BBS Autumn Meeting: Excursion

Home , Marsupella profunda, Metzgeria furcata, Scapania paludicola

MeetingReport Meeting Report – Sussex 2009

birthday. The President conveyed the Society’s congratulations and presented her with a bouquet. The conversazione followed the dinner for which Jean Paton displayed a collection of memorabilia illustrating aspects of her career, and five members provided posters. On Sunday we visited the Francis Rose reserve at Wakehurst Place as guests of the Royal Botanic Gardens Kew.

v Wakehurst Place, Sussex, site of the Sunday n Jonathan Sleath presents Jean Paton with a bouquet BBS Autumn Meeting: excursion. Ian Atherton at the dinner held in her honour. Ian Atherton

University of Sussex,11–13 September 2009 ABSTRACTS OF TALKS David Streeter reports on an excellent Autumn meeting, The distribution of British and Irish liverworts: no species moved. The 10 clusters were named a new analysis – Chris Preston, Colin Harrower & after the species with the best fit. There was one held in honour of Jean Paton’s 80th birthday. Mark Hill cluster of 32 widespread species characterized by Metzgeria furcata, and five clusters which formed a The last classification of the distribution of all he 2009 Annual Meeting was held at x Jeff Duckett series increasingly restricted to the north and west, the British and Irish liverwort species was an the University of Sussex from 11 to 13 The function and evolution of stomata in named after Diplophyllum albicans (40 species). association-analysis published by Michael Proctor September. The occasion was special bryophytes (p. 38) Saccogyna viticulosa (19 species), Marsupella in 1967. This was based on the vice-county records in that the opportunity was taken to emarginata (37 species), Harpalejeunea molleri x Gordon Rothero collated by Jean Paton for the fourth edition of the celebrate the 80th birthday of Jean (30 species) and Bazzania tricrenata (21 species). Feeling the heat? Monitoring change in snowbed Census Catalogue of British Hepatics (1965). The TPaton MBE with a specially designed programme The remaining four groups were those named bryophyte communities in Scotland (p. 40) new analysis presented in this talk was based on of papers and a conference dinner. Altogether 56 after Cephaloziella stellulifera (27 south-western x David Holyoak the 10-km square records held in the BBS database members and guests attended the conference, species), Odontoschisma sphagni (22 species from Marsupella profunda: conservation problems with at BRC. A clustering algorithm was initially used to and it was especially good to welcome Barbara bogs), Preissia quadrata (17 calcicoles of the north a colonist (p. 41) group the 300 species into 10 clusters, the number Crandall-Stotler and Raymond Stotler from South- and west) and Anthelia juratzkana (55 rare montane of clusters being specified at the outset. The ern Illinois University, and Herman Stieperaere from x Barbara Crandall-Stotler species). The Cephaloziella stellulifera group held a similarity of each species to the overall distribution the Nationale Plantentuin van België. Melding molecules and morphology in liverwort high proportion of threatened species, followed by of those in each of the 10 clusters was then tested Publications Committee, Conservation and phylogeny (p. 43) the Anthelia juratzkana group. and species moved if they were more similar to Recording Committee and Council all met on x David Long another cluster than to the one in which they were Chris Preston, CEH Wallingford, Benson Lane, Friday evening with the main programme taking British liverwort floras 1741–1999: from Johann J. currently placed. This testing was repeated until Crowmarsh Gifford, OX10 8BB (e [email protected]) place on Saturday morning and afternoon. The full Dillenius to Jean Paton (p. 44) programme of papers was as follows: x Mark Hill Priorities for bryophyte conservation – Unfortunately, this is the very habitat that supports x Chris Preston, Colin Harrower & Mark Hill Sixty years a queen; vice-county recording since a Scottish perspective – Dave Genney some of our most important oceanic bryophyte The distribution of British and Irish liverworts: a 1947, with observations on where Jean Paton communities. The dense shade under its canopy new analysis (p. 35) went and what she found (p. 46) Bryophyte conservation is a complex area of which the following are just some of the more important kills all but the most shade tolerant species and can x Dave Genney The AGM followed at 1600 and the President’s issues and solutions. devastate a local flora. Control of Rhododendron is Priorities for bryophyte conservation – a Scottish report was published in Field Bryology vol. 100, expensive and requires a number of follow-up visits perspective (p. 35) pp. 66–67. The AGM was followed by a meeting THREATS to be successful. It is important to prioritize control x Sam Bosanquet of the Tropical Bryology Group. The Conference Rhododendron ponticum is a non-native invasive at sites where their impact on native species and Peatland bryophytes in Wales (p. 37) Dinner was the formal celebration of Jean Paton’s shrub that thrives in Scotland’s oceanic woodlands. habitats is greatest.

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v Wakehurst Place, Sussex, site of the Sunday n Jonathan Sleath presents Jean Paton with a bouquet BBS Autumn Meeting: excursion. Ian Atherton at the dinner held in her honour. Ian Atherton

