liverworts mosses
Transcription
liverworts mosses
Key features for identifying liverworts Key features for identifying Mosses Growth 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 filaments or into broader lobes, the lobes can be of different sizes and can be bent over or under each other and formed into flaps 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. Growth form. Leaving aside the very distinctive bogmosses (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. 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 floerkei), Bifid Crestwort (Lophocolea bidentata), Creeping Fingerwort (Lepidozia reptans), part of stem with ‘fingered’ leaves. Above - Left: acrocarpous Scott’s Fork moss (Dicranum scottianum) ; right: pleurocarpous Larger Mouse-tail Moss (Isothecium alopecuroides). Above - Irregular branching in Red-stemmed Feathermoss (Pleurozium schreberi) on the left and regular tri-pinnate branching in Glittering Wood-moss (Hylocomium splendens) on the right; note the red stems. Abbreviations 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 fine 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 (reflexed or squarrose). Some leaves also have teeth on the margin, usually visible only with a hand lens. NR – nationally rare; NS – nationally scarce; BAP – a Biodiversity Action Plan species; S8 – listed on Schedule 8 of the Wildlife and Countryside Act (1981). Further information Books British Mosses and liverworts: a field guide. British Bryological Society (2010). The first comprehensive colour field 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 field 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 advice www.britishbryologicalsociety.org.uk. The British Bryological Society has an excellent website with useful information on publications, courses, field meetings and lots of pictures. www.nwdg.org.uk. The Native Woodland Discussion Group runs courses on Atlantic mosses and liverworts. Advice and Support Plantlife 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 [email protected] © 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. 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). 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 floerkei), never has gemmae and lacks points on the lobes 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. 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 field 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 fingernail to see the stem colour. Hatcher’s Paw-wort (Barbilophozia hatcheri) Michael Lüth Heller’s Notchwort (Anastrophyllum hellerianum) Horned Flap-wort (Lophozia longidens) Michael Lüth Above - Left: a thallose liverwort, Overleaf Pellia (Pellia epiphylla) with fruits; right: a leafy liverwort with round leaves, Autumn Flapwort (Jamesoniella autumnalis). MOSSES Monster Paw-wort (Tetralophozia setiformis) Michael Lüth LIVERWORTS LIVERWORTS 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. Curled Notchwort (Anastrophyllum saxicola) 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. Ciliated Fringewort (Ptilidium ciliare) 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. 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. 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 identified. Leafy liverwort; size: medium-sized but often forming large cushions or patches; branching: irregularly pinnate, occasionally bipinnate, 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. Bryophytes of Scotland’s pine woodlands British Lichen Society MOSSES Introduction This Plantlife field 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 significant 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 flora; 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 flora and affinities 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 floor species can cover rocks and the species on rocks and the bases of trees are often the same. Woodland floor: on the soil of the woodland floor, species have to compete with flowering 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 Woodmoss (Hylocomium splendens), Big Shaggy-moss (Rhytidiadelphus triquetrus), Red-stemmed Feathermoss (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 floor 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 Fringemoss (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 flora on birch is also poor but where there are hazels or aspens the flora can be quite rich, particularly with species of Bristle-mosses (Orthotrichum). Dead wood can often provide a humid habitat and has an interesting flora; 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 finer features on which identification 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 flowers 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. Green Shield-moss (Buxbaumia viridis) Broom Fork-moss (Dicranum scoparium) Dusky Fork-moss (Dicranum fuscescens) Rusty Fork-moss (Dicranum spurium) Wavy Fork-moss (Dicranum undulatum) Glittering Wood-moss Heath Plait-moss (Hypnum jutlandicum) (Hylocomium splendens) 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-astick’. 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. Waved Silk-moss (Plagiothecium undulatum) 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 floor, ledges, boulder tops; note: the whitish-green colour resembling ‘white worms’ and the undulate leaves make this an easy moss to recognise. 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 floor, 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. Red-stemmed Feather-moss (Pleurozium schreberi) 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 roundedtriangular, tapering to a blunt point, no nerve; habitat: woodland floor, ledges, boulder tops 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 fine, toothed apex (lens), leaves often curved in one direction; habitat: on drier open parts of the woodland floor, 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 fine, wispy apex is distinctive, particularly when dry. Common Haircap (Polytrichum commune) Bristly Haircap (Polytrichum piliferum) 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). 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 fine 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. Ostrich-plume Feather-moss (Ptilium crista-castrensis) 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 filaments under the lens; leaves: narrowly triangular, tapering to a long fine point, no nerve, all strongly curved in one direction; habitat: woodland floor, ledges, boulder tops; note: a beautiful and easily recognisable moss and very characteristic of pinewoods. 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 difficult plant to spot despite its size. Big Shaggy-moss (Rhytidiadelphus triquetrus) Pleurocarpous; size: very robust and often forming large patches; branching: irregular and rather untidy; colour: yellow-green, strawcoloured when dry; stem: red but often obscured by the leaves; leaves: triangular, pale yellow-green and again rather untidy; habitat: woodland floor, ledges, boulder tops; note: easily recognised by the large, erect stems and untidy, ‘chaffy’ leaves. 