Bryum calophyllum R.Br.

Matted (or Blunt) Bryum BRYACEAE SYN: Bryum acutiforme Limpr. in I.Hagen, B. axel-blyttii Kaurin ex H.Philib. Status Bryophyte Red Data Book - Vulnerable (2001) [should be revised to Endangered]

BAP Priority Species

Status in Europe: Rare Lead Partner: Plantlife International

UK Biodiversity Action Plan This is the current BAP target following the 2001 Targets Review: T1 - Maintain all extant populations of Matted Bryum. Progress on targets as reported in the UKBAP 2002 reporting round can be viewed by selecting this species and logging in as a guest on the following web page: http://www.ukbap.org.uk/ The full Action Plan for Bryum calophyllum can be viewed on the following web page: http://www.ukbap.org.uk/UKPlans.aspx?ID=166 Work on Bryum calophyllum is supported by

1

Contents 1 Morphology, Identification, Taxonomy & Genetics................................................3

1.1 MORPHOLOGY & IDENTIFICATION...............................................................3 1.2 TAXONOMIC & GENETIC CONSIDERATIONS..................................................5

2 Distribution & Current Status ...........................................................................6 2.1 World ......................................................................................................6 2.2 Europe ....................................................................................................6 2.3 United Kingdom........................................................................................6

2.3.1 England .............................................................................................8 2.3.2 Northern Ireland..................................................................................8 2.3.3 Scotland.............................................................................................8 2.3.4 Wales ................................................................................................9

3 Ecology & Life Cycle........................................................................................9 4 Habitat Requirements ...................................................................................10

4.1 The Landscape Perspective.......................................................................10 Habitats in Europe .....................................................................................10 Habitats in the UK......................................................................................10

4.2 Communities & Vegetation .......................................................................11 4.3 Summary of Habitat Requirements ............................................................13

5 Management Implications ..............................................................................13 Site Safeguard ..........................................................................................13 Advisory...................................................................................................13

6 Threats / Factors Leading to Loss or Decline or Limiting Recovery .......................13 Development ............................................................................................14 Dune Stabilisation......................................................................................14 Vegetation Successes .................................................................................14 Digging of Ditches or other Drainage Works...................................................14 Eutrophication of Water Supplies..................................................................14 Trampling.................................................................................................14 Disturbance due to Motor Vehicles................................................................14 Collection of Specimens ..............................................................................14 coastal defence systems .............................................................................14 Population locality......................................................................................14

7 Recommendations for future work ..................................................................14 Monitor populations....................................................................................14 Maintain contacts with site-managers and landowners ....................................15 Share data................................................................................................15 Ex situ propagation and population re-establishment ......................................15 Coordination of work ..................................................................................15 Site management ......................................................................................15 Further survey work...................................................................................15

8 References ..................................................................................................15 9 Acknowledgements .......................................................................................16 10 Contacts...................................................................................................17 11 Links........................................................................................................17

2

1 Morphology, Identification, Taxonomy & Genetics

1.1 MORPHOLOGY & IDENTIFICATION Bryum calophyllum (Matted or Blunt Bryum) is an attractive acrocarpous moss that grows in foredune slacks and other sandy coastal habitats. It is a small plant, 1-1.5cm tall, which can be distinguished from most other Bryum species by its leaves which are blunt.

Figure 1 - Bryum calophyllum with immature capsules

3

Figure 2 - Gametophytes of Bryum calophyllum.

Figure 3 - Mature and almost mature capsules of Bryum calophyllum.

