is most abundant Racomitrium lanuginosum. cover, but are often exceeded by the
frequent and abundant here. Empetrum nigrum, mostly ssp. her- lichen cover. Vaccinium vitis-idaea
Deschampsia flexuosa is moderately maphroditum, is fairly common, but can be moderately abundant and there
abundant and Galium saxatile is a Vaccinium vitis-idaea is very patchy. is often a little Calluna vulgaris.
good diagnostic species. The lichen Salix herbacea is occasional as is the Deschampsia flexuosa is the only
element is prominent although not so rare moss Kiaeria starkei but both can common grass and species like Carex
extensive as in sub-community H19c. be locally abundant. Other vascular pilulifera and Galium saxatile are at
Cladonia arbuscula is sometimes co- associates are either occasional, as their most infrequent. Racomitrium
dominant with the vascular plants and above, or very occasional. The lichen lanuginosum is accompanied by pref-
C. uncialis is moderately abundant. cover is varied; Cladonia arbuscula erentially common Pleurozium schre-
can be abundant but may be patchy beri, Dicranum scoparium and
This sub-community is more charac- and mixed carpets are more usual Ptilidium ciliare. Lichens are very
teristic of the lime-rich rocks between with C. uncialis, Cetraria islandica, extensive, with the most common
Breadalbane and Clova, although the Cornicularia aculeata and species listed above.
pH of the soil surface is still strongly Sphaerophorus globosus.
This sub-community is widespread on
acid.
This sub-community is found both in granitic and quartzitic mountains.
central and north-west Scotland.

111
H20 Vaccinium myrtillus –
Racomitrium lanuginosum is very important,
Racomitrium lanuginosum heath forming a woolly carpet, and it is found with a
variety of other bulky mosses. Hypnum cupressi-
This community brings together a variety of forme s.l., Hylocomium splendens, Rhytid-
vegetation types in which Vaccinium myrtillus iadelphus loreus and Pleurozium schreberi are all
and/or Empetrum nigrum ssp. hermaphroditum constant and can be prominent. Additionally,
occur, occasionally with other sub-shrubs such as Polytrichum alpinum and Dicranum scoparium
V. vitis-idaea, and are co-dominant with Racom- are found in many stands. Common hepatics are
itrium lanuginosum or hypnaceous mosses. Ptilidium ciliare and Diplophyllum albicans, but
Vaccinium myrtillus and E. nigrum form a low their greatest variety is found in the Bazzania –
mat, usually less than 10 cm high, appearing as a Mylia sub-community. Lichens are less important,
patchy mosaic of bushes among the moss carpet. but Cladonia uncialis and C. arbuscula are most
At lower altitudes Juniperus communis ssp. nana frequent throughout and may be modestly
and Erica cinerea can show local prominence. abundant, with C. gracilis and Cetraria islandica
Among vascular associates Carex bigelowii, also common.
Festuca ovina/vivipara, Deschampsia flexuosa This heath is characteristic of humic, base-poor
and Galium saxatile are all constant and frequent. soils on fairly exposed slopes and summits at
There are few other common herbs, although the moderate to high altitudes, in the cool oceanic
grasses may include frequent Nardus stricta. mountains of north-west Scotland, extending to
Huperzia selago and Potentilla erecta are frequent Skye, and scattered through the Grampians.
in some stands and may be accompanied by Almost always, the bedrocks underlying this
Thymus praecox, Viola riviniana and Carex heath are siliceous in character.
pilulifera. Climatic differences and some modest varia-
Much of the distinctive character of this tion in edaphic conditions influence the floristics,
vegetation type depends on the cryptogams. but this is essentially climax vegetation.

