David T. Holyoak

New national and regional bryophyte records, 26

26 L T Ellis, H Bednarek-Ochyra, R Ochyra, Silvia Calvo Aranda, Maria T Colotti, Maria M Schiavone, Michail V Dulin, P Erzberger, Tulay Ezer, Recep Kara, Rosalina Gabriel, Lars Hedenas, David T Holyoak, P Odor, B Papp, M Sabovljevic, R D Seppelt, V R Smith, Andre Sotiaux, E Szurdoki, Alain Vanderpoorten, J van Rooy, J Żarnowiec Department of Botany, The Natural History Museum, UK, Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Krakow, Poland, 3 Departamento de Biodiversidad y Biologia Evolutiva, Museo Nacional de Ciencias Naturales, Madrid, Spain, Facultad de Ciencias Naturales e IML, San Miguel de Tucuman, Argentina, Institute of Biology Komi Science Centre UB RAS, Syktyvkar, Komi Republic, Russia, Belziger Str. 37, D-10823 Berlin, Germany, Faculty of Science and Arts, Department of Biology, Nigde University, Turkey, Departamento de Ciencias Agrarias, Universidade dos Acores, Angra do Heroismo, Portugal, Department of Cryptogamic Botany, Swedish Museum of Natural History, Stockholm, Sweden, Quinta da Cachopa, Barcoila, Cabecudo, Portugal, Department of Plant Taxonomy and Ecology, Lorand Eotvos University, Budapest, Hungary, Botanical Department, Hungarian Natural History Museum, Budapest, Hungary, Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade, Serbia, Australian Antarctic Division, Channel Highway, Kingston, Tasmania, Australia, Department of Botany, University of Stellenbosch, Matieland, South Africa, National Botanic Garden of Belgium, Domein van Bouchout, Meise, Belgium, Hungarian Natural History Museum, Budapest, Hungary, University of Liege, Institute of Botany, Belgium, National Herbarium, South African National Biodiversity Institute, Pretoria, South Africa, Department of Ecology and Nature Conservation, Institute of Environmental Protection and Engineering, University of Bielsko-Biala, Poland

Morphological, ecological and molecular studies of the intergrading taxa Bryum neodamense and B. pseudotriquetrum (Bryopsida: Bryaceae)

Abstract Studies of Bryum neodamense at most of its localities in the British Isles revealed that plants with morphology intermediate with B. pseudotriquetrum var. pseudotriquetrum frequently accompany B. neodamense. A survey of herbarium specimens found that similar intermediates are widespread in continental Europe and also occur in Asia, Alaska and Greenland. Phylogenetic analyses of DNA sequences from three loci imply that B. neodamense and intermediate plants are nested within the rather variable B. pseudotriquetrum. It is therefore concluded that, despite its striking morphological characters, B. neodamense is an inconstant phenotype of B. pseudotriquetrum, which occurs very locally in highly calcareous habitats that are subject to periodic inundation.

Ephemerum hibernicum sp. nov. (Bryopsida: Ephemeraceae) from Ireland

Abstract Ephemerum hibernicum sp. nov. is described from inundation zones of two lakes and a river in north-central Ireland and a river in western Ireland. It is most similar to E. crassinervium and E. sessile, but differs from both of these in several characters of the leaves. In particular, papillae on the upper lamina are much lower than in most E. crassinervium, while the leaves are longer and wider than in E. sessile, with a wider lamina towards the leaf apex, less well-defined costa and sometimes with large spinose marginal teeth. Ephemerum hibernicum is a rare species and its specialized habitat merits conservation.

A taxonomic revision of some European and Asian bulbiliferous species of Anomobryum (Bryophyta: Bryaceae)

Abstract The taxonomy of Anomobryum julaceum and allied species with axillary bulbils in Europe and Asia is reviewed. A. concinnatum is regarded as a distinct species, occurring in W. and C. Europe, SW., N. and C. Asia and N. America. A. bavaricum has often been confused with A. concinnatum, but differs in its more numerous, small, reddish bulbils and in leaf shape; it is known only from the European Alps. The Asian A. nitidum also has numerous small reddish bulbils but it differs from A. bavaricum in leaf shape and bulbil form. A lectotype is designated for A. nitidum, of which A. gemmigerum and other nominal taxa are regarded as synonyms. Information is presented on geographical ranges and habitats of the four valid species.