4

Bryum is a taxonomically difficult genus in which many species can only be identified when mature capsules are present. B. calophyllum is one of the less critical species because of the ovate leaves with shortly acute to rounded tips, but its secure identification should also be based on all of the following characters: leaves not red at base, narrowly bordered, shortly ovoid capsule pendulous on relatively long seta, autoicous sexuality, endostome with rudimentary cilia, spore size 26-42 µm. Fertile B. calophyllum can usually be identified in the field by experienced bryologists using the combination of a shortly ovoid capsule on a rather long seta with wide concave leaves having shortly acute to blunt apices. Forms of the B. dichotomum group growing on coastal sands may also have short ovoid capsules and blunt concave leaves, but these are commonly brighter green and often have axillary bulbils. Characters that aid in detecting non-fertile B. calophyllum in the field are the small plants with rather wide and concave leaves that are shortly acute to obtuse, leaves rather evenly spaced along the stem and patent to spreading, leaves and stems typically light olive to tan or pinkish. B. pallens can look superficially similar, but it has flat rather than concave leaves that are usually more pointed at the apex. That species is also often a brighter pink to vinous-red colour. B. warneum often resembles B. calophyllum in habitat, size and colour, but differs markedly in its more pointed leaves that are erect to erecto-patent and loosely imbricate. B. marratii resembles B. calophyllum in having wide concave leaves with shortly acute to blunt apices. Plants with mature capsules (which are infrequent in B. marratii) are easily separated by the usually longer capsule of B. calophyllum (typically ovoid cf. almost globular) with a shorter beak on the operculum (conical cf. shortly rostellate). Study of non-fertile plants associated with those bearing capsules has shown that small shoots often cannot be reliably separated (except perhaps by the colour of fresh or living plants: green in B. marratii, strongly tinged olive, tan, reddish or pinkish in B. calophyllum ). Well-grown non-fertile plants (at least 5 mm tall) can usually be distinguished in the herbarium from B. marratii by careful appraisal of a combination of characters, as follows: upper leaves shorter and relatively wider in B. calophyllum (normally with length:breadth <2:1); leaves with acute to blunt apex rather than a blunt to broadly rounded apex; leaf border of narrow cells usually (not always) better developed; leaf margin more often recurved below; costa extending nearer to leaf apex. The temptation to name small plants (<5 mm) on the basis of these differences should be resisted, as errors will sometimes result.

1.2 TAXONOMIC & GENETIC CONSIDERATIONS The north-Scandinavian and arctic Bryum axel-blyttii with acute leaf apices has usually been treated as a separate species but Holyoak (2004) argues that it is a form of B. calophyllum and that B. acutiforme represents intermediate plants. It is reported that all three taxa sometimes occur together in northern Norway (Finmark) accompanied by intermediates (Nyholm 1993). Matted Bryum is monoicous and produces spores freely so that it may be assumed to be an out-breeding species. There is no evidence of vegetative reproduction from gemmae, bulbils or tubers. There have been no studies of its population genetics (e.g. using isozymes) or molecular genetic studies to establish its relationships to other Bryum spp.

5

2 Distribution & Current Status

2.1 WORLD Widespread in arctic and subarctic regions of Europe, Asia, Greenland and North America, occurring further south in North America (reported from Newfoundland, Alberta, Montana, California, N. Dakota), in Europe (see below) and Tibet. Reports from Central Africa (e.g. Smith 1978: 390) are erroneous and probably based on B. afro-calophyllum P.de la Varde from Kenya.

2.2 EUROPE Widespread in the north, in Iceland, Svalbard, Faeroes, Norway, Sweden, Finland, Latvia and northern Russia, occurring sparsely further south in Ireland, Great Britain, Netherlands, Germany, Denmark and Poland. Also reported from Rumania (Transsylvania). French records are erroneous and a report from Sardinia seems unlikely and is thus in need of confirmation. B. calophyllum is listed as ‘Rare’ for Europe as a whole (Schumacker & Martiny 1995: 39), where it has shown marked declines towards its southern range limits. It is currently known from three or four localities in Ireland (National Parks and Wildlife Service, unpublished data). Recorded as extinct at most of its old localities in Germany, with the only recent record from Inseln Borkum (Düll 1994: 87). Rare and very local in Norway and Sweden; two forms treated as conspecific here ('B. axel-blyttii, B. acutiforme') have also been regarded as rare and localised in Scandinavia. In the former USSR it was described as a rather rare arctic-alpine species (Savicz-Ljubitzkaja & Smirnova 1970: 620), but with widespread records from Latvia to arctic Siberia and Central Asia.