112
 H20

Vaccinium vitis-idaea very scarce, but Vaccinium vitis-idaea very common

Calluna vulgaris, Erica cinerea and and all other listed associates scarce.
Juniperus communis nana occasional
with Alchemilla alpina sometimes co-
dominant. Potentilla erecta, Thymus Rich and luxuriant patchwork of Diplophyllum albicans and Anastrepta
praecox, Viola riviniana and Carex bryophytes present among the orcadensis sometimes found but com-
pilulifera frequent. Racomitrium lanuginosum carpet with binations of these other bryophytes
Dicranum scoparium, Plagiothecium rare.
undulatum, Sphagnum capillifolium,
Mylia taylori, Diplophyllum albicans, Racomitrium lanuginosum reduced in Bulky pleurocarps can be common
Pleurozia purpurea, Bazzania tricrena- cover with dominance often passing to but not abundant among the dominant
ta, Scapania gracilis, S. ornithopodi- mixtures of Pleurozium schreberi, Racomitrium lanuginosum carpet, but
oides, Anastrepta orcadensis and Hylocomium splendens and the lichen carpet is richer with fre-
Anthelia julacea frequent. Rhytidiadelphus loreus. quent Cladonia gracilis, Cetraria
islandica and Cornicularia aculeata,
and occasional Cladonia leucophaea,
Sphaerophorus globosus and Alectoria
nigricans.
H20a H20c

Viola riviniana – Thymus praecox Bazzania tricrenata – Mylia taylori H20d H20b
sub-community sub-community
Rhytidiadelphus loreus – Cetraria islandica sub-community
The sub-shrub mat is more varied Here the general floristic features are Hylocomium splendens
than usual in the community. as in H20b though Blechnum spicant sub-community Mixtures of Empetrum nigrum ssp.
Empetrum nigrum hermaphroditum and Juncus trifidus are frequent. The hermaphroditum and Vaccinium myr-
and Vaccinium myrtillus can both be community mosses are well represent- Although Racomitrium lanuginosum is tillus with abundant Racomitrium
abundant, but Alchemilla alpina can ed but distinctively there is a range of generally reduced in cover, the general lanuginosum are usually dominant.
be co-dominant and more locally conspicuous bryophytes, including vegetation features accord well with Grasses like Festuca ovina/vivipara,
Juncus communis ssp. nana or Erica Atlantic hepatics as listed above. the community as a whole. Vaccinium Deschampsia flexuosa, Nardus stricta,
cinerea may occur with sparse shoots Lichens are generally sparse. vitis-idaea is preferentially common, Agrostis canina and Anthoxanthum
of Calluna vulgaris. Herbs characteris- Carex bigelowii and Deschampsia flex- odoratum tend to be prominent, but
This sub-community is very much uosa are frequent and Galium saxatile the richness of vascular plants in H20a
tic of H20, such as Festuca ovina/ confined to the wettest regions and
vivipara, Deschampsia flexuosa and and Festuca ovina/vivipara are more is absent. Bryophytes are not numer-
even then it is restricted to suitable occasional. The moss mat is distinctive ous although community constants are
Galium saxatile, remain very frequent cold and damp aspects.
and are joined preferentially by the with dominance passing to the species all common with frequent Hypnum
herbs listed above. The cryptogam listed above. The Atlantic hepatics cupressiforme s.l. and Polytrichum
flora is poor, with Racomitrium found in H20c are absent. alpinum. The lichen flora is a little
lanuginosum as the typical dominant, richer than usual with the species
This sub-community is local through-
but other pleurocarps are less frequent listed above present.
out the range of H20.
and Diplophyllum albicans is the only This is the most widespread and com-
common hepatic. mon sub-community overall.
This sub-community extends the range
of H20 into the milder foothills of the
western seaboard and Skye.

