Further informationBooksBritish Mosses and liverworts: a fi eld guide. British Bryological Society (2010). The fi rst comprehensive colour fi eld guide to

bryophytes with good keys and hundreds of photos.

Mosses and Liverworts; Gordon Rothero (2005). A brief, general introduction to Scottish bryophytes, part of the ‘Naturally

Scottish’ series published by Scottish Natural Heritage, Battleby.

Bryophytes of native woods – a fi eld guide to common mosses and liverworts of Scotland’s native woods. Carol L Crawford

(2002), Native Woodlands Discussion Group. A small booklet with good colour photos.

Mosses and Liverworts. New Naturalist 97, Porley RD & Hodgetts NG, (2005). Collins. An accessible account of our bryophyte

heritage with a good section on woodlands.

The geographical relationships of British and Irish bryophytes; Hill MO & Preston CD (1998). Journal of Bryology, 20: 127-226.

Information and advicewww.britishbryologicalsociety.org.uk. The British Bryological Society has an excellent website with useful information on

publications, courses, fi eld meetings and lots of pictures.

www.nwdg.org.uk. The Native Woodland Discussion Group runs courses on Atlantic mosses and liverworts.

Advice and SupportPlantlife Scotland can help you in your quest for information and support.

Plantlife Scotland, Balallan House, Allan Park, Stirling, FK8 2QG

Tel: +44 (0) 1786 478509 www.plantlife.org.uk Scotland@plantlife.org.uk

© March 2010 ISBN 978-1-907141-24-9

Plantlife Scotland is part of Plantlife International – the Wild Plant Conservation charity, a charitable company limited by guarantee. Registered in

Scotland (SC038951) and in England and Wales (1059559). Registered company no 3166339.

This guide has been written and illustrated for Plantlife Scotland by Gordon Rothero.

All photos © Gordon Rothero unless otherwise stated. Cover photo © Laurie Campbell.

LIVERWORTS

Key features for identifying liverwortsGrowth form. There are two sorts of liverworts; leafy liverworts have a stem and leaves and resemble a moss, whereas thallose or thalloid liverworts have a simple strap of tissue with no stem or leaves. Leafy liverworts can form erect cushions and turfs while some are creeping and closely apressed to rock or tree. The size of the plant is also important; a number of oceanic liverworts are very, very small.

Leaf shape. This is all-important in leafy liverworts and is much more variable than in mosses. Liverwort leaves can be simple and round, they can be deeply divided into fi laments or into broader lobes, the lobes can be of different sizes and can be bent over or under each other and formed into fl aps or pouches. Many liverworts also have pronounced teeth on the margin of the leaf, visible without a hand lens. Many leafy liverworts also have under-leaves, usually much smaller than the main leaves, and on the lower side of the stem.

Abbreviations

NR – nationally rare; NS – nationally scarce; BAP – a Biodiversity Action Plan species; S8 – listed on Schedule 8 of the Wildlife and Countryside Act (1981).

Photoset Left - A variety of leafy liverwort leaves, clockwise

from top left: Taylor’s Flapwort (Mylia taylori), Prickly

Featherwort (Plagiochila spinulosa), Ciliated Fringewort

(Ptilidium ciliare), White earwort (Diplophyllum albicans -

note upper lobe bent over the top of the lower and the line

of longer cells), Common Paw-wort (Barbilophozia fl oerkei),

Bifi d Crestwort (Lophocolea bidentata), Creeping Fingerwort

(Lepidozia reptans), part of stem with ‘fi ngered’ leaves.

Above - Left: a thallose liverwort, Overleaf Pellia (Pellia

epiphylla) with fruits; right: a leafy liverwort with round

leaves, Autumn Flapwort (Jamesoniella autumnalis).

Key features for identifying Mosses

Growth form. Leaving aside the very distinctive bog-

mosses (Sphagnum), mosses can be split into two

groups, acrocarpous and pleurocarpous. There is a

technical difference between these two forms but in

practical terms, acrocarps usually have erect stems

and grow in cushions or turfs while pleurocarps tend

to grow with main stems parallel to the ground (or rock

or tree trunk) and form wefts. The often dense growth

form of acrocarps means that their sparse branches

are obscured while in pleurocarps the branches are

usually many and easily seen.

Branching. For some of the pleurocarps it is useful

to note what the pattern of branching looks like. Is

it regular with branches more or less opposite each

other on the main stem (pinnate) or irregular? Are

the regular branches branched again (bi-pinnate) and

again (tri-pinnate) giving a fern-like structure?

Colour. Colour and texture are all-important fi eld

characters. Many species have a particular shade,

admittedly usually of green, which coupled with

the structure of stem and leaf, gives a texture which

is what the eye picks up from a distance. Another

important colour is that of the main stem; when

the leaves are dry they become opaque and it may

be necessary to scrape away some leaves with your

fi ngernail to see the stem colour.

Leaf shape. Moss leaves have a variety of shapes but

they tend to be variations on the same theme, with

a relatively broad base tapering to a narrower apex.

Some leaves are long and narrow and taper to a fi ne

point, others have a broad triangular shape tapering

shortly to a sharp point. Other mosses have leaves

with a blunt apex and a few species have round leaves.

Another useful character is whether or not the leaves

are all curved in the same direction (falcate) or bent

back from the stem (refl exed or squarrose). Some

leaves also have teeth on the margin, usually visible

only with a hand lens.

Nerve (or costa). A very useful character is whether

the moss leaf has a nerve or not. The nerve (or costa)

is a thickened rib of tissue running up the centre of

the leaf which looks like a dark line if the leaf is held

against the light and viewed with the hand-lens. It

usually extends beyond halfway up the leaf and may

reach the apex.

Above - Left: acrocarpous Scott’s Fork moss (Dicranum

scottianum) ; right: pleurocarpous Larger Mouse-tail

Moss (Isothecium alopecuroides).