2.3 UNITED KINGDOM B. calophyllum has been recorded historically in England, Scotland and Wales but not in Northern Ireland. Records were listed and reviewed by Holyoak (2001, 2002a, 2002b). They are summarised below for each country in order of vice-counties. The species is rare in Britain, where it has become extinct at all but about 7 of the approximately 21 localities recorded since the mid-nineteenth century, due to habitat changes and coastal development. All sites in England have become extinct, a single small population survives on Anglesey (north-west Wales) and there are three confirmed populations surviving in Scotland. The loss of all English and all but one of the Welsh populations confirmed since 2000 implies that the U.K. Red List status of the species should be revised from Vulnerable to Endangered. Most records of Bryum calophyllum, other than those found during current surveys are now more than 25 years old. During 2000, Bryum calophyllum was not re-found at three of its old sites on the east coast of Scotland but, during 2001, new populations were found on Islay and on the north coast at Bettyhill. This means that of the four confirmed extant sites for Bryum calophyllum in the UK, three are in Scotland (Rothero 2003).

6

Figure 4 – Current and historical distribution of Bryum calophyllum

7

2.3.1 ENGLAND vc4: North Devon Recorded at Braunton Burrows [SS43] 1900. vc58: Cheshire Recorded at Wallasey [SJ29] 1902-1903. vc59: South Lancashire Records from what is now the Sefton Coast area (Southport, Ainsdale, and nearby: SD20, SD21, SD31) were numerous from 1854 to 1933 and there are also a few records inland near Taunton [SD90] in 1865. The only recent record was from Ainsdale [SD31] in 1973. vc62: North-east Yorkshire Recorded at Coatham Marshes [NZ52] 1891-1896. vc66: Durham Recorded at Snook Point, Seaton Carew [NZ52] 1902-1905 and earlier. vc68: North Northumberland Recorded at Holy Island [NU14] in 1924. vc69: Westmorland Recorded from Sandscale Haws [SD27] in 1969.

2.3.2 NORTHERN IRELAND No records.

2.3.3 SCOTLAND vc75: Ayrshire Recorded at Stevenston [NS24] in 1900. vc82: East Lothian Recorded at Aberlady Bay (including Gullane Bay, Gullane Links and Luffness Links) [NT48] 1887-1974. vc85: Fife Recorded at Tentsmuir [NO42] 1853-1963. vc90: Angus Recorded at Barry Links [NO53] 1886-1965. vc95: Moray Recorded at Lossiemouth [NJ27] 1953-1959. vc104: North Ebudes Recorded at Killinallan Bay, Islay [NC37] 2001. vc105: West Ross Recorded at Achanahaird Bay [NC01] 1960-2000. vc108: West Sutherland

8

Recorded at Bettyhill [NC66] 2001. vc109: Caithness Recorded at 'Caithness' [ND26] in 1949 and at Dunnet Links [ND27] 1900-1913.

2.3.4 WALES vc46: Cardiganshire Recorded at Ynyslas [SN69] 1948-1973. vc48: Merioneth Recorded at Morfa Dyffryn [SH52] 1902-1972 and at Towyn Sandhills [SN59] in 1902. vc49: Caernarvonshire Recorded at Minffordd, Afon Glaslyn [SH53] in 1916. vc52: Anglesey Recorded at Newborough Warren [SH46] in 1908 and at Tywyn Aberffraw [SH36] in 2001.