34 FieldBryology No101 | May10 FieldBryology No101 | May10 35 Meeting Report – Sussex 2009 Meeting Report – Sussex 2009

Over- or under-grazing by wild deer or domestic particular range of woodland canopy cover. Direct and identification guides for important habitats. expertise, and nurture the next generation of herbivores can have major implications for wood- monitoring of rare and typical bryophytes is also The project has helped to introduce more people bryologists, mosses and liverworts will be in safest land bryophytes. Over-grazing for long periods carried out. to bryophytes by producing a Wild and Wonderful possible hands. can mean that old trees are not replaced and the World of Scottish Mosses and Liverworts leaflet, Dave Genney, Scottish Natural Heritage canopy becomes too open. Under-grazing can organizing public walks and training rangers to give DATA (e [email protected]) be equally damaging and see bryophytes shaded their own walks and talks. REFERENCES out by dense sapling regeneration or increased Bryophyte distribution data are critical for effective Although this paper is a very brief overview of bracken and bramble cover. Striking the balance bryophyte conservation. It allows better planning bryophyte conservation priorities, it should be Genney, D. (2010). Opinion. Field Bryology 100, 79–80. decisions, informs policy, allows conservation sta- between over- and under-grazing can be difficult to clear how important the BBS is to bryophyte Hodgetts, N. (2009). Conservation News. Field Bryology 97, achieve, and requires coordination and cooperation tus assessments, and can also stimulate interest in conservation. By continuing to collect data, provide 35. between graziers, dear estate managers, foresters bryophytes for many people. Species distribution and Scottish Natural Heritage (SNH). data can come from commissioned surveys, but by far the most important source of data is from Peatland bryophytes in Wales – Sam Bosanquet Cardiganshire site, a peat core revealed basiphilous Climate change has already been discussed as a BBS members. The Threatened Bryophyte Data- bryophytes such as Climacium dendroides and serious threat (Genney, 2010). The Countryside Council for Wales (CCW) is in the base is an invaluable tool for conservation fourth year of a 6-year study of the vegetation, Scorpidium cossonii 35 cm below the current Although the patchwork of mixed-age heather that (Hodgetts, 2009). With widely available data, ecology and condition of Welsh lowland peatlands. vegetation surface, whereas Tomentypnum is results from well-managed muirburn can benefit there is less chance that important species are These are highly varied, ranging from tiny basin now intermixed with Sphagnum squarrosum. many species, burning of hepatic heath can be accidentally damaged. Availability of BBS records mires less than 1 ha in extent, but with exceptionally This indicates relatively recent acidification of the disastrous for internationally important bryophyte on the National Biodiversity Network’s Gateway complex patterning, to vast fen/swamp complexes surface, which is probably an additional threat to communities. In Scotland, muirburn is guided by provides the greatest chance that bryophytes are near the Welsh coast covering 200 ha or more, and this rare moss. the Muirburn Code which promotes avoidance of taken into account and conserved effectively. including swathes of species-poor reedbed. The Bog bryophyte diversity could provide a proxy for sensitive habitats. base-poor Sphagnum mires of mid-Wales contrast bog condition. Welsh peatland bryophytes have with the tufa-forming rich-fens of Anglesey. 191 IMAGE been assigned scores according to a subjective PROTECTED SITES sites have been covered to date, some during the assessment of their restriction to intact raised bogs, Effective bryophyte conservation needs a good PR preceding CCW lowland grassland survey; at least and these scores have been used to rank sites. Many of the most important Scottish sites for campaign. Bryologists must continue to promote 76 remain unsurveyed. The least disturbed Welsh bogs, especially Cors bryophytes are given legal protection though a their importance and beauty so that as many Site floristics and condition are the main focuses of Fochno, have the most consistently high scores for series of protected sites. Sites of Special Scientific people as possible value them. In an ideal world, the survey, requiring detailed mapping. Bryophytes their bog bryophytes. Regenerating peat cuttings Interest (SSSIs) are the building blocks of Scotland’s bryophytes would receive the same level of public are key components of the vegetation and are with Sphagnum recurvum s.l. fen (NVC M4 & M5) protected areas, and just over 1,450 SSSIs form support as birds! However, bryologists can still have therefore studied in reasonable detail. Observations tend to be poor for liverworts, whereas established part of the wider GB series. Of these, bryophytes a big impact, e.g. by raising awareness of the vital include: bog with Sphagnum papillosum is usually richer. are a recognized feature of 65 Scottish sites. The ecosystem services that bryophytes perform, such Liverworts act as a convincing proxy for overall bog important features of each SSSI are described in as flood prevention and as a global carbon sink. Sphagnum magellanicum and S. balticum bryophyte diversity and therefore condition. a citation, which also lists a range of activities that on cut peat surfaces less than 80 years old, can only be carried out with consent from SNH. The suggesting colonization from unknown sources. Scapania paludicola is widespread in mid- and EDUCATION AND TRAINING management requirements for each site are set out S. magellanicum remains widespread but scarce north Wales and is particularly frequent in neutral in a statement that SNH uses to maintain sites in a Bryophyte habitat and management requirements in Wales but seldom produces sporophytes, whilst mires alongside Sphagnum inundatum and healthy condition through close working with their also need to be communicated effectively. This has S. balticum is only known to be extant at a single Sarmentypnum exannulatum. owners. A 6-year cycle of Site Condition Monitoring been advanced over the past 3 years by Plantlife site. The ability of these species to colonize peat Amblystegium radicale grows in humid reedbeds (SCM) determines whether bryophyte features Scotland’s Lower Plants and Fungi Project. This cuttings is encouraging for bog regeneration. in south-west Wales, especially when Riccardia are likely to be maintained in the medium to longer project has provided targeted training events for Tomentypnum nitens has been lost from at least chamedryfolia is abundant. term. To produce this assessment, a set of import- Scottish conservation agency staff on, for example, three of its 10 Welsh sites and has declined at most ant bryophyte habitat attributes are assessed dead wood management. It has also produced a of the others because of scrub encroachment and Sam Bosanquet, Cnwc y Llwyn, Brechfa, Carms. against targets; for example, there should be a range of booklets to promote good management a lack of grazing management. At this species’ only SA32 7QR (e [email protected])

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(1a) (2a)

The function and evolution of stomata in bryophytes – Jeff Duckett, Ken P’ng, Karen Renzaglia & Silvia Pressel