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 ‘stepstair moss’; leaves: broadly triangular, no nerve; habitat: woodland floor, ledges, boulder tops. Little Shaggy-moss 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 flat; colour: pale-green, often whitish when dry; stem: green; leaves: narrow, rounded-triangular, no nerve and with a fine 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. Red Bog-moss (Sphagnum capillifolium) (Rhytidiadelphus loreus) 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 floor, ledges, boulder tops; note: a bit smaller, neater and greener than Big Shaggy-moss (Rhytidiadelphus triquetrus), with hooked shoottips, 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. LIVERWORTS Above - Left: a thallose liverwort, Overleaf Pellia (Pellia epiphylla) with fruits; right: a leafy liverwort with round leaves, Autumn Flapwort (Jamesoniella autumnalis). MOSSES MOSSES Key features for identifying liverworts Key features for identifying Mosses Growth 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 filaments or into broader lobes, the lobes can be of different sizes and can be bent over or under each other and formed into flaps 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. Growth form. Leaving aside the very distinctive bogmosses (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. 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 floerkei), Bifid Crestwort (Lophocolea bidentata), Creeping Fingerwort (Lepidozia reptans), part of stem with ‘fingered’ leaves. Above - Left: acrocarpous Scott’s Fork moss (Dicranum scottianum) ; right: pleurocarpous Larger Mouse-tail Moss (Isothecium alopecuroides). 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? Above - Irregular branching in Red-stemmed Feathermoss (Pleurozium schreberi) on the left and regular tri-pinnate branching in Glittering Wood-moss (Hylocomium splendens) on the right; note the red stems. Abbreviations Further information Books Information and advice www.britishbryologicalsociety.org.uk. The British Bryological Society has an excellent website with useful information on publications, courses, field meetings and lots of pictures. www.nwdg.org.uk. The Native Woodland Discussion Group runs courses on Atlantic mosses and liverworts. Advice and Support Plantlife 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 [email protected] © 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. Woolly Fringe-moss (Racomitrium lanuginosum) 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, fine 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. 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. Red Bog-moss (Sphagnum capillifolium) Skye Bog-moss (Sphagnum skyense) 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. 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. 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 fine 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 (reflexed or squarrose). Some leaves also have teeth on the margin, usually visible only with a hand lens. 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 defined by Hill and Preston (1998). British Mosses and liverworts: a field guide. British Bryological Society (2010). The first comprehensive colour field 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 field 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. Colour. Colour and texture are all-important field 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 fingernail to see the stem colour. Curve-leaved Bow-moss (Dicranodontium uncinatum) 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). 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. 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. Bryophytes of Scotland’s oceanic heath British Lichen Society LIVERWORTS This Plantlife field 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 flora 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 finding out more about this remarkable community, with molecular work revealing two new species in the past few years. 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 finer features on which identification 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 flowers 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. Lindenberg’s Featherwort Orkney Notchwort (Adelanthus lindenbergianus) (Anastrepta orcadensis) Donn’s Notchwort (Anastrophyllum donnianum) © Maren Flagmeier Introduction Where to look for the oceanic heath 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: superficially similar to Anastrophyllum donnianum and Anastrepta orcadensis but the incurved upper leaf margin is unique; currently known from just one site on Islay. Wood’s Whipwort 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. Taylor’s Flapwort (Mylia taylori) (Mastigophora woodsii) Oceanic, NS. Leafy liverwort; size: mediumsized 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. 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. Carrington’s Featherwort (Plagiochila carringtonii) Leafy liverwort; size: medium to large, usually in large swelling cushions; colour: variable, in unshaded places it often purplishred 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: mediumsized 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 flattened, 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. Alpine Notchwort (Anastrophyllum alpinum) Oceanic, NS. Leafy liverwort; size: mediumsized, 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 difficult to separate from it; it is smaller, usually forms denser turfs and may occur in damper sites. Prickly Featherwort (Plagiochila spinulosa) Oceanic. Leafy liverwort; size: mediumsized 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. Lesser Whipwort (Bazzania tricrenata) Leafy liverwort; size: small but may form large cushions or straggling patches; branching: frequent thin branches from bottom of stem (flagellae) 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. Purple Spoonwort (Pleurozia purpurea) 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. Arch-leaved Whipwort (Bazzania pearsonii) 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. Western Earwort (Scapania gracilis) 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. Juniper Prongwort (Herbertus aduncus ssp. hutchinsiae) 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 (flagellae) 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. Cloud Earwort (Scapania nimbosa) 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 purplishred, rarely reddish-green or orange; leaves: two lobed and the lobes divided almost to the base, the smaller upper lobe bent flat 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. Northern Prongwort (Herbertus borealis) 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 flagelliferous branches; note: currently known only from Beinn Eighe, its only European site. Bird’s-foot Earwort (Scapania ornithopodioides) 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 flat 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.