LIVERWORTS MOSSES Curve-leaved Bow- (Dicranodontium uncinatum) Woolly Fringe-moss (Racomitrium lanuginosum) Key features for identifying Key features for identifying liverworts Mosses Growth form. There are two sorts of liverworts; leafy Growth form. Leaving aside the very distinctive bog- liverworts have a stem and leaves and resemble a mosses (Sphagnum), mosses can be split into two moss, whereas thallose or thalloid liverworts have a groups, acrocarpous and pleurocarpous. There is a simple strap of tissue with no stem or leaves. Leafy technical difference between these two forms but in liverworts can form erect cushions and turfs while some practical terms, acrocarps usually have erect stems are creeping and closely apressed to rock or tree. The and grow in cushions or turfs while pleurocarps tend size of the is also important; a number of oceanic to grow with main stems parallel to the ground (or rock liverworts are very, very small. Above - Left: a thallose liverwort, Overleaf Pellia (Pellia Above - Left: acrocarpous Scott’s Fork moss (Dicranum or tree trunk) and form wefts. The often dense growth epiphylla) with fruits; right: a leafy liverwort with round Leaf shape. This is all-important in leafy liverworts and scottianum) ; right: pleurocarpous Larger Mouse-tail form of acrocarps means that their sparse branches leaves, Autumn Flapwort (Jamesoniella autumnalis). is much more variable than in mosses. Liverwort leaves Moss (Isothecium alopecuroides). are obscured while in pleurocarps the branches are can be simple and round, they can be deeply divided into filaments or into broader lobes, the lobes can be usually many and easily seen. of different sizes and can be bent over or under each Branching. For some of the pleurocarps it is useful other and formed into flaps or pouches. Many liverworts to note what the pattern of branching looks like. Is also have pronounced teeth on the margin of the leaf, Acrocarpous moss; size: large and usually forming it regular with branches more or less opposite each Oceanic, NS. Acrocarpous moss; size: large and usually visible without a hand lens. Many leafy liverworts also sprawling patches; branching: unlike most acrocarpous other on the main stem (pinnate) or irregular? Are forming loose cushions within the heath; branching: have under-leaves, usually much smaller than the main mosses, there are frequent irregular branches; colour: the regular branches branched again (bi-pinnate) and obscured; colour: usually a shiny, dark green; stem: often leaves, and on the lower side of the stem. reddish; leaves: very narrowly triangular, tapering to a very usually grey when dry and blackish-green when wet; again (tri-pinnate) giving a fern-like structure? stem: green; leaves: narrowly triangular, tapering to a long, Photoset Left - A variety of leafy liverwort leaves, clockwise long, fine point which is formed mostly from the broad toothed white ‘hair-point’; habitat: an abundant and locally from top left: Taylor’s Flapwort (Mylia taylori), Prickly Colour. Colour and texture are all-important field nerve; habitat: in lower stands of the heath, especially dominant plant in hilly areas, on rocks and in mire and Featherwort (Plagiochila spinulosa), Ciliated Fringewort characters. Many species have a particular shade, where rocky or below crags; note: two very similar species Above - Irregular branching in Red-stemmed Feather- heath; note: an important and unmistakeable component (Ptilidium ciliare), White earwort (Diplophyllum albicans - admittedly usually of green, which coupled with Beaked Bow-moss (Dicranodontium denudatum) and moss (Pleurozium schreberi) on the left and regular of our upland vegetation and a constant in oceanic heath. note upper lobe bent over the top of the lower and the line tri-pinnate branching in Glittering Wood-moss the structure of stem and leaf, gives a texture which Orange Bow-moss (Dicranodontium asperulum) also occur of longer cells), Common Paw-wort (Barbilophozia floerkei), (Hylocomium splendens) on the right; note the red stems. is what the eye picks up from a distance. Another in oceanic heath. Bifid Crestwort (Lophocolea bidentata), Creeping Fingerwort important colour is that of the main stem; when (Lepidozia reptans), part of stem with ‘fingered’ leaves. the leaves are dry they become opaque and it may be necessary to scrape away some leaves with your fingernail to see the stem colour. Red Bog-moss (Sphagnum capillifolium) Skye Bog-moss (Sphagnum skyense) Abbreviations Leaf shape. Moss leaves have a variety of shapes but NR – nationally rare; NS – nationally scarce; RDL – Red Data List 2001; S8 – listed on Schedule 8 of they tend to be variations on the same theme, with the Wildlife and Countryside Act. Oceanic indicates that this is an oceanic species as defined by Hill a relatively broad base tapering to a narrower apex. Some leaves are long and narrow and taper to a fine and Preston (1998). 