Above - Irregular branching in Red-stemmed Feather-

moss (Pleurozium schreberi) on the left and regular

tri-pinnate branching in Glittering Wood-moss

(Hylocomium splendens) on the right; note the red stems.

Photoset above - A variety of moss-leaf shapes, clockwise

from top left: Dotted Thyme-moss (Rhizomnium

punctatum), Catherine’s Moss (Atrichum undulatum),

Little Shaggy-moss (Rhytidiadelphus loreus), Yellow

Fringe-moss (Racomitrium aciculare), Common Striated

Feather-moss (Eurhynchium striatum), Cypress-leaved

Plait-moss (Hypnum cupressiforme).

Moss or liverwort?Initially this is a tricky question but with a little experience it ceases to be a problem. Thallose liverworts are

easy but leafy liverworts can be passed over as mosses by the uninitiated. In most mosses the leaves grow all

around the stem but in most liverworts the main leaves are in two ranks down each side of the stem, sometimes

with a line of smaller under-leaves below. Most moss leaves are roughly triangular, wide at the base and narrow

to the apex; few liverworts are like that. Moss leaves never have lobes whereas many leafy liverworts do. Most

mosses with round leaves have a nerve; no leafy liverworts have a nerve but a few have lines of cells running up

the centre of the leaf.

MOSSES

Leafy liverwort NS; size: very small and forming thin patches

of upright stems; colour: yellow or yellow green with shoot

tips red with gemmae; leaves: tiny with two lobes but hardly

visible even with a lens; habitat: on well-rotted, rather damp

logs; note: not easy to see without putting your nose to the

log but once spotted easily recognised by the erect stems

with red tips.

Leafy liverwort NS; size: small and forming loose patches

of normally procumbent stems; colour: mid or dark green

with some shoot tips red with gemmae; leaves: rectangular

in outline but with two pointed lobes with an acute gap

between them, the leaves near the stem apex having dark red

gemmae (lens); habitat: often present on the older, sprawling

stems of juniper and also on thin peaty soil on rocks; note:

the red gemmae and the relatively long sharp lobes are

characteristic.

Leafy liverwort NR; size: medium sized but often forming

large dense cushions of erect stems; colour: yellow or

greenish brown, occasionally with a reddish tinge; leaves:

complex, two lobed with the smaller upper lobe folded up over

the larger lower lobe, the lower lobe is concave and the top

edge of the upper lobe is turned in towards the stem; habitat:

on thin peaty soil on rocks and in scree; note: a rare species

of scree, the dense patches of erect stems with closely set,

bilobed leaves are usually easily identifi ed.

Leafy liverwort; size: small to medium-sized forming tight

patches of erect stems; colour: mid or yellow green, often with

some shoot tips red with gemmae; leaves: rounded-rectangular

in outline but with three or four lobes, each lobe usually ending

in a little sharp point (lens); underleaves: usually visible with

a lens, with two long pointed lobes; habitat: on thin peaty soil

on rocks and in scree; note: a very similar and equally common

species, Common Paw-wort (Barbilophozia fl oerkei), never has

gemmae and lacks points on the lobes

Leafy liverwort NS; size: small to medium sized but often

forming large dense cushions of erect stems; colour: yellow

or sometimes orange or dark brown; leaves: inserted close

together across the stem with up to four long pointed lobes with

a narrow gap in between, lobes erect and pointing up the stem;

underleaves: large and with two lobes; habitat: on thin peaty

soil on rocks and in scree; note: a characteristic species of dry

scree in eastern hills and readily recognised by the tight cushions

of erect stems and narrow lobes on the closely-set leaves.

Leafy liverwort; size: medium-sized but often forming large

cushions or patches; branching: irregularly pinnate, occasionally bi-

pinnate, at right-angles to the stem, the branches blunt at the end

colour: usually some shade of yellow but may be variegated red,

green or brownish; leaves: two lobes but the most noticeable feature

is the margin which has lots of long narrow teeth (cilia) easily visible

with a lens; underleaves: similar in shape to the main leaves but

much smaller; habitat: on well-drained sites with other bryophytes

amongst ericaceous shrubs, in heathy grassland and in woodland;

note: confusion only possible with the related but much smaller

Tree Fringewort (Ptilidium pulcherimum) and Wood’s Whipwort

(Mastigophora woodsii) a scarce western oceanic species of humid

sites which has longer branches tapering to a point.

Heller’s Notchwort (Anastrophyllum hellerianum)

Horned Flap-wort (Lophozia longidens)

Curled Notchwort (Anastrophyllum saxicola)

Hatcher’s Paw-wort (Barbilophozia hatcheri)

Monster Paw-wort (Tetralophozia setiformis)

Ciliated Fringewort (Ptilidium ciliare)

LIVERWORTS

Mic

ha

el

Lü

th

Mic

ha

el

Lü

th

Mic

ha

el

Lü

th

Bryophytes of

Scotland’s pine woodlands

British Lichen Society

Introduction

This Plantlife fi eld guide will help those who want to

go a little further in identifying the carpets of mosses

and liverworts that are such an obvious feature of

many of our Scots pine woodlands. This guide deals

with some of the typical species and a few of the more

uncommon and rare species of particular habitats

within the woodland. It is focused largely on the

woodlands of the east of Scotland, so, for the pine

woods of the west, particularly Beinn Eighe, the guides

for Atlantic woodland would be more useful.

What are pine woodlands?

They are semi-natural stands of woodland where Scots

pine forms a signifi cant proportion of the canopy.

Within these woodlands pine or birch may be the

dominant tree and there will also be rowan, aspen

and more locally alder and willows, especially along

water-courses. Some woodlands, mostly in the east

of Scotland, have a patchy understory of juniper and

occasionally of hazel.

What are mosses and liverworts?

Mosses and liverworts (collectively known as

bryophytes) are two of the oldest groups of land plants

and have had millions of years to evolve a variety of

species that have colonised almost all habitats apart

from the sea. Most have a simple structure with a

main stem and more or less frequent branches covered

in leaves. They do not have roots but absorb water and

minerals directly into the (usually) single layer of cells

in the leaves. Though some bryophytes are strongly

coloured, often red or purple, most are some shade

of green and an appreciation of the many shades of

green is a useful quality in anyone wishing to identify

these small plants.