It was most fitting that this celebration of Jean understanding of stomatal evolution and function in v Fig. 1. Cryo-SEM images, surface views. (a, b) Paton’s 80th birthday should include a presentation bryophytes. On the one hand, genomic data-mining Paraphymatoceros minutus. on stomata. Two of Jean’s most seminal early works should reveal whether or not bryophytes possess (1b) (a) Closed stoma from inside were on stomata in British bryophytes (Paton, 1957; the same stomatal genes as higher plants and, an involucre; (b) newly opened Paton & Pearce, 1957); 52 years later these remain on the other, cryo-scanning electron microscopy stoma immediately above indispensably authoritative accounts. Apart from a (cryo-SEM) coupled with ion milling and X-ray an involucre. (c) Phaeoceros flurry of excellent cytological and functional studies microanalysis (enabling direct detection of possible (2b) laevis, open stoma flanked – Sack & Paolillo (1973) and Garner & Paolillo (1973) potassium fluxes between guard cells and epi- by desiccated and shrunken showed that the stomata of Funaria are able to open dermal cells like those in higher plants), can now be epidermal cells well above and close, at least when young and that, like those used for incisive functional studies. the dehiscence point on a in higher plants, they are sensitive to abscissic acid; Our current research using cryo-SEM is pro- sporophyte. Bars, 20 μm. while Maier’s (1974) and Boudier’s (1988) findings ducing novel functional data that lead to exciting, v Fig. 2. Cryo-SEM images, that the stomata of Sphagnum lack subsomatal conflicting and challenging hypotheses awaiting the cryo-fractured preparations. air spaces lead to the conclusion that they should application of genomic technologies. In Sphagnum, (a, b) Anthoceros agrestis. (a) Sporophyte within an best be considered as pseudostomata (Cox et the role of the numerous pseudostomata is (1c) involucre with mucilage-filled al., 2004) – the last half century saw virtually no facilitation of sporophyte desiccation (Duckett et al., intercellular spaces (arrowed) new advances in our understanding of bryophyte 2009; Beerling & Franks, 2009). A detailed study (2c) in the assimilatory tissues; (b) stomata. This was almost certainly due to a lack across hornworts has revealed that stomata open gas-filled intercellular spaces of suitable experimental technologies; thus, Jean’s after the sporophytes emerge from the involucres, (arrowed) above an involucre; decision to leave stomata and thereafter pursue thereafter remaining in this condition and being spores and pseudoelaters liverworts was a very wise move. unaffected by darkness and desiccation (Fig. 1a–c); embedded in mucilage (S). (c) It is widely assumed that stomata, one of the key there is no evidence of potassium fluxes between Podocarpus nivalis, gas-filled innovations in land plants, are homologous across the guard cells and surrounding cells. An even more intercellular spaces in a very land plants (Raven, 2002). This hypothesis fits surprising discovery is that the intercellular spaces young leaf before stomatal ontogeny. Bars, (a, c) 20 μm, neatly with 21st century total evidence phylogenies in the outer assimilatory layers in the sporophytes (b) 50 μm. that place liverworts firmly at the base of the land are initially mucilage-filled (Fig. 2a) (as is always the plant tree (Shaw & Renzaglia, 2004; Renzaglia et case in hornwort gametophytes) and that water finding that, in most species, a possible closing Jeff Duckett & Silvia Pressel, School of Biological al., 2007). Accordingly, stomata first evolved in evaporates producing air-filled spaces only after mechanism is absent. and Chemical Sciences, Queen Mary University mosses (± Sphagnum) and are thereafter present the stomata open (Fig. 2b). The same observations Thus we present three novel, challenging and of London E1 4NS, UK; Ken P’ng, Department of from polytrichalean mosses and hornworts into on the leaves of vascular plants show that here, in conflicting questions for future research. (1) Did Materials, Queen Mary University of London; Karen higher plants. Embedded in this notion of stomatal dramatic contrast, the intercellular spaces are gas- stomata first appear in bryophytes as structures Renzaglia, Plant Biology Department, Southern homology is a further assumption that their principal filled from the outset and long before the stomata that facilitated sporophyte desiccation and spore Illinois University, Carbondale, IL 62901, USA function (and the selection pressure driving the open (Fig. 2c). We therefore conclude that the dispersal, and was their role in gaseous regulation (e [email protected]; [email protected]; same) is regulation of gaseous exchange (Raven, principal role of hornwort stomata is facilitation of in vascular plants a later secondary acquisition? (2) [email protected]; [email protected]) 2002), though this is certainly not the case in sporophyte desiccation, as previously suggested Did stomata in vascular plants evolve independently REFERENCES Sphagnum where the pseudostomata are covered by Lucas & Renzaglia (2002). This very different of those in bryophytes? (3) Or did stomata first Beerling, D.J. & Franks, P.J. (2009). Evolution of stomatal with the calyptra until the sporophytes are almost function, taken together with the strikingly different evolve in early land plants as sites of gaseous function in ‘lower’ land plants. New Phytologist 183, mature (Cox et al., 2004; Duckett & Ligrone, 2004; origins of the intercellular spaces, suggests that exchange and were subsequently co-opted in 921–925. Duckett et al., 2009). hornwort stomata are not the same as those in moss and hornwort sporophytes as structures Boudier, P. (1988). Differenciation structurale de l’eiperme du However, with new tools at hand we now stand higher plants. Studies in progress on mosses are facilitating drying out, capsule dehiscence and sporogone chez Sphagnum fimbriatum Wilson. Annales at the break of a new dawn for major advances in yielding similar results together with the additional spore discharge? des Sciences Naturelles Botanique, Paris 8, 143–156.