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. Further information Another useful character is whether or not the leaves Books are all curved in the same direction (falcate) or bent British Mosses and liverworts: a field guide. British Bryological Society (2010). back from the stem (reflexed or squarrose). Some The first comprehensive colour field guide to with good keys and hundreds of photos. leaves also have teeth on the margin, usually visible Mosses and Liverworts; Gordon Rothero (2005). A brief, general introduction to Scottish bryophytes, only with a hand lens. part of the ‘Naturally Scottish’ series published by Scottish Natural Heritage, Battleby. Photoset above - A variety of moss-leaf shapes, clockwise Nerve (or costa). A very useful character is whether Bryophytes of native woods – a field guide to common mosses and liverworts of ’s native woods. from top left: Dotted Thyme-moss (Rhizomnium Carol L Crawford (2002), Native Woodlands Discussion Group. A small booklet with good colour photos. the moss leaf has a nerve or not. The nerve (or costa) punctatum), Catherine’s Moss (Atrichum undulatum), Mosses and Liverworts. New Naturalist 97, Porley RD & Hodgetts NG, (2005). Collins. is a thickened rib of tissue running up the centre of Little Shaggy-moss (Rhytidiadelphus loreus), Yellow An accessible account of our heritage with a good section on woodlands. the leaf which looks like a dark line if the leaf is held Fringe-moss (Racomitrium aciculare), Common Striated The geographical relationships of British and Irish bryophytes; Hill MO & Preston CD (1998). Journal of Bryology, 20: 127-226. against the light and viewed with the hand-lens. It Feather-moss (Eurhynchium striatum), Cypress-leaved usually extends beyond halfway up the leaf and may Bog-moss; size: medium sized and forming swelling Oceanic, NR. Bog-moss; size: large, twice the size of Bryophytes of Information and advice Plait-moss (Hypnum cupressiforme). reach the apex. www.britishbryologicalsociety.org.uk. The British Bryological Society has an excellent cushions or hummocks of erect stems but in the oceanic Red Bog-moss (Sphagnum capillifolium), and forming website with useful information on publications, courses, field meetings and lots of pictures. heath usually as a loose turf under the heather; branching: loose patches in the turf; branching: a bunch of long Scotland’s oceanic heath www.nwdg.org.uk. The Native Woodland Discussion Group runs courses on Atlantic mosses and liverworts. a bunch of tight branches at the top of the stem (capitulum) branches at the top of the stem (capitulum) giving an Moss or liverwort? and whorls of branches at intervals below; colour: often untidy appearance and whorls of long branches below; Advice and Support red but in the oceanic heath more frequently green with colour: often red or pink but with some variegated green ; Plantlife Scotland can help you in your quest for information and support. Initially this is a tricky question but with a little experience it ceases to be a problem. Thallose liverworts are varying amounts of red; leaves: branch leaves are narrowly leaves: branch leaves are narrowly triangular ; habitat: in Plantlife Scotland, Balallan House, Allan Park, Stirling, FK8 2QG easy but leafy liverworts can be passed over as mosses by the uninitiated. In most mosses the leaves grow all triangular; habitat: an abundant and locally dominant grassy heath and often associated with Juniper Prongwort Tel: +44 (0) 1786 478509 www.plantlife.org.uk [email protected] around the stem but in most liverworts the main leaves are in two ranks down each side of the stem, sometimes plant in hilly areas in both mire and heath; note: other ( aduncus ssp. hutchinsiae), and Carrington’s © March 2010 ISBN 978-1-907141-23-2 with a line of smaller under-leaves below. Most moss leaves are roughly triangular, wide at the base and narrow Plantlife Scotland is part of Plantlife International – the Wild Plant Conservation charity, a charitable company limited by guarantee. species of Sphagnum occur in this community but this is Featherwort (Plagiochila carringtonii); note: the size, Registered in Scotland (SC038951) and in England and Wales (1059559). Registered company no 3166339. to the apex; few liverworts are like that. Moss leaves never have lobes whereas many leafy liverworts do. Most the most frequent species. colour, long branches and habitat are useful characters; mosses with round leaves have a nerve; no leafy liverworts have a nerve but a few have lines of cells running up outside of the west of Scotland only known from one site in This guide has been written and illustrated for Plantlife Scotland by Gordon Rothero the centre of the leaf. Wales and one in Ireland. British Society All photos © Gordon Rothero, unless otherwise stated. Cover photo © Laurie Campbell. LIVERWORTS Where to look for the oceanic heath Introduction Lindenberg’s Featherwort Notchwort Donn’s Notchwort