113
H21 Calluna vulgaris –
locus for the ‘mixed northern hepatic mat’.
Vaccinium myrtillus – Sphagnum Species such as Scapania gracilis, Mylia
capillifolium heath taylori and Diplophyllum albicans can be found
throughout, but the Mastigophora – Herbertus
This community has a mixed canopy of sub- sub-community has an additional range of Atlantic
shrubs, usually 30-50 cm high, with a damp layer species, forming a unique vegetation found at high-
of luxuriant bryophytes. Calluna vulgaris is usual- er altitudes in north-west Scotland where summer
ly the dominant ericoid, although Vaccinium myr- temperatures are lower and rainfall higher. Lichens
tillus is constant and Empetrum nigrum, almost are fairly insignificant, Cladonia impexa being the
always ssp. hermaphroditum, very frequent. Erica only species occurring commonly throughout.
cinerea is also frequent, but patchy. Other sub- This heath is highly characteristic of fragmen-
shrubs are only occasional. tary humic soils, developed in situations with a
Deschampsia flexuosa and Potentilla erecta are cool but equable climate and a consistently shady
constant and very common though usually present and extremely humid atmosphere. It is almost
as sparse, scattered individuals. More distinctive- wholly confined to low to moderate altitudes
ly Blechnum spicant is constant and Solidago through the oceanic mountains of north-west
virgaurea and Listera cordata frequent. There are Scotland and on Skye, with outliers on Orkney, in
only occasional records for other vascular south-west Scotland and the Lake District.
associates. It is largely restricted to steep, sunless slopes of
The bryophytes form an extensive and lush car- north-west to easterly aspect, often with rock
pet. Constant throughout are bulky hypnaceous outcrops and blocky talus, among which crevices
mosses such as Hypnum cupressiforme s.l., provide additional shade. In some situations this
Rhytidiadelphus loreus, Pleurozium schreberi and may not be a climax community but a result of
Hylocomium splendens, with Plagiothecium undu- woodland clearance, but towards the upper end of
latum, Dicranum scoparium and D. majus also its altitudinal limits this heath appears to form a
very common. Particularly distinctive is the high natural component of vegetation patterns con-
frequency and local abundance of Sphagnum trolled largely by variations in local climates and
capillifolium. Racomitrium lanuginosum becomes soils. It is sometimes lightly grazed, but burning
more frequent at higher altitudes. The most spec- is very damaging and recovery is probably
tacular enrichment in this element comes from extremely slow. It seems certain that the extent of
oceanic hepatics and this community is a major this community has been reduced by burning.

114
 H21

Empetrum nigrum hermaphroditum frequent and locally Empetrum nigrum hermaphroditum local and combina-
abundant among the sub-shrubs with especially rich and tions of listed cryptogams rare, but Dicranum scoparium
luxuriant cryptogam carpets among which there is frequent common with frequent fronds of Pteridium aquilinum.
Racomitrium lanuginosum, Mylia taylori, Scapania gracilis,
Bazzania tricrenata, Pleurozia purpurea, Diplophyllum
albicans, Anastrepta orcadensis, Mastigophora woodsii,
Herbertus aduncus hutchinsiae, Cladonia uncialis and
C. arbuscula.

H21b H21a

Mastigophora woodsii – Herbertus aduncus ssp. Calluna vulgaris – Pteridium aquilinum

hutchinsiae sub-community sub-community

Calluna vulgaris is usually the most abundant sub-shrub, This sub-community occurs in sites which cannot support
but the canopy is short and more mixed than in H21a. The the full range of hepatics. Calluna vulgaris is generally a
bryophytes are extremely well developed. Among the moss- strong dominant in this taller and more species-poor
es all the community constants occur frequently. The hepat- heath. Vaccinium myrtillus is very common with Erica
ics, however, are most abundant, tingeing the vegetation cinerea and Vaccinium vitis-idaea occasional. Other vascu-
with a variety of colours. They include the species listed lar plants are sparse, but distinctive is Pteridium aquil-
above with other rarer Atlantic hepatics. inum with occasional Oxalis acetosella, Viola riviniana
and Luzula sylvatica. Bryophytes can have fairly high
This sub-community is restricted in range, being confined
cover, but comprise almost entirely the community
to the more shaded and humid habitats in north-west
constants.
Scotland.
This sub-community is found throughout the range of H21.