38 FieldBryology No101 | May10 FieldBryology No101 | May10 39 Meeting Report – Sussex 2009 Meeting Report – Sussex 2009

(1a) (2a)

The function and evolution of stomata in bryophytes – Jeff Duckett, Ken P’ng, Karen Renzaglia & Silvia Pressel

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Cox, C.J., Goffinet, B., Shaw, A.J. & Boles, S.B. (2004). Paton, J.A. (1957). The occurrence, structure and functions This project, funded by Scottish Natural Heritage can only provide information on changes within the Phylogenetic relationships among the mosses based on of the stomata in British bryophytes. I. Occurrence and and carried out in cooperation with the University of community, not on spatial changes. The results here heterogeneous Bayesian analysis of multiple genes from structure. Transactions of the British Bryological Society Bergen and the Royal Botanic Gardens Edinburgh indicate that the bryophyte communities are much 3, 228–242. multiple genomic compartments. Systematic Botany 29, was designed to do two things. The first was to the same now as in 1989–1990, but with some 234–250. Paton, J.A. & Pearce, J.V. (1957). The occurrence, structure revisit most of the snowbed sites that I visited in 1989 significant increase in the frequency and cover of and functions of the stomata in British bryophytes. II. Duckett, J. G. & Ligrone, R. (2004). There are many ways of and 1990, and resample the snowbed communities some vascular plants. Comparison of photos taken Functions and physiology. Transactions of the British making water-conducting cells but what about stomata? I studied then, in an attempt to discern any change in 1989 with those from 2007 also provides evidence Field Bryology 82, 32–33. Bryological Society 3, 242–259. within the communities. The second was to set up of some marked changes, particularly in the cover Raven, J.A. (2002). Selection pressures on stomatal Duckett, J. G., Pressel, S., P’ng K. & Renzaglia, K.S. (2009). evolution. New Phytologist 153, 371–386. a series of baseline transects across a number of of vascular plants like Juncus trifidus, Deschampsia Exploding a myth: the capsule dehiscence mechanism snowbeds plus taking a series of site photographs flexuosa, D. cespitosa and Nardus stricta. and the function of pseudostomata in Sphagnum. New Renzaglia, K.S., Duff, R.J., Ligrone, R., Shaw, J., Mishler, so that any future change can be monitored. It Setting up of the baseline transects was a time- Phytologist 183, 1053–1063. B.D. & Duckett, J.G. (2007). Bryophyte phylogeny: advancing the molecular and morphological frontiers. was planned that sites in the Cairngorms would be consuming exercise, trying to select a suitable line Garner, D.L.B. & Paolillo, D.J. (1973). On the functioning of Bryologist 110, 179–213. covered in 2007 and a selection of sites elsewhere with good natural markers at each end and trying stomates in Funaria. Bryologist 76, 423–427. Sack, F. & Paolillo, D.J. (1973). Structure and development in the Highlands surveyed in 2008. This was broadly to snatch location photos in the spells when the Lucas, J.R. & Renzaglia, K.S. (2002). Structure and function of walls in Funaria stomata. American Journal of Botany successful, although the weather was not kind and clouds lifted. These are dynamic sites, but it is of hornwort stomata. Microscopy and Microanalysis 8, 70, 1019–1030. two further transects were not completed until hoped that the large boulders selected as markers Supplement 2, 1090–1091. Shaw, A.J. & Renzaglia, K.S. (2004). Phylogeny and 2009 (thanks to funding from the BBS). By the end will remain for some time. If possible, a selection of Maier, K. (1974). Ruptur der Kapselwand bei Sphagnum. diversification of bryophytes. American Journal of Botany of the field work some 20 transects had been set these transects will be revisited in 5 years or so, to Plant Systematics and Evolution 123, 13–24. 91, 1557–1581. up involving almost 500 quadrats and 115 of the test out how viable relocating the line is, although 1989–90 plots had been resampled. Most of the funding is always going to be a problem. Another sites involved are well above 1,000 m and not close fundamental problem may be finding the right Feeling the heat? Monitoring change in to the road, so this involved some very long days, combination of skills in a surveyor, as it helps to snowbed bryophyte communities in Scotland – but fortunately in wonderful places. have both a fondness for being alone in the big hills Gordon Rothero It is important to stress that the resampling did and a moderate obsession with Marsupella! not involve fixed plots; quadrat data were gathered When snow falls it rarely stays in the same place in similar stands on the same sites in both periods, Gordon Rothero, Stronlonag, Glenmassan, By for long, and this is particularly true of snow in the but not from exactly the same places. So, the data Dunoon, Argyll PA23 8RA (e [email protected]) Scottish hills. Here the wind blows the snow onto lee slopes where it can accumulate to prodigious depths. As the prevailing winds are from the south Marsupella profunda: conservation problems occurring in Britain; (d) in Biodiversity: The UK and west, the snow tends to accumulate on slopes with a colonist – David Holyoak Steering Group Report (1995) it was placed on with north and north-easterly aspects, slopes which the ‘Short List of Globally Threatened/Declining also receive limited insolation. On the bigger hills the The small liverwort Marsupella profunda Lindb. was Species’, classed as Critically Endangered in snow patches persist right into the summer, leaving added to the British flora by Paton (1990), who Britain and as a ‘Species of global conservation a short growing season, limiting the growth of higher recognized it amongst specimens of M. sprucei concern’, and an Action Plan for its conservation plants and favouring vegetation dominated by lower (Limpr.) H.Bern. she had collected in West Cornwall was presented. Most of these conservation plants. However, winters in Scotland have been between 1965 and 1971. designations were established on the assumption noticeably less snowy since at least 1980 and snow Because of the restricted range of M. profunda that its true status was already known, i.e. prior to patches have melted out earlier; the presumption and its apparent rarity throughout Europe, it is: (a) any really extensive targeted surveys being made of must be that this is having some effect on snowbed listed on Schedule 8 of the Wildlife and Countryside its status in the UK or elsewhere. vegetation. Act (as amended); (b) listed on Appendix 1 of the In 1990, M. profunda was not known to survive Bern Convention; (c) listed on Annex II of the EC at any of its Cornish sites, which had become v Ciste Mhearad, Cairngorm, a classic snowbed site. Habitats and Species Directive (Council Directive overgrown with gorse or bramble or been destroyed, Gordon Rothero 92/43/EEC); it is the only Annex II ‘priority’ species although Jean Paton suggested that ‘it must surely