Alpine Notchwort Lesser Whipwort Arch-leaved Whipwort Juniper Prongwort Northern Prongwort This habitat has a patchy distribution from Islay in (Adelanthus lindenbergianus) This Plantlife field guide will help those who want to ( orcadensis) (Anastrophyllum donnianum) (Anastrophyllum alpinum) (Bazzania tricrenata) (Bazzania pearsonii) (Herbertus aduncus ssp. hutchinsiae) (Herbertus borealis) the south, up the west coast to Ben Hope, including go a little further in identifying the carpets of mosses hills in the Hebrides with more limited outposts on and liverworts that make our western hills such Orkney and . There are also isolated stands special places. This guide deals with species making further east on the larger hills extending to the main up Scotland’s oceanic heath, a globally rare plant Cairngorms. Most sites are on N or NE-facing slopes community. from 200m up to 900m where rain falls in excess of What are mosses and liverworts? 200 days in a year. Mosses and liverworts (collectively known as The majority of Scotland’s oceanic heath occurs on bryophytes) are two of the oldest groups of land moderate to steep slopes with an aspect from NW and have had millions of years to evolve a variety of through to NE, almost always where there are some species that have colonised almost all habitats apart exposed rocks and frequently where the heath has from the sea. Most have a simple structure with a Oceanic, NS. Leafy liverwort; size: medium- developed over large block scree. A number of the sized, forming open turfs of erect stems Oceanic. Leafy liverwort; size: medium-sized main stem and more or less frequent branches covered lower level sites are quite grassy, with only residual amongst other bryophytes; colour: usually with slender stems but often forming deep, in leaves. They do not have roots but absorb water amounts of heather but such sites probably represent dark brown but can be brownish green, glossy; dense cushions or large straggling patches; and minerals directly into the (usually) single layer of relicts that were formerly dominated by ericaceous Flagmeier © Maren leaves: rounded and very concave, forming branching: frequent thin branches from cells in the leaves. Most bryophytes are some shade Leafy liverwort; size: medium, usually shrubs and altered by subsequent management. a cup-shape, the apex has a small notch, all Leafy liverwort; size: small but may form large bottom of stem (flagellae) with tiny leaves of green but the liverworts in Scotland’s oceanic heath growing as an open turf of erect stems Oceanic, NS. Differs from Bazzania Oceanic, NR, RDL. Distinguished from They usually have frequent Juniper Prongwort Oceanic, NR, RDL, S8. Leafy liverwort; Oceanic, NS. Leafy liverwort; size: leaves turned down in the same direction; cushions or straggling patches; branching: colour: usually a distinctive reddish- orange amongst other bryophytes; colour: often a habitat: on thin peaty soil amongst ericaceous tricrenata in usually forming denser and colour but can be green or yellow green tinged Herbertus aduncus ssp. hutchinsiae are usually strongly coloured, ranging from yellow and (Herbertus aduncus ssp. hutchinsiae) but only patchy size: medium-sized, forming open turfs of medium-sized, forming open turfs of erect frequent thin branches from bottom of stem rather pale green but can be a dull yellow or shrubs, in block scree or at the base of crags on more ‘succulent-looking’ patches, the base with red; leaves: triangular but deeply divided by its usually bright orange colour, the orange, through to reds and purples, to almost black. stands of other species. The higher sites, where neat, erect stems amongst other bryophytes; stems amongst other bryophytes; colour: (flagellae) pale with tiny leavescolour: usually brown, frequently has red gemmae at the N and NE-facing slopes; note: almost always into two long, pointed lobes, usually all turned more regularly curved and asymmetric blaeberry is dominant and where Donn’s Notchwort colour: usually dark brown but can be usually dark brown but can be brownish mid-green but can be yellowish; leaves: of the leaves widely crossing the stem when What is Scotland’s oceanic heath? stem apex; leaves: rounded but irregular with Donn’s Notchwort (Anastrophyllum in the same direction underleaves: similar in leaves, smaller underleaves and by the (Anastrophyllum donnianum) and Alpine Notchwort brownish-green, glossy; leaves: rounded and green, glossy; leaves: rounded-triangular rounded-triangular and asymmetric with three viewed from above; underleaves held away and usually shallowly lobed at the apex, the donnianum) but much less common, easily shape and size to the main leaves; habitat: Oceanic heath is a neglected habitat of global (Anastrophyllum alpinum) can occur, are subject but concave so that leaves seem narrow, the small teeth at the narrow apex, often closely from the stem and untoothed. numerous flagelliferous