Why are the mosses and liverworts important?

Bryophytes in woodland are an excellent indicator

of habitat quality and contribute much to the

functioning of the woodland ecosystem, as well as

giving character and aesthetic appeal. The UK has

some 1100 species of bryophyte, approximately 65%

of the European fl ora; in comparison our vascular

plants total only 15%. Our remnants of Atlantic

woodland have as great a diversity of bryophytes as

almost anywhere else on the planet. Our pinewoods

are less diverse but have a more continental fl ora and

affi nities with the forests of Scandinavia and include a

number of Biodiversity Action Plan (BAP) species that

are rare or absent elsewhere in the UK.

Bryophyte communities

A number of the plants illustrated in this guide are

very precise as to the habitat they need but it is

possible to group species into broad communities

within the pinewoods. It needs to be emphasised that

there will be an overlap as large woodland fl oor species

can cover rocks and the species on rocks and the bases

of trees are often the same.

Woodland fl oor: on the soil of the woodland fl oor,

species have to compete with fl owering plants and

with the accumulation of leaf litter and so the mosses

here tend to be large and relatively fast-growing.

Where the woodland is heathy, as is common

under pine and birch, the dominant bryophytes are

large pleurocarpous species like Glittering Wood-

moss (Hylocomium splendens), Big Shaggy-moss

(Rhytidiadelphus triquetrus), Red-stemmed Feather-

moss (Pleurozium schreberi), Heath Plait-moss

(Hypnum jutlandicum), Ostrich-plume Feather-moss

(Ptilium crista-castrensis) and, in more humid woods,

Red Bog-moss (Sphagnum capillifolium), Little

Shaggy-moss (Rhytidiadelphus loreus) and Waved

Silk-moss (Plagiothecium undulatum). These species

are often abundant and occur to the exclusion of all

else. In drier woodland with a more complete pine

canopy, the fl oor may be more open with areas of bare

pine needles; here there are often dense cushions of

Broom Fork-moss (Dicranum scoparium) and Dusky

Fork-moss (Dicranum fuscescens) running up onto the

tree bases.

Rocks, crags and scree: Within pine woodland there

are often areas of crag and scree which have some

characteristic species. Grey carpets of Woolly Fringe-

moss (Racomitrium lanuginosum) are common and

cushions of Dusky Fork-moss (Dicranum fuscescens)

are frequent and there may be patches of the liverwort

Hatcher’s Paw-wort (Barbilophozia hatcheri). Two

characteristic species of scree are the liverworts

Monster Paw-wort (Tetralophozia setiformis) which

is quite common here and the rare Curled Notchwort

(Anastrophyllum saxicola).

Trees and logs: Bryophytes do not like pine bark and

are usually limited to the base of the tree. The fl ora on

birch is also poor but where there are hazels or aspens

the fl ora can be quite rich, particularly with species

of Bristle-mosses (Orthotrichum). Dead wood can

often provide a humid habitat and has an interesting

fl ora; in damp places species like Wood-rust (Nowellia

curvifolia) and Palmate Germanderwort (Riccardia

palmata) are common and good logs may have the

tiny liverwort Heller’s Notchwort (Anastrophyllum

hellerianum) and the rare moss Green Shield-moss

(Buxbaumia viridis).

‘Bryologising’

Although identifying species initially needs a

modicum of determination, it is possible to quite

quickly become familiar with the common mosses and

liverworts in pine woodlands. They have the virtue of

being available all the year round and grow in really

nice places. Though some of the species are very

distinct, even from some metres away, the process

and the enjoyment will be enhanced if you get used

to using a hand-lens (at least x10) to reveal the fi ner

features on which identifi cation sometimes depends.

In the descriptions, where a lens is necessary, it is

indicated by (lens); remember, hand-lens to the eye

and move the plant into focus.

In these guides, English names have been used

alongside the Latin; these are not ‘common names’

in the same sense as those for fl owers because they

are all recent inventions and as yet are hardly used

except in publications like this! Latin names should

always be used for recording purposes to avoid any

ambiguity.

MOSSES

Pleurocarpous; size: robust and usually forming

large patches; branching: sparse and irregular;

colour: typically a whitish green; stem: green;

leaves: rounded-triangular, tapering shortly to

a point and with distinct undulations running

across the leaf, no nerve; habitat: woodland fl oor,

ledges, boulder tops; note: the whitish-green colour

resembling ‘white worms’ and the undulate leaves

make this an easy moss to recognise.

Pleurocarpous; size: robust and usually forming

large patches; branching: irregular and rather

untidy; colour: mid to yellow-green and ‘chaffy’

when dry; stem: red; leaves: rather rounded-

triangular, tapering to a blunt point, no nerve;

habitat: woodland fl oor, ledges, boulder tops

Acrocarpous; size: very large and forming large

hummocks or lawns; branching: sparse and

obscure; colour: dark green; leaves: a white base

that clasps the stem, above narrowly triangular to

a shortly pointed apex and strongly toothed; the

leaves are opaque and look thick because of the

numerous lamellae on the upper surface; habitat:

in damper areas and boggy ground; note: the

largest of several similar species in the woodlands;

also illustrated is Bristly Haircap (Polytrichum

piliferum) a much smaller plant with a white hair

point which grows in drier places like rock tops

(N.B. the pictures are not to scale).

Pleurocarpous; size: robust and often forming

large patches; branching: very closely pinnate and

so feather-like hence the English name; colour:

mid to yellow-green; stem: green and furry with

green fi laments under the lens; leaves: narrowly

triangular, tapering to a long fi ne point, no nerve,

all strongly curved in one direction; habitat:

woodland fl oor, ledges, boulder tops; note: a

beautiful and easily recognisable moss and very

characteristic of pinewoods.