Notes on some rare and newly recorded bryophytes of metalliferous mine sites in Ireland

There have been few records from Ireland of the so-called ‘copper mosses’ or metallophytes, those few bryophyte species characteristic of substrata rich in copper and other heavy metals. None were recorded in the classical compilations by McArdle (1904) and Lett (1915), the first records being of Cephaloziella massalongi found at Allihies in Co. Cork by H. Milne-Redhead in 1955 (Castell, 1957) and east of Bunmahon in Co. Waterford by R. D. Fitzgerald and D. M. Synnott in 1966 (Paton, 1967). Subsequent visitors have added Pohlia andalusica from both of the same two sites (Holyoak, 2003). During 1999–2005 an extensive programme of research on Irish bryophytes was undertaken by National Parks and Wildlife Service (NPWS), resulting in many new records although few involved ‘copper mosses’ (Holyoak, 2006). Continuation of this work into Co. Cork in 2006 and Co. Waterford in 2007 resulted, however, in the first Irish records of Cephaloziella nicholsonii, Ditrichum cornubicum and Scopelophila cataractae among other finds at old copper mine sites. This led to suspicion that other populations of metallophyte bryophytes may hitherto have been overlooked elsewhere in Ireland. A wide-ranging survey of bryophytes at Irish metalliferous mine sites was therefore undertaken by NPWS during April to early June 2008, which produced numerous new finds, including Ditrichum plumbicola new to Ireland. The present paper summarises these new records of metallophyte bryophytes along with the others made by NPWS surveys since 1999; it also comments on the record of Ditrichum lineare from Allihies. The likely role of accidental transport by man in facilitating dispersal of metallophyte bryophytes is discussed. Taxonomy and nomenclature follow Hill et al. (2008) for bryophytes and Stace (1997) for phanerogams. Cephaloziella integerrima W. CORK (H3): on unshaded part-bare damp ground by track edge on old copper-mine site, ca 128 m alt., Allihies, V588457, 28 September 2006, DTH 06-324B & NL (BBSUK, DBN) (Blackstock, 2007). WATERFORD (H6): on almost unshaded damp clayey and humic soil of bank of ditch in copper-mine spoil on disturbed hillslope, ca 15 m alt., Knockmahon, X437990, 20 April 2008, DTH 08-129 (BBSUK, DBN), c.fr., also collected at several similar places nearby, where the population was apparently large. E. DONEGAL (H34): on unshaded thin soil among quartzite cobbles in disturbed area of grassland near buildings (with galvanised roofing), ca 50 m alt., Dunree Head, C286390, 8 August 2002, DTH 02-786 (BBSUK, DBN), c.fr. (Blackstock, 2003). The only previous records from Ireland were from a roadside bank east of Coumshingaun Lough in the Comeragh Mountains (H6), where it was discovered in 1966 by Mrs J. Appleyard and J. G. Duckett (Holyoak, 2003) and collected again in 1968 (Mrs J. A. Paton). This locality was apparently not metalliferous; it was not refound in 2007 or 2008. The recent records thus increase the number of sites reported from Ireland from one to four and show the species is still extant, with at least one large population. All three recent records are from substrata likely to have high levels of copper (two sites) and zinc (one site). Most of the older British records were not from metalliferous substrata (Macvicar, 1926; Paton, 1999), as in continental Europe (Damsholt, 2002), but all of about 15 British records made since 1990 are from metalliferous mine sites in Cornwall, often from ground conspicuously contaminated with copper (Holyoak, 1997, 2000, and unpublished). Cephaloziella massalongi N. KERRY (H2): part-shaded soil on slope beneath saplings at edge of old copper mine, ca 20 m alt., Ross Island, V946880, 13 May 2005, DTH 05119 (BBSUK, DBN) (Blackstock, 2006); on N.-facing bank of top part of old lead-mine shaft, on rock (carbonate?) and thin overlying soil, lightly shaded by Betula saplings, ca 25 m alt., N. shore of Muckross Lake, V948859, 24 April 2008, DTH 08-162 (DBN). W. CORK (H3): on damp patch of open humic soil amongst gravelly copper mine-spoil near cliff top, ca 30 m alt., coast NW. of Allihies, V577459, 1 June 2006, DTH 06-108 (DBN). WICKLOW (H20): partshaded crevices of NW.-facing stone wall below ‘buddle’ at old lead mine, ca 285 m alt., Vale of Glendasan (Old Hero Mine), T098981, 15 April 2008, DTH 08-100 (BBSUK, DBN). DUBLIN (H21): partly bare soil among low rocks on slope of lead-mine spoil, slightly shaded by Agrostis, ca 215 m alt., Ballycorus disused lead mine, O225208, 13 April 2008, DTH 08-89 (BBSUK); also collected nearby from another small population (DBN). Previous records from Ireland are from two localities at old copper mines: Mountain Mine near Allihies (H3) and coastal sites E. of Bunmahon (H6) (Holyoak, 2003); strong populations persisted at both localities in 2007–2009. New Journal of Bryology (2009) 31: 267–282