3 Ecology & Life Cycle The regular spore production in B. calophyllum and apparent absence of bulbils or tubers in the species leaves little doubt that the species initially becomes established from spores. The rather large size of the spores (26-42 µm in diameter) probably implies that their dispersal distances tend to be less than for most mosses, but strong coastal winds in late autumn and winter may nevertheless distribute them widely. Large spores might also contain large food reserves to support early growth stages, or possibly to allow several years of spore dormancy. Very little is known about the longevity of individual plants and patches of B. calophyllum . At some sites the patches may be lost after a few years due to shading as taller vegetation becomes established. The species is monoicous (autoicous). Capsules are frequent, at least in Britain, Ireland, Germany and Scandinavia. They mature from late summer to early winter in Britain (Smith 1978: 390, A.C. Crundwell in Hill et al. 1994: 84), in October in Ireland (DT Holyoak unpublished) and in summer or autumn in Scandinavia (Nyholm 1993). It is probably a ‘short-term shuttle’ species, exploiting gaps on damp, open, calcareous sand and disappearing if a stable, closed vegetation develops. For all of its known Scottish populations, natural processes involving wind-blown or water-deposited sand on the sites means that there is a regular supply of new gaps to be colonised and this supply may be augmented by the grazing regime of sheep or rabbits. Bryum calophyllum can exploit these gaps through its production of abundant spores which are shifted around with the sand. Population levels at any one site are likely to fluctuate according to the amount of suitable habitat available at any one time. The spore bank is vital in enabling the plant to rapidly colonise new areas of suitable habitat as soon as they become available (Rothero 2003).

9

4 Habitat Requirements

4.1 THE LANDSCAPE PERSPECTIVE HABITATS IN EUROPE Recent records of B. calophyllum from Ireland are from coastal dune or machair areas on damp calcareous sand with sparse open vegetation (National Parks and Wildlife Service, unpublished data). In Germany it formerly occurred in coastal sand-dune areas, sporadically at saline localities inland, with a record also from mud of a dry fish-pond (Limpricht 1895: 307). In Scandinavia it grows on moist or wet sandy or gravelly soil beside the sea, streams or lakes, in the south and in arctic regions (Nyholm 1993: 173). Also reported in northern Scandinavia as 'B. axel-blyttii' growing on moist sandy or gravelly soil beside lakes, streams, etc. in mountains and in arctic regions, often with Pohlia filum or Aongstroemia longipes, sometimes in mixed tufts with [typical] B. calophyllum (Nyholm 1993: 171, 172). In arctic regions of former USSR it grows on moist and wet sandy, silty or gravelly soils along seacoasts, on flood-terraces, in arctic meadows with Dupontia, in Hypnum-dominated mires, tundras and arctic deserts; in continental-arctic regions on soil (often sandy or silty) of saline land, periodically flooded river shores and flooded places in grassy mires (Abramova et al. 1961: 414, Savicz-Ljubitzkaja & Smirnova 1970: 620-621). In W. Siberian Arctic on sand in herb-moss tundra and on bare sand in disturbed places (Dyachenko et al. 1999). Also in W. Siberian Arctic, reported as 'B. axel-blyttii' growing on sand in dwarf shrub-herb community on river terrace (Czernyadjeva 2001: 133). HABITATS IN THE UK In Britain B. calophyllum grows on moist, mainly bare sandy soils, particularly in coastal dune slacks which are at least moderately calcareous; a single inland record (1865) was from an old gravel pit (Church et al. 2001: 67); also occasionally on damp sand in estuaries (A.C. Crundwell in Hill et al. 1994: 84). Habitat notes on old specimens from England and Wales consistently refer to sand or dunes, with a few noting dune-slacks. Several specimens were evidently growing on sand, often with Bryum pseudotriquetrum. The very large gatherings made by PW Richards at Morfa Dyffryn were of plants growing scattered on partly bare sand. The only population still known in Wales (Holyoak 2002a) grows on partly bare damp sand with low, open, early-successional vegetation in a dune-slack just beneath a blow-out at Tywyn Aberffraw. At Achnahaird in West Ross (Scotland), Bryum calophyllum occurs in both open communities on bare sand and in quite closed areas of sandy grassland, but all areas are associated with flushing by surface water from calcareous shell-sand slopes above. The site on Islay is broadly similar in ecology although the flushing here is from a single small burn that flows through the dunes although there is a major difference in the local occurrence here of halophytic species like Glaux maritima and Juncus gerardii. At Bettyhill, like Achnahaird, the site is well above the most extreme tides, but Bryum calophyllum again occurs on open shell-sand on terraces beside, and presumably created by, a small burn (Rothero 2003).