Pleurocarpous; size: very robust and often

forming large patches; branching: irregular

and rather untidy; colour: yellow-green, straw-

coloured when dry; stem: red but often obscured

by the leaves; leaves: triangular, pale yellow-green

and again rather untidy; habitat: woodland fl oor,

ledges, boulder tops; note: easily recognised by

the large, erect stems and untidy, ‘chaffy’ leaves.

Pleurocarpous; size: robust and usually forming

large patches; branching: irregular; colour: mid to

yellow-green; stem: red, springy; leaves: broadly

triangular, no nerve; curved in one direction

so that the stem ends look hooked; habitat:

woodland fl oor, ledges, boulder tops; note: a bit

smaller, neater and greener than Big Shaggy-moss

(Rhytidiadelphus triquetrus), with hooked shoot-

tips, and usually in more humid sites, so more

frequent in the western woods.

Bog-moss; size: medium sized and forming

swelling cushions or hummocks of erect

stems and often as a loose turf under heather;

branching: a bunch of tight branches at the top

of the stem (capitulum) and whorls of branches

below; colour: often red or pink but variegated

green in more shaded sites; leaves: branch leaves

are narrowly triangular ; habitat: an abundant

and locally dominant plant in more humid

woodland, in valleys or on N-facing slopes note:

other species of Bog-moss may occur in the

woodland but this is the most frequent species.

NR, BAP, S8; fruit: The leaves of this strange moss

are too tiny to see and it is only the large capsule

that attracts attention, resembling a ‘bug-on-a-

stick’. This capsule is 5mm or so long on a stem of

about the same length and held erect when young

and at more of an angle when mature. Bright green

in the winter, brown and peeling when mature in

the summer; habitat: most sites are on rotting logs

or stumps but it has been found on old wood-ant

nests and rarely on bark of living trees; note: a

Schedule 8 species so should not be disturbed.

Acrocarpous; size: medium-sized and usually

forming cushions and occasionally tight turfs;

branching: obscured; colour: mid-green, leaves:

long, very narrowly triangular with a toothed apex;

leaves often curved in one direction; habitat: on

drier open parts of the woodland fl oor, often on pine

litter in bare areas and often around the base of pine

trees, but also on rocks and logs; note: the much

larger Greater Fork-moss (Dicranum majus) is also

common in more humid woodland, it has very long,

regularly curved leaves.

Acrocarpous; size: medium-sized and usually

forming cushions and occasionally tight turfs;

branching: obscured; colour: mid-to dark green,

leaves: long, very narrowly triangular with a

fi ne, toothed apex (lens), leaves often curved in

one direction; habitat: on drier open parts of the

woodland fl oor, often on pine litter in bare areas

and often around the base of pine trees, but also

on rocks and logs; note: easily confused with

Broom Fork-moss (Dicranum scoparium) but the

fi ne, wispy apex is distinctive, particularly when

dry.

Acrocarpous NS, BAP; Size: medium-sized and

forming open turfs or just scattered stems; branching:

obscured; colour: yellow to mid-green, leaves:

tapering abruptly from a broad base to a fi ne apex,

irregularly undulate and concave giving the shoots a

fat appearance; habitat: most frequent on thin peat

under open, ‘leggy’ heather on moderate to steep

slopes but also occurring in damper heath; note: a

very different ‘habit’ from other Dicranum species and

might be overlooked as the common moss Swans-neck

Thyme-moss (Mnium hornum) when dry.

Acrocarpous NS, BAP; size: medium to robust

and forming dense cushions which can be large;

branching: obscured; colour: yellow to olive-green,

leaves: long, narrowly triangular with a rather short,

toothed apex, regularly and strongly undulate;

habitat: in boggy areas within the woodland where

there has been little disturbance; note: the olive-green

colour and the habit and habitat are useful characters

but this is a diffi cult plant to spot despite its size.

Pleurocarpous; size: robust and usually forming

large patches; branching: regular with branches

branched again, like a small fern; colour: mid to

yellow-green, often ‘chaffy’ when dry; stem: red,

springy; new growth emerging from old frond and

may form several layers of fronds, hence ‘step-

stair moss’; leaves: broadly triangular, no nerve;

habitat: woodland fl oor, ledges, boulder tops.

Pleurocarpous; size: small and forming untidy

patches or straggling stems; branching: irregularly

pinnate with all the branches in the same plane

so the shoots look fl at; colour: pale-green, often

whitish when dry; stem: green; leaves: narrow,

rounded-triangular, no nerve and with a fi ne

pointed tip which is turned down giving the

shoots a ‘plaited’ appearance; habitat: a common

species amongst the heather and blaeberry in

heathy woodland; note: Cypress-leaved Plait-moss

(Hypnum cupressiforme), a very common species

in a variety of habitats, is similar but is usually

greener and has less regular branching.

Green Shield-moss (Buxbaumia viridis)

Waved Silk-moss

(Plagiothecium undulatum)

Rusty Fork-moss (Dicranum spurium)Dusky Fork-moss (Dicranum fuscescens)

Ostrich-plume Feather-moss

(Ptilium crista-castrensis)

Broom Fork-moss (Dicranum scoparium)

Red-stemmed Feather-moss

(Pleurozium schreberi)

Common Haircap (Polytrichum commune)

Bristly Haircap (Polytrichum piliferum)

Wavy Fork-moss (Dicranum undulatum) Glittering Wood-moss

(Hylocomium splendens)

Heath Plait-moss (Hypnum jutlandicum)

Big Shaggy-moss

(Rhytidiadelphus triquetrus)

Little Shaggy-moss

(Rhytidiadelphus loreus)

Red Bog-moss (Sphagnum capillifolium)

Further informationBooksBritish Mosses and liverworts: a fi eld guide. British Bryological Society (2010).

The fi rst comprehensive colour fi eld guide to bryophytes with good keys and hundreds of photos.