Bruchia vogesiaca Schwägr. (Bruchiaceae) on Bodmin Moor, East Cornwall: a moss new to Britain

On 13 July 2006 four unfamiliar but striking moss capsules were noticed while I was examining bryophytes on the bank of a small river on Bodmin Moor, East Cornwall, U.K. (v.-c. 2; National Grid square SX17). A small specimen was collected (DTH 06-231, part of which will be lodged in BBSUK) and the living plants were photographed. Microscopic study later the same day revealed a close match with illustrations and descriptions of Bruchia vogesiaca (e.g. in Crum & Anderson, 1981) and this was confirmed by comparisons with old specimens from France in the author’s herbarium. Distinctive features noted on the Bodmin Moor plants include the sporophyte 6–7 mm long overall, suberect; seta about 4 mm long, pale, flexuose; capsule cylindrical with conspicuous neck as long as the theca; operculum rostrate; calyptra mitrate covering only the upper one-third of the capsule. The leaves were poorly preserved and they were not examined in detail to avoid damaging the tiny voucher specimen, but they were seen to be narrow with a stout excurrent costa. The Bodmin Moor plants lacked mature spores so that separation from additional species of Bruchia known in North America relied on other characters, especially the exserted sporophytes resulting from the length of the seta greatly exceeding 2 mm (cf. Crum & Anderson, 1981). Fuller descriptions of the species are given by Limpricht (1891), Crum & Anderson (1981), Rushing (1986) and Sérgio, Jansen & Séneca (1998). The last of these papers shows that the species has a persistent protonema that may produce uniseriate gemmae. The locality was revisited on 28 July 2006 in order to search for more B. vogesiaca and to record details of the habitat and associated plants. Despite an extensive search, only seven more capsules were found, all within about 1.5 m of the original discovery. Because of its apparent rarity the precise location is not being published, although Natural England has full details. The plants grow on steep, damp, gritty but humic soil of a near-vertical bank about 1 m high beside a small river. The site is fully exposed to sunlight for part of each day. The river runs through an area of grassland closely grazed by cattle, with a few patches of Ulex europaeus L. and other scrub. The vegetation accompanying B. vogesiaca formed an incomplete, low cover, composed mainly of Dicranella rufescens (Dicks.) Schimp., with smaller amounts of Jungermannia gracillima Sm., Atrichum tenellum (Röhl.) Bruch & Schimp., Pellia sp. (immature thalli) and rare stems of Pohlia camptotrachela (Renauld & Cardot) Broth. Other bryophytes within 1 m were Calypogeia arguta Mont. & Nees, Cephalozia bicuspidata (L.) Dumort., Fissidens cf. curnovii Mitt., Pellia epiphylla (L.) Corda and Philonotis sp. (immature stems); phanerogams close by (mainly on top of the bank) included Agrostis capillaris L., Danthonia decumbens (L.) DC., Euphrasia nemorosa (Pers.) Wallr., Juncus bulbosus L., Plantago lanceolata L., Potentilla erecta (L.) Raeusch., Prunella vulgaris L. and Wahlenbergia hederacea (L.) Rchb. All of the associated plants are widespread on Bodmin Moor and the assemblage, as a whole, points to acidic soil conditions. The steep low river banks in the area result from fluvial erosion and they are sometimes submerged completely when the river overflows. Considerable erosion of the banks in the vicinity is being caused by cattle. Descriptions of the habitat in continental Europe also refer to open, damp, acidic substrata and to disturbance resulting from trampling by cattle (Sérgio et al., 1998; Hugonnot & Ully, 2003). The ecological requirements and habitat types occupied by French populations are described in detail at http:/www.ecologie.gouv.fr/IMG/natura2000/ habitats/pdf/tome6/1385.pdf In Europe B. vogesiaca has a restricted range that extends in a band from Austria through southern Germany, France and Spain to central Portugal. Bonnot (1967, 1968) and Sérgio et al. (1998) have mapped its localities, the latter authors reporting it from only 17 of the 50650 km U.T.M. grid squares in Europe. There is only one record each from Austria and Germany, both more than 100 years old. However, there has been an increase in the number of records from France in recent decades from both the Massif Central, where Hugonnot & Ully (2003) report two sites in Département du Cantal, and from the Vosges (Frahm, 2005). There are also three recent new records in northern Portugal (Garcia & Sérgio, 2004). The Red Data Book of European Bryophytes (Schumacker & Martiny, 1995) treats B. vogesiaca as ‘Endangered’. In recognition of its rarity and vulnerability it is included on Appendix I of the Bern Convention and on Annex 2 of the EC Habitats and Species Directive. The species is rare elsewhere, with records in North America from New York State, Maine and New Hampshire (Crum & Anderson, 1981; Rushing, 1986; Allen, 1992) and a single record in eastern China (Gao, Crosby & Si, 1999). Fortunately, the locality at which B. vogesiaca occurs on Bodmin Moor lies within the extensive Bodmin Moor, North Site of Special Scientific Interest. To judge from Journal of Bryology (2007) 29: 135–136