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(1) Instances of accidental damage on SSSIs habitat management activity, but it can also be x Employees of Goonvean Ltd has exposed the companies to potential risk of counter-productive (as in East Cornwall). Quarrying moving a large boulder with M. prosecution. by the China-clay companies provides most of the profunda growing on it (February (2) Discovery of M. profunda in large quantities new habitat for the species in East Cornwall. Hence, 2007). The boulder was on a site on the route for a new road at Greensplat led to a partnership with the clay companies should be being used as a spoil tip for china- extensive translocation (under licence) of rocks with developed so they have an interest in protecting M. clay waste at Trelavour Downs, East Cornwall. The translocation patches of the plant. profunda rather than in seeking to prevent it being was carried out under licence from (3) Discovery of M. profunda on the location for a surveyed on their land. Natural England. David Holyoak new quarry spoil tip led to last-minute translocation of boulders with patches of the plant (under licence) David T. Holyoak, Quinta da Cachopa, Barcoila, still be around somewhere’. A search at the same Tregonning Hill SSSI the largest population seen to the edge of a neighbouring SSSI. 6100-014 Cabeçudo, Portugal (e holyoak9187@ Cornish sites in early 1993 by Nick Hodgetts, was of just 15 cm2 on a single boulder in 1998. In (4) M. profunda currently occurs on the site of a hotmail.com) Jean and Ron Porley resulted in the species being 1999, when the site was proposed as a candidate planned ‘eco-village’ of 1,000 new homes. REFERENCES refound in very small quantity at Tregonning Hill, but SAC, there was even less of it (1.5 cm2), and by In conclusion, discovery of additional large UK Steering Group (1995). Biodiversity: The UK Steering nowhere else. 2005 it had become extinct at the site because of populations of M. profunda in Cornwall suggests Group Report, Vol. 2: Action Plans. London: HMSO. A field study of M. profunda in Cornwall during encroaching brambles and no suitable unshaded that it is now much less immediately threatened Holyoak, D.T. (1996). Status and conservation of the March–April 1996 was carried out under English habitat remains. globally than it appears to be on paper. Hence, liverwort Marsupella profunda in Cornwall, 1996. Pool, Nature’s Species Recovery Programme (Holyoak, At Lower Bostraze and Leswidden the extent of conservation efforts should focus on maintenance Redruth: Cornish Biological Records Unit (Institute of 1996). This study revealed that a tiny population still open habitat has declined progressively since 1996 or provision of newly bared habitat for the regional Cornish Studies). 30 pp. Unpublished. persisted on granitic rocks at Tregonning Hill, but as gorse and heather have colonized clay surfaces. metapopulations, with much less concern over Holyoak, D.T. (1999). Status, ecology and conservation found much larger populations at two ‘new’ sites Glyphosate herbicide has been used annually to individual sites that have small populations. of the liverwort Marsupella profunda in Cornwall, in West Penwith (Leswidden and Lower Bostraze) slow vegetation succession on the best areas. New Statutory protection of sites as SSSI and SAC 1999. Work carried out under English Nature Species where it mainly grew on clay-waste surfaces. Jean ground was cleared mechanically at both sites. has sometimes proved to be valuable (e.g. in West Recovery Programme contract no. F14-01-648. 37 pp. participated in the Leswidden find and checked all Intensive annual management has slowed declines Cornwall), since it can help to ensure appropriate + 8 pp. of photos. Unpublished. voucher specimens. Realization that the species is of M. profunda at both sites, but by 2009 a need an early-successional colonist on China-clay spoil for more drastic clearance work at Lower Bostraze led to further surveys for English Nature, resulting in was becoming apparent. Melding morphology and molecules in liverwort relationships within liverworts, such as Forrest et new finds of large populations at working clay pits The three SSSI sites in East Cornwall all showed phylogeny – Barbara Crandall-Stotler al. (2006) and the collaborative ‘Assembling the in East Cornwall (Holyoak, 1999). major declines by 2009, despite intermittent scrub Liverwort Tree of Life’ project (see http://bryophytes. Subsequent conservation actions on paper to clearance. However, adjacent unprotected areas Although many tools in the past have modified our plant.siu.edu/posters_ATOL.html), have challenged protect M. profunda in West Cornwall have included had gained much larger populations near two of the views of bryophyte phylogeny, none have impacted traditional, morphology-based concepts of most notification of Tregonning Hill as a SSSI (on 23 SSSIs. our systematic schemes and concepts of evolution taxonomic units and led to major modifications December 1994) and notification of Lower Bostraze The Second Report under Article 17 on the im- as much as those of molecular biology. The in liverwort classification. Our recently published and Leswidden as a SSSI (on 8 November 1996). plementation of the European Community Direct- mitochondrial intron classification scheme (Crandall-Stotler et al., 2009) In East Cornwall three sites (at Carclaze, Prosper Pit ive on the Conservation of Natural Habitats and of An intron is a region of data of Groth-Malonek attempts to integrate morphological and molecular and south of Treviscoe) were also notified as SSSIs Wild Fauna and Flora (92/43/EEC) (JNCC, 2007) DNA within a gene that is et al. (2005) and multi- data sets and reconcile their incongruence. Some and at the same time proposed as a candidate SAC was based largely on monitoring of the SAC sites not translated into protein. locus DNA sequencing of the unforeseen relationships now recognized in during 2000, despite strong recommendations to for M. profunda, so it was able to give only a gloomy studies of Qiu et al. the classification include the placement of Blasia avoid notifying sites at working clay pits (Holyoak, report on prospects. This emphasizes that even the (2006), for example, have unequivocally established at the base of the complex thalloid lineage, and 1999). Since 2000, the clay companies have mainly limited and artificial bureaucratic objective of report- that mosses, liverworts and hornworts comprise the placement of Monoclea and Riccia in derived refused access to their land in East Cornwall for any ing favourable conservation status in the UK was separate lineages, as sometimes argued (Crandall- lineages, close to Dumortiera and Conocephalum, wide-ranging surveys of the species. not being met by designating SACs for M. profunda. Stotler, 1986), and that liverworts are the first respectively. Conservation results on the ground have only M. profunda has also become a nuisance to the divergence of land plants. Studies of mature sporophytes of Phyllothallia partly justified the actions taken on paper. At clay companies: Phylogenetic studies focused on resolving show that it has several anatomical characters in