Mosses and Liverworts; Gordon Rothero (2005). A brief, general introduction to Scottish bryophytes,

part of the ‘Naturally Scottish’ series published by Scottish Natural Heritage, Battleby.

Bryophytes of native woods – a fi eld guide to common mosses and liverworts of Scotland’s native woods.

Carol L Crawford (2002), Native Woodlands Discussion Group. A small booklet with good colour photos.

Mosses and Liverworts. New Naturalist 97, Porley RD & Hodgetts NG, (2005). Collins.

An accessible account of our bryophyte heritage with a good section on woodlands.

The geographical relationships of British and Irish bryophytes; Hill MO & Preston CD (1998). Journal of Bryology, 20: 127-226.

Information and advicewww.britishbryologicalsociety.org.uk. The British Bryological Society has an excellent

website with useful information on publications, courses, fi eld meetings and lots of pictures.

www.nwdg.org.uk. The Native Woodland Discussion Group runs courses on Atlantic mosses and liverworts.

Advice and SupportPlantlife Scotland can help you in your quest for information and support.

Plantlife Scotland, Balallan House, Allan Park, Stirling, FK8 2QG

Tel: +44 (0) 1786 478509 www.plantlife.org.uk Scotland@plantlife.org.uk

© March 2010 ISBN 978-1-907141-23-2

Plantlife Scotland is part of Plantlife International – the Wild Plant Conservation charity, a charitable company limited by guarantee.

Registered in Scotland (SC038951) and in England and Wales (1059559). Registered company no 3166339.

This guide has been written and illustrated for Plantlife Scotland by Gordon Rothero

All photos © Gordon Rothero, unless otherwise stated. Cover photo © Laurie Campbell.

LIVERWORTS

Key features for identifying liverwortsGrowth form. There are two sorts of liverworts; leafy liverworts have a stem and leaves and resemble a moss, whereas thallose or thalloid liverworts have a simple strap of tissue with no stem or leaves. Leafy liverworts can form erect cushions and turfs while some are creeping and closely apressed to rock or tree. The size of the plant is also important; a number of oceanic liverworts are very, very small.

Leaf shape. This is all-important in leafy liverworts and is much more variable than in mosses. Liverwort leaves can be simple and round, they can be deeply divided into fi laments or into broader lobes, the lobes can be of different sizes and can be bent over or under each other and formed into fl aps or pouches. Many liverworts also have pronounced teeth on the margin of the leaf, visible without a hand lens. Many leafy liverworts also have under-leaves, usually much smaller than the main leaves, and on the lower side of the stem.

Abbreviations

NR – nationally rare; NS – nationally scarce; RDL – Red Data List 2001; S8 – listed on Schedule 8 of the Wildlife and Countryside Act. Oceanic indicates that this is an oceanic species as defi ned by Hill and Preston (1998).

Photoset Left - A variety of leafy liverwort leaves, clockwise

from top left: Taylor’s Flapwort (Mylia taylori), Prickly

Featherwort (Plagiochila spinulosa), Ciliated Fringewort

(Ptilidium ciliare), White earwort (Diplophyllum albicans -

note upper lobe bent over the top of the lower and the line

of longer cells), Common Paw-wort (Barbilophozia fl oerkei),

Bifi d Crestwort (Lophocolea bidentata), Creeping Fingerwort

(Lepidozia reptans), part of stem with ‘fi ngered’ leaves.

Above - Left: a thallose liverwort, Overleaf Pellia (Pellia

epiphylla) with fruits; right: a leafy liverwort with round

leaves, Autumn Flapwort (Jamesoniella autumnalis).

Key features for identifying Mosses

Growth form. Leaving aside the very distinctive bog-

mosses (Sphagnum), mosses can be split into two

groups, acrocarpous and pleurocarpous. There is a

technical difference between these two forms but in

practical terms, acrocarps usually have erect stems

and grow in cushions or turfs while pleurocarps tend

to grow with main stems parallel to the ground (or rock

or tree trunk) and form wefts. The often dense growth

form of acrocarps means that their sparse branches

are obscured while in pleurocarps the branches are

usually many and easily seen.

Branching. For some of the pleurocarps it is useful

to note what the pattern of branching looks like. Is

it regular with branches more or less opposite each

other on the main stem (pinnate) or irregular? Are

the regular branches branched again (bi-pinnate) and

again (tri-pinnate) giving a fern-like structure?

Colour. Colour and texture are all-important fi eld

characters. Many species have a particular shade,

admittedly usually of green, which coupled with

the structure of stem and leaf, gives a texture which

is what the eye picks up from a distance. Another

important colour is that of the main stem; when

the leaves are dry they become opaque and it may

be necessary to scrape away some leaves with your

fi ngernail to see the stem colour.

Leaf shape. Moss leaves have a variety of shapes but

they tend to be variations on the same theme, with

a relatively broad base tapering to a narrower apex.

Some leaves are long and narrow and taper to a fi ne

point, others have a broad triangular shape tapering

shortly to a sharp point. Other mosses have leaves

with a blunt apex and a few species have round leaves.

Another useful character is whether or not the leaves

are all curved in the same direction (falcate) or bent

back from the stem (refl exed or squarrose). Some

leaves also have teeth on the margin, usually visible

only with a hand lens.

Nerve (or costa). A very useful character is whether

the moss leaf has a nerve or not. The nerve (or costa)

is a thickened rib of tissue running up the centre of

the leaf which looks like a dark line if the leaf is held

against the light and viewed with the hand-lens. It

usually extends beyond halfway up the leaf and may

reach the apex.

Above - Left: acrocarpous Scott’s Fork moss (Dicranum

scottianum) ; right: pleurocarpous Larger Mouse-tail

Moss (Isothecium alopecuroides).

Above - Irregular branching in Red-stemmed Feather-

moss (Pleurozium schreberi) on the left and regular

tri-pinnate branching in Glittering Wood-moss

(Hylocomium splendens) on the right; note the red stems.