Notes on two nominal taxa of Ephemerum (Bryophyta: Pottiaceae) described from France by Charles Douin

Charles Isidore Douin (b. 1858, d. 1944) made careful studies of Ephemerum from northern France and published detailed observations that are of lasting value (Douin, 1906, 1907, 1912). A recent review of several European species of Ephemerum separated E. minutissimum Lindb. at species rank from E. serratum (Hedw.) Hampe, mainly on the basis of characters of the mature spores (as previously advocated by Risse, 1996, 1997, and followed by several authors); it also synonymised E. stellatum H.Philib. with E. serratum s. str. (Holyoak, 2010). It was noted that two other names proposed for E. stellatum by Douin remained of uncertain identity as synonyms of either E. serratum s. str. or E. minutissimum because the types had not been found. These types have since been located at the National Botanical Garden of Belgium (BR), which houses Douin’s original herbarium. This note describes results of study of these type specimens. Ephemerum serratum var. subintegrum [-a] Douin, Mém. Soc. Sci. Nat. Cherbourg 35: 285. 1906 [not 1900 as given by Holyoak, 2010, p. 127]. The protologue gave very little information on this taxon, merely stating: ‘Dans les allées du bois de Dangeau s’en voit une autre à feuilles petites et presque entières (var. subintegra Douin in herb.)’. Later, Douin (1907, p. 243) noted that E. stellatum was collected ‘c. fr. dès 1901 dans le bois de Dangeau; mais l’absence du fameux caractère des feuilles étalés en étoile et divergentes m’avait toujours empêché de l’appeler ainsi: de là nom de E. serratum var. subintegra Douin que je luis avais donné’ (a numbered footnote refers to the protologue of the latter name). Hence, it is clear that Douin (1907) reinterpreted the name as that of a synonym or form of E. stellatum rather than of E. serratum. BR-BRYO 361890-80 from Douin’s herbarium apparently represents the holotype of this taxon, although it is not labelled as a type. The small green packet bears the original data: ‘Eph. serratum v subintegra//petit Bois jus Dangeau//9.11.00 [or 01]’ (illegible handwriting is underlined). The packet contains flakes of silty soil, some of which have crumbled to a powder, with scattered immature plants of the ecostate E. serratum s. str. or E. minutissimum. Three plants without capsules that were separated had most of their leaves bearing rather small teeth, a few also had a single large tooth and one small leaf was edentate. Two larger plants with immature capsules were also studied, one of them having long narrow ecostate leaves with very large teeth. The lack of any mature spores in the entire gathering prevents reidentification of the taxon as either E. serratum s. str. or E. minutissimum, although it clearly represents one or the other of these. Ephemerum stellatum var. denticulatum Douin, Bull. Soc. Bot. France 59: 734 [name only], 735. 1912. The protologue stated only ‘j’ai récolté dernierment dans le bois de Dionval près SaintPiat (E.-et-L.) une forme à feuilles nettement denticulées dans la moitié supérieure au moins (var. denticulatum in herb. Douin)’. This name was overlooked by the compilers of Index Muscorum (van der Wijk et al., 1962, 1967). BR-BRYO 010879-15 from Douin’s herbarium apparently represents the holotype of this taxon, although it is not labelled as a type. The outer packet bears the original data: ‘xx Ephemerum stellatum Phil var. denticulatum D//Chemin du Bois au dessus de Dionval près St Piat (E.&L.)//26.10.1912 France’. The protologue was published in the proceedings of the ‘Séance du 13 Décembre 1912’ according to the running heading, so the date of collection on 26 October 1912 on the specimen suggests that the ‘récolté dernierment’ [collected lately, or not long ago] was literally true. The smaller of two main inner packets is labelled ‘beau’ and contains soil flakes with numerous Ephemerum plants. Two plants from it with relatively large brown capsules were removed for study; both have ecostate leaves that are denticulate to weakly toothed; the first plant has apparently immature spores (41 mm long) but they lack a ‘veil’ and are rather strongly Correspondence to: David Thomas Holyoak, Quinta da Cachopa, Barcoila, 6100-014 Cabeçudo, Portugal. Email: holyoak9187@hotmail.com Bryological notes