10

Figure 5 - Habitat of Bryum calophyllum in dune slack with bare sand exposed.

4.2 COMMUNITIES & VEGETATION Table 1 records habitat details and associated plants in 50 x 50 cm quadrats at the last surviving Welsh site (from Holyoak 2002a). This population was growing with open, low vegetation which corresponds to NVC type SD13 of Rodwell (2000: 214), the Sagina nodosa - Bryum pseudotriquetrum dune-slack community, in which common associates include Aneura pinguis, Bryum pseudotriquetrum, other Bryum spp., Carex arenaria, Juncus articulatus, Leontodon saxatilis, Sagina nodosa and Salix repens. Table 1. Habitat characteristics and associated plants in 50 x 50 cm quadrats with Bryum calophyllum at Tywyn Aberffraw, Anglesey in September 2001. QUADRAT NO. 1 2 3 4 SLOPE (°) 0-3 2-4 0-2 2 ASPECT SW NW W SW VEGETATION HEIGHT MEAN 0-5 0-6 0-6 0-7 (CM) MAXIMUM 22 21 20 22 PERCENTAGE COVER

Bare Sand 40 35 20 57 Vascular Plants Agrostis stolonifera 3 5 4 3 Anagallis tenella 3 5 - 1 Carex arenaria 4 - 6 - Carex viridula ssp. Oedocarpa 6 2 - 9 Equisetum arvense 2 3 1 1 Equisetum variegatum - - 2 - Euphrasia sp. <1 1 1 - Hydrocotyle vulgaris 2 - 3 1 Juncus articulatus 4 6 8 4 Leontodon saxatilis - - 2 - Plantago coronopus <1 - - -

11

QUADRAT NO. 1 2 3 4 SLOPE (°) 0-3 2-4 0-2 2 ASPECT SW NW W SW VEGETATION HEIGHT MEAN 0-5 0-6 0-6 0-7 (CM) MAXIMUM 22 21 20 22 PERCENTAGE COVER

Potentilla anserine - 3 3 - Radiola linoides - - <1 <1 Sagina nodosa - - - <1 Salix repens 25 6 13 19 Bryophytes Bryum calophyllum 2 2 2 2 Bryum pseudotriquetrum 2 5 4 2 Bryum warneum - - <1 - Campylium stellatum var.

stellatum - 12 - -

Climacium dendroides 1 - - - Pellia endiviifolia 5 15 30 1 Riccardia incurvata 1 - - -

Table 2. Species associated with Bryum calophyllum on two or more of six stands on three sites. 1998 & 2001 (Rothero 2003).

SPECIES 1A 1B 1C 2A 2B 3 f Bare sand A A A O A A 6 Bryum spp O O O O F F 6 Barbula spadicea F O F O O 5 Juncus bulbosus F O F F O 5 Sagina nodosa O F O O O 5 Aneura pinguis O O O O 4 Bellis perennis F O O O 4 Festuca rubra O A F A 4 Pohlia wahlenbergii O R F O 4 Pellia endiviifolia R F O 3 Petalophyllum ralfsii F O O 3 Poa annua F R F 3 Potentilla anserina O R F 3 Agrostis stolonifera A F 2 Barbula fallax O O 2 Barbula unguiculata O O 2 Carex arenaria F O 2 Carex nigra R R 2 Eleocharis palustris F F 2 Holcus lanatus O O 2 Homalothecium lutescens F R 2 Hydrocotyle vulgaris O O 2 Mentha aquatica F R 2 Parnassia palustris O O 2 Riccardia latifrons R R 2 Site 1) Achnahaird Bay (1998); 2) Killinallan (2001); Bettyhill (2001). Letters are DAFOR values.