Photoset above - A variety of moss-leaf shapes, clockwise

from top left: Dotted Thyme-moss (Rhizomnium

punctatum), Catherine’s Moss (Atrichum undulatum),

Little Shaggy-moss (Rhytidiadelphus loreus), Yellow

Fringe-moss (Racomitrium aciculare), Common Striated

Feather-moss (Eurhynchium striatum), Cypress-leaved

Plait-moss (Hypnum cupressiforme).

Moss or liverwort?Initially this is a tricky question but with a little experience it ceases to be a problem. Thallose liverworts are

easy but leafy liverworts can be passed over as mosses by the uninitiated. In most mosses the leaves grow all

around the stem but in most liverworts the main leaves are in two ranks down each side of the stem, sometimes

with a line of smaller under-leaves below. Most moss leaves are roughly triangular, wide at the base and narrow

to the apex; few liverworts are like that. Moss leaves never have lobes whereas many leafy liverworts do. Most

mosses with round leaves have a nerve; no leafy liverworts have a nerve but a few have lines of cells running up

the centre of the leaf.

MOSSES

Oceanic, NS. Acrocarpous moss; size: large and usually

forming loose cushions within the heath; branching:

obscured; colour: usually a shiny, dark green; stem: often

reddish; leaves: very narrowly triangular, tapering to a very

long, fi ne point which is formed mostly from the broad

nerve; habitat: in lower stands of the heath, especially

where rocky or below crags; note: two very similar species

Beaked Bow-moss (Dicranodontium denudatum) and

Orange Bow-moss (Dicranodontium asperulum) also occur

in oceanic heath.

Bog-moss; size: medium sized and forming swelling

cushions or hummocks of erect stems but in the oceanic

heath usually as a loose turf under the heather; branching:

a bunch of tight branches at the top of the stem (capitulum)

and whorls of branches at intervals below; colour: often

red but in the oceanic heath more frequently green with

varying amounts of red; leaves: branch leaves are narrowly

triangular; habitat: an abundant and locally dominant

plant in hilly areas in both mire and heath; note: other

species of Sphagnum occur in this community but this is

the most frequent species.

Acrocarpous moss; size: large and usually forming

sprawling patches; branching: unlike most acrocarpous

mosses, there are frequent irregular branches; colour:

usually grey when dry and blackish-green when wet;

stem: green; leaves: narrowly triangular, tapering to a long,

toothed white ‘hair-point’; habitat: an abundant and locally

dominant plant in hilly areas, on rocks and in mire and

heath; note: an important and unmistakeable component

of our upland vegetation and a constant in oceanic heath.

Oceanic, NR. Bog-moss; size: large, twice the size of

Red Bog-moss (Sphagnum capillifolium), and forming

loose patches in the turf; branching: a bunch of long

branches at the top of the stem (capitulum) giving an

untidy appearance and whorls of long branches below;

colour: often red or pink but with some variegated green ;

leaves: branch leaves are narrowly triangular ; habitat: in

grassy heath and often associated with Juniper Prongwort

(Herbertus aduncus ssp. hutchinsiae), and Carrington’s

Featherwort (Plagiochila carringtonii); note: the size,

colour, long branches and habitat are useful characters;

outside of the west of Scotland only known from one site in

Wales and one in Ireland.

Curve-leaved Bow-moss (Dicranodontium uncinatum) Woolly Fringe-moss (Racomitrium lanuginosum)

Red Bog-moss (Sphagnum capillifolium) Skye Bog-moss (Sphagnum skyense)

MOSSES

Bryophytes of

Scotland’s oceanic heath

British Lichen Society

Introduction

This Plantlife fi eld guide will help those who want to

go a little further in identifying the carpets of mosses

and liverworts that make our western hills such

special places. This guide deals with species making

up Scotland’s oceanic heath, a globally rare plant

community.

What are mosses and liverworts?

Mosses and liverworts (collectively known as

bryophytes) are two of the oldest groups of land plants

and have had millions of years to evolve a variety of

species that have colonised almost all habitats apart

from the sea. Most have a simple structure with a

main stem and more or less frequent branches covered

in leaves. They do not have roots but absorb water

and minerals directly into the (usually) single layer of

cells in the leaves. Most bryophytes are some shade

of green but the liverworts in Scotland’s oceanic heath

are usually strongly coloured, ranging from yellow and

orange, through to reds and purples, to almost black.

What is Scotland’s oceanic heath?

Oceanic heath is a neglected habitat of global

importance. Heather and blaeberry are major

components but what makes oceanic heath special

is its ground layer of mosses that only thrive in the

oceanic climate of the hills in the west of Scotland.

This ground layer is also home to a diverse fl ora of

large leafy liverworts that have a highly restricted

global distribution. These rare liverwort species

normally occur alongside more common species, such

as Woolly Fringe-moss (Racomitrium lanuginosum),

Red Bog-moss (Sphagnum capillifolium) and

more widespread liverworts like Taylor’s Flapwort

(Mylia taylori) and Orkney Notchwort (Anastrepta

orcadensis).

This habitat is also known as ‘Scottish liverwort heath’,

the ‘Northern hepatic mat’ or the ‘oceanic-montane

heath’.

Why is Scotland’s oceanic heath important?

Oceanic heath is globally rare. In Europe, it is only

well-developed in Western Scotland, western Ireland

and, to a much more limited extent, in SW Norway.

Globally, some of the same species occur in a similar

community above the tree line in the Sino-Himalaya

and North-west America.

Most of the constituent liverwort species have a

remarkably disjunct global distribution. For example

Cloud Earwort (Scapania nimbosa), also occurs in W

Ireland (rare), SW Norway (one site) Nepal, Sikkim and

Yunnan. The community is also so limited in its UK

distribution that most species are nationally scarce

and some are nationally rare. Northern Prongwort

(Herbertus borealis) is limited to just one site in

Europe on Beinn Eighe and Lindenberg’s Featherwort

(Adelanthus lindenbergianus) has one site on Islay

(elsewhere in Europe only in W Ireland). We are still

fi nding out more about this remarkable community,

with molecular work revealing two new species in the

past few years.