42 FieldBryology No101 | May10 FieldBryology No101 | May10 43 Meeting Report – Sussex 2009

42 FieldBryology No101 | May10 FieldBryology No101 | May10 43 Meeting Report – Sussex 2009 Meeting Report – Sussex 2009

common with elements of the Pallaviciniales where molecular markers to produce a meaningful, total who visited him in Oxford in 1736. His major work It took 58 years for the next liverwort flora to it is now resolved, rather than the Treubiales where evidence phylogeny. Historia Muscorum was published (in Latin) in appear, by Benjamin Carrington in 1874–1875. it had been placed, and new studies of Pleurozia Barbara Crandall-Stotler, Department of Plant 1741, but with a later English edition in 1768. It Although he had a drawing of Marchantia confirm that its elaborate water sacs are ventral, not Biology, Southern Illinois University, Carbondale, was a remarkable publication, both in its excellent polymorpha on the front cover, his work covered dorsal, and its strange tubes are unique structures, IL 62901-6509, USA (e [email protected]) illustrations, and its detailed Latin descriptions only 29 leafy liverworts as it was sadly abandoned not sterile perianths. (using polynomial names). Dillenius was a field due to his ill health. However, his name is REFERENCES In contrast to these examples in which related botanist and this shines through his descriptions. commemorated in some of his discoveries like Crandall-Stotler, B. (1986). Morphogenesis, develop- taxa have highly divergent morphologies, in the Dillenius’ friend Carl Linnaeus’ Species Plantarum Plagiochila carringtonii and Radula carringtonii. The mental anatomy and bryophyte phylogenetics: contra- Jungermanniidae morphological convergence indications of monophyly. Journal of Bryology 14, 1–23. was published 12 years later and is the starting next, Mordecai Cubitt Cooke was a remarkable often obscures underlying genetic divergence. point for liverwort nomenclature. He published the man, a polymath – he dabbled in many things and Crandall-Stotler, B., Stotler, R. E. & Long, D. G. (2009). For example, Jungermannia and Jamesoniella, Phylogeny and classification of the Marchantiophyta. first binomials for plants, including 44 species of was a prolific author. He was first and foremost an traditionally placed in the same family based on their Edinburgh Journal of Botany 66, 155–198. liverwort. Linnaeus’ liverworts were largely based outstanding mycologist based at Kew gardens and similar facies, are consistently resolved in different De Roo, R. T., Hedderson, T. A. & Söderström, L. on Dillenius’ work as described by Pekka Isoviita in was certainly not a hepaticologist; his biographer suborders in molecular studies. A similar scenario (2007). Molecular insights into the phylogeny of the 1970. Therefore Dillenius’ Historia Muscorum and John Ramsbottom of Kew commented that his occurs in analyses of Lophozia (De Roo et al., 2007), leafy liverwort family Lophoziaceae Cavers. Taxon 56, his herbarium are extremely important in correctly Hepatic Flora (1907) ‘did not add to his reputation’. which support its transfer to the Scapaniaceae, the 301–314. interpreting Linnaeus’ liverwort names, as in the In 1912, Cooke did an interview for the Morning recognition of several segregate genera, and the Forrest, L.L., Davis, E.C., Long, D.G., Crandall- case of Jungermannia undulata, now Scapania Post on his 87th birthday which led to erroneous transfer of Leiocolea to the Jungermanniaceae. Stotler, B.J., Clark, A. & Hollingsworth, M.L. (2006). undulata, first described by Dillenius but given a report of his death – a notice in the Kew Bulletin While such realignments may seem spurious, Unraveling the evolutionary history of the liverworts binomial by Linnaeus. stated that ‘he peacefully passed away at his critical re-evaluations of morphological features often (Marchantiophyta): multiple taxa, genomes and analysis. Our second hepaticologist is the Scotsman residence’; Cooke later said the report was ‘much Bryologist 109, 303–334. provide additional support for them. Thus, despite a James Dickson who worked as a nurseryman in exaggerated’! Groth-Malonek, M., Pruchner, D., Grewe, F. & Knoop, V. growing reliance on molecular tools to classify and/ London and became interested in bryophytes. William H. Pearson worked his whole life in the (2005). Ancestors of trans-splicing mitochondria introns or identify organisms, morphological studies must He made several field trips in Scotland and textile industry, but must have had a lot of spare support serial sister group relationships of hornworts still play a critical role in deciphering relationships and mosses with vascular plants. Molecular Biology and described his new discoveries in his 4-volume time as he published both his Hepaticae of the and documenting biodiversity. Intrinsic characters Evolution 22, 117–125. Fasciculus Plantarum Cryptogamicarum Britanniae British Islands in 1899–1902 and many papers on of spermatid architecture, sporeling and branch Qui, Y.-L., Li, L., Wang, B. & other authors (2006). (1785–1801). One example of his new species is liverworts from far-flung corners of the globe, such ontogeny, androecial and gynoecial organization, The deepest divergences in land plants inferred from Jungermannia adunca (now Herbertus aduncus) as New Zealand and New Caledonia, and was one capsule wall anatomy and sporogenesis are reliable phylogenomic evidence. Proceedings of the National described by Dickson from ‘Alpibus Scoticis’, but of the founders of the BBS in 1922. Pearson’s flora markers of evolution that should be melded with Academy of Sciences, U S A 103, 15511–15516. later demonstrated by Johannes Proskauer that was a considerable advance on his predecessors; his type specimen actually came from western it was the first truly comprehensive flora. The North America, not Scotland, probably collected by drawings lacked the critical detail of Hooker’s, and British liverwort floras 1741–1999: from Johann of liverwort species has steadily increased and now Archibald Menzies. the book was soon overshadowed by Macvicar’s as J. Dillenius to Jean A. Paton – David G. Long stands at 299. My four ‘giants of hepaticology’ and The second ‘giant’ is Sir William Hooker, who a practical flora. Henry W. Lett is perhaps the least their ground-breaking liverwort floras in this story became one of Britain’s most famous botanists well-known of our liverwort flora-writers, but he was a Britain has more liverwort floras than any other are J.J. Dillenius, W.J. Hooker, S.M. Macvicar and and Director of Kew Gardens, but before that he very important figure in the development of bryology country in the world, no less than 10 spanning of course Jean Paton. These are the authors who had been Professor of Botany in Glasgow for 21 in Ireland. He was a contemporary of Pearson and 259 years. Jean has some very distinguished have made major leaps forward in our knowledge. years and was author of many important works Macvicar, and acknowledged the help he got from predecessors, so writing a new flora demands The others in between have largely synthesized on cryptogamic plants, particularly bryophytes Macvicar. Perhaps his greatest bryological moment some very special qualities: exceptional ability, the work of earlier authors, apart perhaps for W.H. and ferns. One of the most important is his British was the discovery of Adelanthus dugortiensis on patience, attention to detail, sheer dedication and Pearson. Jungermanniae (1816), which covered only leafy Achill Island (now A. lindenbergianus). not least the ability to draw plants. Jean, fortunately, The first important British bryologist was Johann liverworts, with exceedingly detailed descriptions The next of our ‘giants’ is Symers Macvicar has all these qualities, but has certainly needed J. Dillenius, who was Sherardian Professor of and illustrations, for example Jungermannia whose work (1912) became the Standard British them over the 34 years spent researching and Botany in Oxford from 1734 until he died in 1747. hutchinsiae (now Jubula) which Hooker named after Liverwort Flora for 78 years – it is concise and writing her flora. Over these 259 years the number He was a correspondent and friend of Linnaeus Miss Hutchins of Bantry who sent him specimens. practical, still very useful today for its taxonomic