12

4.3 SUMMARY OF HABITAT REQUIREMENTS In Britain B. calophyllum grows on moist, mainly bare sand, particularly in coastal dune slacks which are at least moderately calcareous. It is associated with low, open, early-successional vegetation.

5 Management Implications In 2003 Plantlife International published an illustrated leaflet giving an account of various threatened bryophytes of coastal dune slacks as part of the 'Back from the Brink Management Series', entitled: Looking after rare mosses and liverworts in coastal dune slacks. This leaflet is intended to increase awareness of the species along with other dune-slack bryophytes. It describes Bryum calophyllum, discusses threats to its populations and provides details of contacts able to supply additional information. SITE SAFEGUARD It is important to:

Ensure that all sites for B. calophyllum which are not currently SSSIs but which are at all likely to be threatened by any kind of damaging activity are considered for notification as SSSIs.

Ensure the safeguard of all B. calophyllum sites through the preparation of

Site Management Statements and the negotiation of Management Agreements.

Encourage managers of land where conservation is the primary concern to

appreciate the importance of this species and to implement action to optimise its chances of survival and encourage expansion of its range. In particular, the needs for mobile sand, a seasonally high water table, unpolluted ground water, little or no trampling and very short vegetation should be stressed.

Prevent disruption or pollution of fresh water sources on sites where this may

have an adverse affect on populations or on the integrity of the site.

ADVISORY It is important to:

Inform all relevant landowners, statutory authorities and local authorities of

the presence of the plant, to ensure that they are aware of and fulfill their responsibilities for its management and safeguard.

Advise relevant landowners and site managers of the requirements of the

species and measures needed to prevent damage to its populations. In particular, the needs for mobile sand, a seasonally high water table, unpolluted ground water and very short vegetation should be stressed.

6 Threats / Factors Leading to Loss or Decline or Limiting Recovery The loss of Bryum calophyllum from a number of its known sites is generally attributed to the loss or degradation of its habitat. This can occur through a number of ways.

13

DEVELOPMENT In the past, sites were lost to residential and industrial developments and to holiday developments (particularly golf course construction) and the associated increase in recreational pressures. DUNE STABILISATION Dune stabilisation is a serious threat that has probably led to loss of several sites. It promotes vegetation succession at dune-slacks in which this species grows. In some cases, coastal protection works has increased dune stabilisation. Dunes can also be stabilized for agriculture or silviculture. VEGETATION SUCCESSES Vegetation succession is a potential threat at all sites, particularly associated with a decline in rabbit populations or reduction in the supply of wind-blown sand. DIGGING OF DITCHES OR OTHER DRAINAGE WORKS A seasonally high water table is an important feature of many sites and survival of the species is unlikely if they are drained. EUTROPHICATION OF WATER SUPPLIES Low-nutrient status and short vegetation characterise sites with B. calophyllum. Increase in the nutrient status of waters could threaten it directly and through causing competing plants to grow taller. Usage of artificial fertilisers on farmland or golf courses should be avoided near to known sites for B. calophyllum. TRAMPLING B. calophyllum is easily damaged by trampling. Unlike Petalwort (Petalophyllum ralfsii), it does not grow on trampled pathways. DISTURBANCE DUE TO MOTOR VEHICLES Limited disturbance of the ground surface may be beneficial but if too extensive or too regular it may reduce populations or destroy the habitat. Illegal motorcycle scrambling has been noted as a potential threat. However, increased mobility of sand due to vehicles and other disturbance, as well as natural processes, often maintains the open vegetation conditions needed by B. calophyllum, although too rapid accretion of sand may result in at least temporary losses of its populations. COLLECTION OF SPECIMENS Thoughtless collection of specimens might continue to pose a threat, especially to small populations. COASTAL DEFENCE SYSTEMS The constructions of coastal defences can lead to dramatic changes in habitat (Rothero 2003). POPULATION LOCALITY All the Bryum calophyllum populations are very local and small changes in management at any of the sites, particularly affecting the quality of the water run-off to the sites, could threaten them (Rothero 2003).