Where to look for the oceanic heath

This habitat has a patchy distribution from Islay in

the south, up the west coast to Ben Hope, including

hills in the Hebrides with more limited outposts on

Orkney and Shetland. There are also isolated stands

further east on the larger hills extending to the main

Cairngorms. Most sites are on N or NE-facing slopes

from 200m up to 900m where rain falls in excess of

200 days in a year.

The majority of Scotland’s oceanic heath occurs on

moderate to steep slopes with an aspect from NW

through to NE, almost always where there are some

exposed rocks and frequently where the heath has

developed over large block scree. A number of the

lower level sites are quite grassy, with only residual

amounts of heather but such sites probably represent

relicts that were formerly dominated by ericaceous

shrubs and altered by subsequent management.

They usually have frequent Juniper Prongwort

(Herbertus aduncus ssp. hutchinsiae) but only patchy

stands of other species. The higher sites, where

blaeberry is dominant and where Donn’s Notchwort

(Anastrophyllum donnianum) and Alpine Notchwort

(Anastrophyllum alpinum) can occur, are subject

to long periods of snow-lie which may give some

protection from frost.

‘Bryologising’

Although identifying species initially needs a

modicum of determination, it is possible to quite

quickly become familiar with most of the indicator

species for Scotland’s oceanic heath. The lower

stands of this community have the virtue of being

available all the year round and grow in really nice

places. Though some of the species are very distinct,

even from some metres away, the process and the

enjoyment will be enhanced if you get used to using

a hand-lens (at least x10) to reveal the fi ner features

on which identifi cation sometimes depends. In the

descriptions, where a lens is necessary it is indicated

by (lens). Remember, hand-lens to the eye and move

the plant into focus.

In these guides, English names have been used

alongside the Latin; these are not ‘common names’

in the same sense as those for fl owers because they

are all recent inventions and as yet are hardly used

except in publications like this! Latin names should

always be used for recording purposes to avoid any

ambiguity.

LIVERWORTS

Oceanic, NS. Leafy liverwort; size: medium-

sized but often forming large cushions or

patches; branching: irregular but frequent,

the branches tending to narrow towards the

end colour: usually some shade of yellow

but may be variegated green or brownish;

leaves: two lobes, the upper larger than the

lower but the most noticeable feature is the

margin which has lots of long narrow teeth

(cilia) easily visible with a lens underleaves:

similar in shape to the main leaves but much

smaller; habitat: on thin peaty soil amongst

ericaceous shrubs, in block scree or at the

base of crags on N and NE-facing slopes,

also occasionally in heathy woodland and

wooded ravines; note: confusion only possible

with Ciliated Fringewort (Ptilidium ciliare),

a species of drier heaths which has shorter,

blunt branches.

Leafy liverwort; size: medium to large,

usually in large swelling cushions; colour:

variable, in unshaded places it often purplish-

red variegated with yellow and brown, usually

green in shade; leaves: large and round, often

pressed together at the apex and with very

large cells (lens); habitat: on thin peaty soil

amongst ericaceous shrubs, in block scree

or at the base of crags on N and NE-facing

slopes, also frequent in heathy woodland and

wooded ravines; note: a handsome liverwort,

generally common in the west and also

frequent in this community.

Oceanic, NS. Leafy liverwort; size: medium-

sized but often forming large cushions or

patches with erect stems; colour: pale or

yellow-green, often whitish when dry; leaves:

round and placed vertically on the stem so

that the upper faces are pressed against

each other so the shoots look fl attened, the

upper margin running down the stem where it

joins (decurrent); habitat: on thin peaty soil

amongst ericaceous shrubs, in block scree

or at the base of crags on N and NE-facing

slopes, also occasionally in heathy woodland

and wooded ravines.

Oceanic. Leafy liverwort; size: medium-

sized but often forming large cushions or

extensive pure patches; colour: yellow- or

mid-green; leaves: rounded but upper

margin rather straight and running down

the stem (decurrent) in a slight curve, apex

and lower margin with spine-like teeth,

aromatic when crushed and inrolled when

dry; habitat: on rocks and trees in shaded

habitats in the west and common in the

oceanic heath community.

Oceanic. Leafy liverwort; size: medium

to large forming loose turfs or straggling

through other bryophytes; colour: usually

a distinctive dark reddish-purple but can be

yellow- or brownish-green in shade; leaves:

complex, bilobed with a smaller, almost

tubular, upper lobe over a large concave

lower lobe; habitat: on thin peaty soil

amongst ericaceous shrubs, in block scree

or at the base of crags on N and NE-facing

slopes, in wet heath, on blanket bog, also in

heathy woodland and wooded ravines; Note:

Unmistakeable, frequent in this community

but also abundant in wet heath and mires.

Oceanic. Leafy liverwort; size: medium

sized but often forming large cushions or

extensive pure patches; colour: a rather

dull yellow- or brownish-green; leaves: two

lobed but with the upper lobe bent over the

lower and upper part bent back so it stands

up from the stem, margin usually strongly

toothed (lens); habitat: on thin peaty soil

amongst ericaceous shrubs, in block scree,

on acidic rocks and trees; note: a common

western species often present and locally

abundant in stands of Scotland’s oceanic

heath at lower levels.

Oceanic, NS. Leafy liverwort; size: medium

sized, usually in small patches mixed with

other bryophytes and only rarely forming

large pure patches; colour: usually a purplish-

red, rarely reddish-green or orange; leaves:

two lobed and the lobes divided almost to

the base, the smaller upper lobe bent fl at

over the lower and both lobes tapering to a

rounded apex, margin with long curved teeth

(lens); habitat: on thin peaty soil amongst

ericaceous shrubs, in block scree or at the

base of crags on N and NE-facing slopes; note:

easily distinguished from Bird’s-foot Earwort

(Scapania ornithopodioides), with which it

usually grows, by the colour and long, curved

teeth on the leaves.