branches; note: the upper margin incurved towards the stem, base of the leaf is concave but the upper part confused with it but the cup-shaped leaves on thin peaty soil amongst ericaceous shrubs, importance. Heather and blaeberry are major to long periods of snow-lie which may give some all leaves turned down in the same direction; apex has a small notch, all leaves turned set and overlapping but sometimes widely currently known only from Beinn Eighe, components but what makes oceanic heath special curves away from the stem so that the whole are distinctive – if you are not sure, then it is spaced; underleaves: rounded and held close in block scree or at the base of crags on N and its only European site. protection from frost. habitat: on thin peaty soil amongst heather looks convex; habitat: on thin peaty soil down in the same direction; habitat: on is its ground layer of mosses that only thrive in the not Anastrophyllum alpinum. Joergensen’s the stem, usually with teeth visible at the apex NE-facing slopes, also in heathy woodland on NE-facing slopes; note: superficially amongst ericaceous shrubs, in block scree thin peaty soil amongst ericaceous shrubs, Notchwort (Anastrophyllum joergensenii) has and wooded ravines; note: confusion is only oceanic climate of the hills in the west of Scotland. ‘Bryologising’ (lens); habitat: on thin peaty soil amongst similar to Anastrophyllum donnianum and or at the base of crags on N and NE-facing in block scree or at the base of crags on N only recently been described as distinct from ericaceous shrubs, in block scree or at the base possible with the next species or with Straw This ground layer is also home to a diverse flora of Although identifying species initially needs a Anastrepta orcadensis but the incurved slopes, also frequent in heathy woodland and and NE-facing slopes; note: the commonest Anastrophyllum alpinum and is difficult to of crags on N and NE-facing slopes; note: This Prongwort (Herbertus stramineus), another large leafy liverworts that have a highly restricted modicum of determination, it is possible to quite upper leaf margin is unique; currently known wooded ravines; note: generally common in species of this community at higher separate from it; it is smaller, usually forms is a widespread species not limited to this oceanic species but one which favours more global distribution. These rare liverwort species quickly become familiar with most of the indicator from just one site on Islay. the west and also frequent in this community. altitudes and locally abundant. denser turfs and may occur in damper sites. community but frequent in it. base-rich sites. normally occur alongside more common species, such species for Scotland’s oceanic heath. The lower as Woolly Fringe-moss (Racomitrium lanuginosum), stands of this community have the virtue of being Wood’s Whipwort Taylor’s Flapwort (Mylia taylori) Carrington’s Featherwort Prickly Featherwort Purple Spoonwort Western Earwort Cloud Earwort ’s-foot Earwort Red Bog-moss (Sphagnum capillifolium) and available all the year round and grow in really nice (Mastigophora woodsii) (Plagiochila carringtonii) (Plagiochila spinulosa) (Pleurozia purpurea) (Scapania gracilis) (Scapania nimbosa) (Scapania ornithopodioides) more widespread liverworts like Taylor’s Flapwort places. Though some of the species are very distinct, (Mylia taylori) and Orkney Notchwort (Anastrepta even from some metres away, the process and the orcadensis). enjoyment will be enhanced if you get used to using This habitat is also known as ‘Scottish liverwort heath’, a hand-lens (at least x10) to reveal the finer features the ‘Northern hepatic mat’ or the ‘oceanic-montane on which identification sometimes depends. In the heath’. descriptions, where a lens is necessary it is indicated by (lens). Remember, hand-lens to the eye and move Why is Scotland’s oceanic heath important? the plant into focus. Oceanic heath is globally rare. In , it is only In these guides, English names have been used well-developed in Western Scotland, western Ireland alongside the Latin; these are not ‘common names’ and, to a much more limited extent, in SW . in the same sense as those for flowers because they Globally, some of the same species occur in a similar are all recent inventions and as yet are hardly used Oceanic, NS. Leafy liverwort; size: medium- community above the tree line in the Sino-Himalaya except in publications like this! Latin names should and North-west America. sized but often forming large cushions or always be used for recording purposes to avoid any patches; branching: irregular but frequent, Oceanic, NS. Leafy liverwort; size: medium Most of the constituent liverwort species have a ambiguity. the branches tending to narrow towards the sized, usually in small patches mixed with Oceanic, NS. Leafy liverwort; size: remarkably disjunct global distribution. For example end colour: usually some shade of