7 Recommendations for future work MONITOR POPULATIONS Monitoring of all U.K. populations should be carried out at three-yearly intervals, i.e. if possible in autumns of 2007-08, 2010-11, etc.

14

MAINTAIN CONTACTS WITH SITE-MANAGERS AND LANDOWNERS Ensure that all relevant site-managers and land-owners remain aware of populations of Bryum calophyllum on land they control, of the conservation needs of the species and of any threats, potential threats or other significant developments. SHARE DATA Information on current population status, threats, ecology, etc. should be supplied to Threatened Bryophyte Database, JNCC and BRC for incorporation into national databases. It is also necessary to provide information on current status annually or as it becomes available to World Conservation Monitoring Centre. EX SITU PROPAGATION AND POPULATION RE-ESTABLISHMENT The loss of all English and all but one of the Welsh populations confirmed since 2000 implies that the U.K. Red List status of the species should be revised from Vulnerable to Endangered. Urgent consideration should therefore be given to population re-establishment of the species and also to establishment of ex situ cultures. COORDINATION OF WORK The requirements of this species should be considered in the delivery of the action plans for coastal sand dunes, the moss Bryum warneum and the liverwort Petalophyllum ralfsii. SITE MANAGEMENT Bryum calophyllum needs damp, open, calcareous sand. It cannot compete with larger species and disappears when a significant humic content builds up in the sand. It exploits gaps in the vegetation created by dynamic processes of water and wind that are usually part of sand dune systems and will be threatened by anything that tends to stabilize the habitat. Any management which encourages the maintainance of these dynamic processes will benefit Bryum calophyllum (Rothero 2003). FURTHER SURVEY WORK There are a large number of other sand-dune systems in the west of Scotland, particularly on the islands, that have never been surveyed for Priority Bryophyte species. The fact that a large population of Bryum calophyllum could exist undetected at a well-known botanical site like Bettyhill suggests that further survey work in the west is an urgent priority (Rothero 2003)..

8 References Abramova, A.L., Savicz-Ljubitzkaja, L.I. & Smirnova, Z.N. 1961. [Handbook for the

determination of the mosses of arctic Russia]. Moscow & Leningrad: Inst. bot. V. L. Komarovii Akad. Sci. USSR [in Russian].

Blockeel, T.L. & Long, D.G. 1998. A check-list and census catalogue of British and Irish bryophytes. Cardiff: British Bryological Society.

Breeds, J. & Rogers, D. 1998. Dune management without grazing: a cautionary tale. Enact, Managing Land for Wildlife 6: 18-22.

Church, J.M., Hodgetts, N.G., Preston, C.D. & Stewart, N.F. 2001. British red data books, mosses and liverworts. Peterborough: JNCC.

Corley, M.F.V. & Hill, M.O. 1981. Distribution of Bryophytes in the British Isles. A Census Catalogue of their occurrence in Vice-Counties. Cardiff.: British Bryological Society.

Czernyadjeva, I.V. 2001. Moss flora of Yamal Peninsula (West Siberian Arctic). Arctoa 10: 121-150.

Dixon, H.N. 1954. The student’s handbook of British mosses. 3rd ed. Eastbourne: Sumfield & Day.

15

Düll, R. 1985. Distribution of the European Mosses, Part II. Bryologische Beiträge 5: 110-232.

Düll, R. 1994. Deutschlands Moose. ... 2. Teil. Bad Münstereifel - Olerath: IDH-Verlag. Dyachenko, A.P., Morozova, L.M., Stepanova, A.V. & Magomedova, M.A. 1999.

Contributions to the mos flora of the Yamal Peninsula (Siberian Arctic). Arctoa 8: 73-78.