Oceanic, NR, RDL, S8. Leafy liverwort;

size: medium-sized, forming open turfs of

neat, erect stems amongst other bryophytes;

colour: usually dark brown but can be

brownish-green, glossy; leaves: rounded and

the upper margin incurved towards the stem,

all leaves turned down in the same direction;

habitat: on thin peaty soil amongst heather

on NE-facing slopes; note: superfi cially

similar to Anastrophyllum donnianum and

Anastrepta orcadensis but the incurved

upper leaf margin is unique; currently known

from just one site on Islay.

Leafy liverwort; size: medium, usually

growing as an open turf of erect stems

amongst other bryophytes; colour: often a

rather pale green but can be a dull yellow or

brown, frequently has red gemmae at the

stem apex; leaves: rounded but irregular

and usually shallowly lobed at the apex, the

base of the leaf is concave but the upper part

curves away from the stem so that the whole

looks convex; habitat: on thin peaty soil

amongst ericaceous shrubs, in block scree

or at the base of crags on N and NE-facing

slopes, also frequent in heathy woodland and

wooded ravines; note: generally common in

the west and also frequent in this community.

Oceanic, NS. Leafy liverwort; size:

medium-sized, forming open turfs of erect

stems amongst other bryophytes; colour:

usually dark brown but can be brownish

green, glossy; leaves: rounded-triangular

but concave so that leaves seem narrow, the

apex has a small notch, all leaves turned

down in the same direction; habitat: on

thin peaty soil amongst ericaceous shrubs,

in block scree or at the base of crags on N

and NE-facing slopes; note: the commonest

species of this community at higher

altitudes and locally abundant.

Oceanic, NS. Leafy liverwort; size: medium-

sized, forming open turfs of erect stems

amongst other bryophytes; colour: usually

dark brown but can be brownish green, glossy;

leaves: rounded and very concave, forming

a cup-shape, the apex has a small notch, all

leaves turned down in the same direction;

habitat: on thin peaty soil amongst ericaceous

shrubs, in block scree or at the base of crags on

N and NE-facing slopes; note: almost always

with Donn’s Notchwort (Anastrophyllum

donnianum) but much less common, easily

confused with it but the cup-shaped leaves

are distinctive – if you are not sure, then it is

not Anastrophyllum alpinum. Joergensen’s

Notchwort (Anastrophyllum joergensenii) has

only recently been described as distinct from

Anastrophyllum alpinum and is diffi cult to

separate from it; it is smaller, usually forms

denser turfs and may occur in damper sites.

Leafy liverwort; size: small but may form large

cushions or straggling patches; branching:

frequent thin branches from bottom of stem

(fl agellae) pale with tiny leaves colour: usually

mid-green but can be yellowish; leaves:

rounded-triangular and asymmetric with three

small teeth at the narrow apex, often closely

set and overlapping but sometimes widely

spaced; underleaves: rounded and held close

the stem, usually with teeth visible at the apex

(lens); habitat: on thin peaty soil amongst

ericaceous shrubs, in block scree or at the base

of crags on N and NE-facing slopes; note: This

is a widespread species not limited to this

community but frequent in it.

Oceanic, NS. Differs from Bazzania

tricrenata in usually forming denser and

more ‘succulent-looking’ patches, the base

of the leaves widely crossing the stem when

viewed from above; underleaves held away

from the stem and untoothed.

Oceanic. Leafy liverwort; size: medium-sized

with slender stems but often forming deep,

dense cushions or large straggling patches;

branching: frequent thin branches from

bottom of stem (fl agellae) with tiny leaves

colour: usually a distinctive reddish- orange

colour but can be green or yellow green tinged

with red; leaves: triangular but deeply divided

into two long, pointed lobes, usually all turned

in the same direction underleaves: similar in

shape and size to the main leaves; habitat:

on thin peaty soil amongst ericaceous shrubs,

in block scree or at the base of crags on N and

NE-facing slopes, also in heathy woodland

and wooded ravines; note: confusion is only

possible with the next species or with Straw

Prongwort (Herbertus stramineus), another

oceanic species but one which favours more

base-rich sites.

Oceanic, NR, RDL. Distinguished from

Herbertus aduncus ssp. hutchinsiae

by its usually bright orange colour, the

more regularly curved and asymmetric

leaves, smaller underleaves and by the

numerous fl agelliferous branches; note:

currently known only from Beinn Eighe,

its only European site.

Oceanic, NS. Leafy liverwort; size:

medium to large, usually in small patches

mixed with other bryophytes and only

rarely forming large pure patches; colour:

usually a characteristic ‘liver’ red colour;

leaves: two lobed and the lobes divided to

the base, the smaller upper lobe bent fl at

over the lower and both lobes tapering to a

rounded apex, margin with straight teeth

(lens); habitat: on thin peaty soil amongst

ericaceous shrubs, in block scree or at the

base of crags on N and NE-facing slopes,

also occasionally in heathy woodland and

wooded ravines.

Lindenberg’s Featherwort

(Adelanthus lindenbergianus)

Wood’s Whipwort

(Mastigophora woodsii)

Alpine Notchwort

(Anastrophyllum alpinum)

Donn’s Notchwort

(Anastrophyllum donnianum)

Prickly Featherwort

(Plagiochila spinulosa)

Orkney Notchwort

(Anastrepta orcadensis)

Taylor’s Flapwort (Mylia taylori)Carrington’s Featherwort

(Plagiochila carringtonii)

Lesser Whipwort

(Bazzania tricrenata)

Arch-leaved Whipwort

(Bazzania pearsonii)

Juniper Prongwort

(Herbertus aduncus ssp. hutchinsiae)

Northern Prongwort

(Herbertus borealis)

Purple Spoonwort

(Pleurozia purpurea)

Western Earwort

(Scapania gracilis)

Cloud Earwort

(Scapania nimbosa)

Bird’s-foot Earwort

(Scapania ornithopodioides)

© M

are

n F

lag

me

ier