yellow Oceanic. Leafy liverwort; size: medium other bryophytes and only rarely forming Leafy liverwort; size: medium to large, Oceanic, NS. Leafy liverwort; size: medium- Oceanic. Leafy liverwort; size: medium- Oceanic. Leafy liverwort; size: medium medium to large, usually in small patches Cloud Earwort (Scapania nimbosa), also occurs in W but may be variegated green or brownish; to large forming loose turfs or straggling large pure patches; colour: usually a purplish- usually in large swelling cushions; colour: sized but often forming large cushions or sized but often forming large cushions or sized but often forming large cushions or mixed with other bryophytes and only Ireland (rare), SW Norway (one site) Nepal, Sikkim and leaves: two lobes, the upper larger than the variable, in unshaded places it often purplish- patches with erect stems; colour: pale or extensive pure patches; colour: yellow- or through other bryophytes; colour: usually extensive pure patches; colour: a rather red, rarely reddish-green or orange; leaves: lower but the most noticeable feature is the rarely forming large pure patches; colour: Yunnan. The community is also so limited in its UK red variegated with yellow and brown, usually yellow-green, often whitish when dry; leaves: mid-green; leaves: rounded but upper a distinctive dark reddish-purple but can be dull yellow- or brownish-green; leaves: two two lobed and the lobes divided almost to margin which has lots of long narrow teeth the base, the smaller upper lobe bent flat usually a characteristic ‘liver’ red colour; distribution that most species are nationally scarce green in shade; leaves: large and round, often round and placed vertically on the stem so margin rather straight and running down yellow- or brownish-green in shade; leaves: lobed but with the upper lobe bent over the (cilia) easily visible with a lens underleaves: over the lower and both lobes tapering to a leaves: two lobed and the lobes divided to and some are nationally rare. Northern Prongwort pressed together at the apex and with very that the upper faces are pressed against complex, bilobed with a smaller, almost similar in shape to the main leaves but much the stem (decurrent) in a slight curve, apex lower and upper part bent back so it stands rounded apex, margin with long curved teeth the base, the smaller upper lobe bent flat (Herbertus borealis) is limited to just one site in large cells (lens); habitat: on thin peaty soil tubular, upper lobe over a large concave smaller; habitat: on thin peaty soil amongst each other so the shoots look flattened, the and lower margin with spine-like teeth, up from the stem, margin usually strongly (lens); habitat: on thin peaty soil amongst over the lower and both lobes tapering to a amongst ericaceous shrubs, in block scree lower lobe; habitat: on thin peaty soil Europe on Beinn Eighe and Lindenberg’s Featherwort ericaceous shrubs, in block scree or at the upper margin running down the stem where it aromatic when crushed and inrolled when toothed (lens); habitat: on thin peaty soil ericaceous shrubs, in block scree or at the rounded apex, margin with straight teeth or at the base of crags on N and NE-facing amongst ericaceous shrubs, in block scree (Adelanthus lindenbergianus) has one site on Islay base of crags on N and NE-facing slopes, joins (decurrent); habitat: on thin peaty soil dry; habitat: on rocks and trees in shaded or at the base of crags on N and NE-facing amongst ericaceous shrubs, in block scree, base of crags on N and NE-facing slopes; note: (lens); habitat: on thin peaty soil amongst slopes, also frequent in heathy woodland and (elsewhere in Europe only in W Ireland). We are still also occasionally in heathy woodland and amongst ericaceous shrubs, in block scree habitats in the west and common in the slopes, in wet heath, on blanket bog, also in on acidic rocks and trees; note: a common easily distinguished from Bird’s-foot Earwort ericaceous shrubs, in block scree or at the wooded ravines; note: a handsome liverwort, finding out more about this remarkable community, wooded ravines; note: confusion only possible or at the base of crags on N and NE-facing oceanic heath community. heathy woodland and wooded ravines; Note: western species often present and locally (Scapania ornithopodioides), with which it base of crags on N and NE-facing slopes, with molecular work revealing two new species in the with Ciliated Fringewort (Ptilidium ciliare), generally common in the west and also slopes, also occasionally in heathy woodland Unmistakeable, frequent in this community abundant in stands of Scotland’s oceanic usually grows, by the colour and long, curved also occasionally in heathy woodland and past few years. a species of drier heaths which has shorter, frequent in this community. and wooded ravines. but also abundant in wet heath and mires. heath at lower levels. teeth on the leaves. wooded ravines. blunt branches.