Hill, M.O. 1988. A bryophyte flora of north Wales. Journal of Bryology 15: 377-491. Hill, M.O., Preston, C.D. & Smith, A.J.E. 1994. Atlas of the Bryophytes of Britain and

Ireland. 3. Mosses (Diplolepideae). Colchester: Harley Books. Holyoak, D.T. 2001. Coastal mosses of the genus Bryum. Report to Plantlife on work

carried out during 2000. London: Plantlife Report number 178 [unpublished]. Holyoak, D.T. 2002a. Coastal mosses of the genus Bryum. Report to Plantlife on work

carried out in Wales during 2001. London: Plantlife Report 203: 1-49. Holyoak, D.T. 2002b. Coastal mosses of the genus Bryum. Report to Plantlife on work

carried out in England during 2001. London: Plantlife Report 206: 1-43. Jóhannsson, B. 1995. Íslenskir mosar. Hnokkmosaætt. Reykjavik: Fjölrit Náttúrufræ

istofnunar 27. Limpricht, K.G. 1895. Die Laubmoose Deutschlands, Oesterreichs und der Schweiz. II.

Leipzig: Kummer. Nyholm, E. 1993. Illustrated Flora of Nordic Mosses. Fasc. 3. Bryaceae, etc. Copenhagen

& Lund: Nordic Bryological Society. Rodwell, J.S. (ed.) 2000. British Plant Communities. vol. 5. Maritime communities and

vegetation of open habitats. Cambridge: Cambridge University Press. Rothero, G. 2003. Blunt Bryum Bryum calophyllum. Species Dossier report to Scottish

National Heritage. Savicz-Ljubitzkaja, L.I. & Smirnova, Z.N. 1970. [The Handbook of the Mosses of the

U.S.S.R. The Mosses Acrocarpous.] Academy of Sciences of USSR & Komarov Botanical Institute.

Schumacher, R. & Martiny, P. (eds.) 1995. Red Data Book of European Bryophytes. Part 2. Threatened Bryophytes in Europe including Macaronesia. Trondheim: European Committee for the Conservation of Bryophytes.

Smith, A.J.E. 1978. The moss flora of Britain and Ireland. Cambridge: Cambridge University Press.

Smith, A.J.E. 2004. The moss flora of Britain and Ireland. 2nd ed. Cambridge: Cambridge University Press.

Steere, W.C. 1978. The Mosses of Arctic Alaska. Bryophytorum Bibliotheca 14: i-x, 1-508.

Stewart, N. (ed.) 1995. Red Data Book of European Bryophytes. Part 1: Introductory section & background. European Committee for the Conservation of Bryophytes.

Wilson, W. 1855. Bryologia Britannica; containing the mosses of Great Britain and Ireland ... London: Longman, Brown, Green, and Longmans.

9 Acknowledgements Tom Blockeel, John Breeds, Kathryn Childerhouse, the late Alan Crundwell, Phil Davey, Prof. Jeff Duckett, Ruth Davis, Dr Jenny Duckworth, Alan Hale, Nick Hodgetts, Geraldine Holyoak, Dr Nadezhda Konstantinova, Dr Neil Lockhart, Dr David Long, Mrs Jean Paton, Roy Perry, Ron Porley, Dr Chris Preston, Gordon Rothero, Dr Phil Smith, Dr Tony Smith, Dr Gill Stevens, Nick Stewart, Dr Ray Tangney, the late Dr Harold Whitehouse, Sally Whyman, Paul Wisse, Dan Wrench; National Parks and Wildlife Service, Sefton Coast Life Project; curators of bryophytes at BBSUK, BM, DBN, E, NMW, S. All photographs by David Holyoak.

16

10 Contacts Plantlife International The Wild Plant Conservation Charity 14 Rollestone Street Salisbury Wiltshire SP1 1DX Tel: 01722 342730

or contact enquiries: enquiries@plantlife.org.uk

11 Links

British Bryological Society http://www.britishbryologicalsociety.org.uk/.

Plantlife International’s Back from the Brink (species recovery) programme is supported by: Natural England, Scottish Natural Heritage & the Countryside Council for Wales.

ISBN: 1 904749-67-4

Original draft by David Holyoak First draft dated January 2005

Partial Revision December 2006 Edited by Plantlife International Last revised February 13, 2007

17