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H22 Vaccinium myrtillus – especially common and several other species
locally abundant. A variety of other mosses and
Rubus chamaemorus heath hepatics are variable in their occurrence. Lichens
are typically less prominent, although Cladonia
This heath has a mixed cover of sub-shrubs over a arbuscula is constant and can show modest
moist cover of bryophytes similar to that abundance.
of Calluna vulgaris – Vaccinium myrtillus – This heath is characteristic of wet, base-poor
Sphagnum capillifolium heath (H21). However, peats at moderate to high altitudes (mainly between
here the canopy is not as tall, being mostly 500 m and 800 m), where there is protection against
between 10 and 30 cm high, and Calluna vulgaris extremes of dryness and winter cold by virtue of an
is not invariable in its dominance (Vaccinium oceanic influence or locally prolonged snow-lie.
myrtillus is dominant in the Polytrichum – Galium The profiles found beneath this community are
sub-community). Empetrum nigrum ssp. her- typically poorly-developed, often consisting of
maphroditum is constant, as is V. vitis-idaea just a layer of bryophyte or ericoid humus resting
(although less frequent), and V. uliginosum is rare. directly on blocky talus, derived from a variety of
Erica cinerea is absent. pervious bedrocks. It is almost entirely confined to
The vascular associates are distinctive because, the central and north-west Highlands of Scotland.
with constant Deschampsia flexuosa, there is In the former region it is typical of early snow-beds
frequently a little Rubus chamaemorus and where it is mainly present as the Polytrichum –
Cornus suecica. Eriophorum vaginatum can be Galium sub-community. In the north-west High-
locally abundant and there are records for lands, where the climate is ameliorated by the
Potentilla erecta, Melampyrum pratense, oceanic climate, this heath is generally represented
Listera cordata, Juncus squarrosus and Nardus by the Plagiothecium – Anastrepta sub-community.
stricta. Climatic and edaphic factors maintain this
Bryophytes are always conspicuous and heath as a climax vegetation in most situations,
sometimes very abundant. Dicranum scoparium although at its lowest limits it falls within the alti-
and the hypnaceous mosses Pleurozium schreberi, tudinal range of historical pine forest. It is some-
Hylocomium splendens and Rhytidiadelphus times affected by grazing and burning where these
loreus are the most consistent and constant, treatments are applied to the surrounding heaths.
although Sphagnum spp. can also have a high Burning is deleterious to the floristic richness of
cover, with the constant S. capillifolium being the community.

116
 H22

Carex bigelowii frequent in small amounts with a rich Carex bigelowii and cryptogams listed opposite all very
and extensive patchwork of cryptograms, among which scarce, but Gallium saxatile and Blechnum spicant frequent
Racomitrium lanuginosum, Plagiothecium undulatum, as scattered individuals and Polytrichum commune very
Ptilidium ciliare, Anastrepta orcadensis, Barbilophozia common among usually plentiful hypnaceous mosses.
floerkii, Cladonia bellidiflora, C. uncialis, C. leucophaea,
C. gracilis and C. impexa are very common.

H22b H22a

Plagiothecium undulatum – Anastrepta orcadensis Polytrichum commune – Galium saxatile sub-community

sub-community
Vaccinium myrtillus is generally dominant in a low sub-
Calluna vulgaris is often a strong dominant in a taller shrub canopy with Calluna vulgaris and/or Empetrum
canopy with Empetrum nigrum ssp. hermaphroditum nigrum ssp. hermaphroditum sub-dominant. Vaccinium
occasionally abundant. Vaccinium myrtillus usually has low vitis-idaea is fairly common but of low cover. Cornus sueci-
cover and V. vitis-idaea occurs unevenly. Cornus suecica, ca and Rubus chamaemorus are most consistently frequent
Rubus chamaemorus and Deschampsia flexuosa are all here though not abundant. Deschampsia flexuosa is rather
more patchy than in H22a, though Eriophorum vaginatum is patchy and Blechnum spicant and Galium saxatile are
common as scattered shoots. Carex bigelowii and Huperzia preferentially common. Hypnaceous mosses, especially
selago are preferential at low frequencies. The cryptogams Hylocomium splendens and Rhytidiadelphus loreus, are
are distinctive with both hypnaceous mosses and plentiful with Sphagnum capillifolium patchily abundant.
Sphagnum spp. prominent together with a number of Lichens are rare apart from scattered Cladonia arbuscula.
Atlantic hepatics including the species listed above. Lichens
This sub-community is largely restricted to the central
are also more numerous in this sub-community with the
Highlands.
above species present and Cladonia arbuscula constant.
This sub-community occurs throughout the range of H22
and is particularly well-developed in the north-west
Highlands.

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through which the three country nature conservation agencies –
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