Genetic variation in the Red-Listed moss Ditrichum cornubicum Paton (Ditrichaceae) and implications for its conservation

Ditrichum cornubicum is a rare and threatened acrocarpous moss found on copper-rich mine waste, characteristic of the EU protected habitat ‘Calaminarian grassland of the Violetalia calaminariae’. The species was once thought to be a British endemic, being known from two former copper/tin mine sites in Cornwall, until the discovery of a population in 2006 at Allihies Mountain Mine, Co. Cork, Ireland, a former copper mine. In light of this discovery, two theories of possible introduction from Britain to Ireland were put forward: (1) an introduction from the 1800s and (2) a more recent introduction (2000s). Only male plants of the species are known, and reproduction and dispersal are therefore solely through asexual propagules and fragmentation. In order to address the conservation questions of the origin of the Irish population and to determine whether diversity exists within this rare species, genetic fingerprinting (amplified fragment length polymorphism) was carried out on the three known global populations. Percent polymorphism was found to be 7.29% and Neis total gene diversity (HT) was 0.0356. AMOVA revealed that, of the small amount of variation found, the majority was among the three populations (98%). The presence of five private alleles in the Allihies population suggests a longer period of isolation than would fit either theory of introduction. Cluster analyses reveal that the Allihies population is more genetically distinct than the two British populations are to each other, thus heightening the conservation priority for this population.

Bryum austriacum Köckinger, Holyoak & Suanjak, a new bulbiliferous species from the Alps (Bryaceae, Bryopsida)

Abstract Bryum austriacum Köckinger, Holyoak & Suanjak, sp. nov. is described from the eastern Alps. It is a small, delicate, pale-coloured moss, with mainly imbricate and concave leaves, very large, and often thin-walled mid-leaf cells and abundant dark red-brown ovoid bulbils occurring singly in each leaf axil. The new species is currently known from 19 localities in Austria and one in S. Germany. It is almost exclusively restricted to sites above the tree-line, growing as a pioneer mainly on humus- and base-rich soil in gaps of alpine meadows or among subnival cushion vegetation.

Bryum apiculatum (Bryaceae) in East Cornwall: a moss new to Europe

In January 2007 an unfamiliar small Bryum was found at the western edge of Retire Common, East Cornwall, UK (v.-c. 2; National Grid reference SW997631). It attracted attention because clusters of rounded brown gemmae lacking leaf primordia were present in the upper leaf axils (Fig. 1). Specimens collected on 28 and 30 January and 24 April 2007 have since been identified as B. apiculatum Schwägr. (Holyoak 07-09, 07-15 and 07-110, parts of which are lodged in BBSUK), a species hitherto unknown in Europe. The species is widespread in tropical and subtropical regions of the world, often growing as a weed colonising disturbed soil. In Cornwall it was established at the edge of heathland on a small car-parking area and in a cattle grid along a track. Identification characters of this notoriously variable moss are discussed.