44 FieldBryology No101 | May10 FieldBryology No101 | May10 45 Meeting Report – Sussex 2009 Meeting Report – Sussex 2009

notes and acute observations. He married into a artist (in inking the plates) are a major contribution 700 v Fig. 1. Total numbers of records of wealthy family and was able to live on his wife’s to her liverwort flora. In her own words, over‘ the mosses (open bars) and liverworts 600 estate in Moidart in the oceanic west of Scotland. years, Pat spent hundreds of hours inking most of (solid bars) in each year. He was the first hepaticologist to become really the figures’. 500 that were published with the vice- familiar with our oceanic liverworts. He discovered I would summarize the important attributes of county records are included. many new species in Scotland. His work is a model a good liverwort flora as follows, and all of these 400 There were two phases of high of a good synoptic flora. qualities are exemplified by Jean’s book: recording activity (Fig. 1). In the Although Tony Smith’s liverwort flora (1990) has 300 x clear, precise descriptions, incorporating data first phase, 1952–1972, there was been superseded by a later book, his contribution to from living plants much activity in Ireland, with more Number of records of Number 200 British bryology has nevertheless been enormous, x well-constructed keys using most easily- than 500 Irish vice-country records particularly his pivotal role in the BBS mapping observed characters; alternative keys where 100 in 1958 and 1967. After 1971, Irish scheme which led to the excellent Atlas. During the useful, e.g. gemmae in Scapania vice-county recording was at a war, as a schoolboy Tony and his mother had a very 0 x attention to detail, e.g. oil bodies, cuticles much lower level, averaging only fortunate escape when they had to flee Singapore x full details of reproductive structures, gemmae, 1950 1960 1970 1980 1990 2000 49 per year from 1972 to 1997. before the Japanese invasion. In the confusion androecia, gynoecia, perianths, sporophytes Year of publication The nadir was 1997, with two Irish they boarded the wrong ship and eventually got to x accurate cell range measurements vice-county records. Australia, but the ship they should have boarded of the Recorders for 1997–2006 (published 1998– was lost at sea. x understanding and description of variation within 2007). CONTRIBUTION OF INDIVIDUAL RECORDERS Our fourth ‘giant’ is with us today; Jean and her species For the calculation of totals for individual recorders, outstanding contributions to hepaticology over x diagnostic taxonomic notes for critical taxa RESULTS recorders were given points for records, 1 for a solo many decades is the reason for our celebrations. x information on fruiting, frequency of reproduction At the end of the process, there were 22,532 usable record, 0.5 for contributing to a pair (A & B), 0.333 Eric Watson in his ‘Foreward’ summarized some x cytological information records. Numerous early records were of varieties for a triplet (A, B & C) and 0.25 for a quadruplet of the qualities of her work very aptly: ‘the quality x precise ecological information that are no longer recognized. Records from notices (A, B, C & D). Using this scoring system, the top and completeness of the illustrations… far surpass x good summary of geographical distribution of new species and from revisions were mostly not 20 recorders accounted for 12,580 records (Table what is to be found in any other British bryophyte x high-quality illustrations of all features of plants published with the vice-county records. Only those 1), more than half of the total 22,532 records. It flora known to me’, ‘Mrs Paton’s long and intimate x good illustrated glossary will come as no surprise to those familiar with the experience of these plants in the field’, [the Table 1. Top 20 recorders over the period 1947–1999 period that Jean Paton and Alan Crundwell were illustrations] ‘a triumph indeed’ and ‘a landmark in David Long, The Herbarium, Royal Botanic the most prolific contributors. the study of British liverworts’. I would like also to Garden, Inverleith Row, Edinburgh EH3 5LR Recorder No. Recorder No. On average, the most prolific recorder in each mention Jean’s husband, ‘Pat’, whose skills as an (e [email protected]) Paton, J.A. 1,924 King, A.L.K. 551 year found about 16% of the records. In 11 out of the 53 years, the top recorder found more than Sixty years a Queen: vice-county recording since DATA AND DATA PROCESSING Crundwell, A.C. 1,265 Fitzgerald, R.D. 542 20% of the total. From 1969 to 1972 and again in 1947, with observations on where Jean Paton 1977 and 1980, this person was Jean Paton, with a In an effort to eliminate the resulting backlog and Long, D.G. 792 Hill, M.O. 519 went and what she found – Mark Hill clear maximum of 40% of the records in 1971. She check for other omissions, I decided to capture Appleyard, J. 743 Blockeel, T.L. 456 is indeed the queen of vice-county recording. Starting in 1947 after an interruption during the all vice-county records from 1947–1999, by Warburg, E.F. 731 Ratcliffe, D.A. 386 Second World War, lists of vice-county records submitting photocopied pages for optical character ACKNOWLEDGEMENTS have been published annually up to the present. recognition (OCR). This activity was performed Wallace, E.C. 710 Rose, F. 367 I am grateful to Joint Reprographic Services for optical Records since 2000 have been added electronically by the Research Councils’ Joint Reprographic Lobley, E.M. 602 Perry, A.R. 353 character recognition, and to Tom Blockeel and David to the BBS database held at the Biological Records Services at Swindon. On 2 November 2007, they Long for comments on an earlier draft. Centre (BRC). Prior to that, some were added while very efficiently processed the 860 printed pages. Fitzgerald, J.W. 592 Synnott, D.M. 328 others were not. Inclusion was not systematic, and All I had do was to transform electronic output into Duncan, U.K. 567 Milne-Redhead, H. 307 Mark Hill, CEH Wallingford, Benson there was a gap during 1990–1999 following the usable data. For the analysis that follows, additional Lane, Crowmarsh Gifford, OX10 8BB Corley, M.F.V. 557 Jones, E.W. 287 Atlas of the Bryophytes of Britain and Ireland. statistics were added from the annual reports (e [email protected])

46 FieldBryology No101 | May10 FieldBryology No101 | May10 47 Meeting Report – Sussex 2009 Meeting Report – Sussex 2009

46 FieldBryology No101 | May10 FieldBryology No101 | May10 47 Meeting Report – Sussex 2009

v Left. Jeff Duckett (yes, v Left. Jonathan Sleath admires patches of Phaeoceros laevis. it is Jeff) and Mark Pool Ian Atherton search for Syntrichia , Centre left. Phaeoceros laevis. David Holyoak latifolia. Ian Atherton , Bottom centre and right. BBS members in the Francis Rose v Right. There’s always Reserve at Wakehurst Place. Jean Paton is just in shot on the far one! Has Mark Hill right). Ian Atherton (centre) & Robin Stevenson (right) spotted something the rest of the group stream side were much admired together with fine missed? Ian Atherton Dichodontium pellucidum. Tilgate Wood is one , Centre left. Harpanthus of the best Central Wealden woodlands with fine scutatus. John Birks outcrops of the Lower Tunbridge Wells Sand long , Bottom left. famous for their Atlantic bryophytes. The wood’s Dichodontium particular speciality, Harpanthus scutatus, was soon pellucidum. Des found together with Scapania umbrosa, to delight Callaghan our special guest. Diligent searches of material post the excursion also revealed Blepharostoma EXCURSION TO WAKEHURST PLACE, trichophyllum mixed with other species. Several nice colonies of Tunbridge Filmy-fern, Hymenophyllum On Sunday 39 members joined the excursion to tunbrigense, growing well on the rock outcrops RBG Kew’s Wakehurst Place Gardens and Tilgate were also admired. Altogether 64 species were Wood, which Kew have designated the Francis recorded and added to Wakehurst’s database, with Rose Reserve. We were met and welcomed by thanks to Richard Fisk for keeping the day’s tally. Andy Jackson, head of RBG Kew, Wakehurst Place I would like to express my thanks to Andy Jackson who stayed with the party for the rest of the day. and his staff for the trouble that they took to make Car parks are well known to be irresistible to a the visit so enjoyable and especially for providing dedicated few and both Didymodon nicholsonii and the transport to the wood. Syntrichia latifolia dutifully made the list even before the excursion started! David Streeter On the walk through the gardens to the wood, John Maynard Smith Building, University of Sussex, the large patches of Phaeoceros laevis by the Brighton BN1 9QG (e [email protected])

48 FieldBryology No101 | May10 FieldBryology No101 | May10 49 Meeting Report – Sussex 2009

48 FieldBryology No101 | May10 FieldBryology No101 | May10 49