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Meetings of the BBS - 1998

Summer Field Meeting 1998

Islay and Jura, July/August 1998


Account of meeting - Week 1 - Isle of Islay, 25-31 July

Participants: John Blackburn, Tom Blockeel, Agneta Burton, Blanka Buryová (from Prague), Richard Gulliver (local secretary), David and Geraldine Holyoak, Frank Lammiman, David Long, Seán O’Leary, Jean Paton, Mark Pool, Christine Rieser, David Rycroft, Graeme Smith, Philip Stanley, Rod Stern and Harold Whitehouse.

Headquarters: White Hart Hotel, Port Ellen.

All excursions were in VC 102 (South Ebudes).


The majority of the members attending met at Kennacraig in time for the 18.00 Port Ellen ferry. The weather was windy and drizzling, but spirits were generally high. An evening briefing took place at Richard and Mavis Gulliver’s home on the western outskirts of Port Ellen, after which members went to their assorted lodgings hopeful of a fine day to follow.


Kintra and points west, NR3248 to NR3048

The day did not start particularly well, the party gathering at Kintra in lashing rain. No sooner had they struggled from their cars, however, than the rain stopped; by the time lunch was taken, on sandy banks at Port Alsaig, the sun was beating down.

The ground west of Kintra consists largely of moorland, but with considerable basic influence locally on the coast due to blown shell-sand. As is usual on such excursions, the party took a long time to cover the first few hundred metres. Initially, the bryoflora was not of consuming interest, being made up largely of common calcifuge species. Highlights included Frullania microphylla and F. teneriffae (from coastal rocks), and Entosthodon attenuatus (Funaria attenuata), E. obtusus (F. obtusa), Lophozia incisa and Nardia geoscyphus from the heathy ground above. An already respectable list was given a considerable fillip when a flush just east of Port Alsaig was reached. Plants of interest here included Drepanocladus cossonii, Scorpidium scorpioides and Thuidium delicatulum; sandy rocks nearby, sometimes damp, produced a number of calcicoles, such as Bryoerythrophyllum (Barbula) ferruginascens and Jungermannia paroica.

Lunch over, the party divided. Tom and Mark investigated some rocks just above the shore; the rest of the group struck uphill, passing an old concrete reservoir (or similar) which produced common calcicoles, such as Didymodon rigidulus (Barbula rigidula) and Orthotrichum anomalum, from its sides. David Holyoak also recorded fruiting Splachnum ampullaceum from hereabouts. The party coalesced in the stunted (and, in part, very wet) woodland just to the west. Cololejeunea minutissima* was found here, growing on sallow; Lejeunea patens was seen not far away. A move further westward, onto more open ground, soon turned up Cephalozia leucantha; not long after this, a rock outcrop was checked and was proved to be basic by the presence of Hymenostylium (Gymnostomum) recurvirostrum and G. calcareum. Most of the party then descended into a large gully leading down towards the coast west of the Allt Fada (NR306480); this had an interesting, but not dramatic flora, perhaps the most noteworthy species being Jungermannia pumila. The total ‘bag’ from this excursion was an impressive 153 species (106 mosses and 47 liverworts).

David Long spent the day ‘freelancing’ south-west of Port Ellen. He visited the Carraig Fhada area (NR3444) and the vicinity of Port an Eas (NR3342); the highlights of another useful list (62 mosses, 18 liverworts) were Jubula hutchinsiae, Marchesinia mackaii, Dicranum scottianum, Fissidens polyphyllus and Zygodon viridissimus var. stirtonii.

The evening saw all those present being entertained by the Gullivers to a meal at their home. This was not only excellently prepared, but also efficiently organised; not many people would have been capable of feeding sixteen hungry bryologists at once in a domestic setting!


Duich Moss (NR3355, NR3356 etc.)

The party met initially at the end of a rough track a few hundred metres south of Laggan Bridge. This track proved very interesting, occupying most of those present for a considerable time. Species seen here included Aongstroemia longipes*, Didymodon ferrugineus* (Barbula reflexa), Fossombronia incurva*, Riccardia incurvata and a number of commoner ruderals. After following the track almost to the shed at its terminus, members turned south-westwards onto the Moss proper. Initially this proved disappointing, the most noteworthy bryophyte seen being the locally abundant Campylopus introflexus. Members persevered, however, and after a slightly tricky stream crossing (roughly at NR333561) came to better ground. It was not long before Sphagnum austinii (S. imbricatum ssp. austinii) was found; soon afterwards S. fuscum (albeit in small quantity) turned up as well. Cephalozia loitlesbergeri was found by Jean growing among Sphagnum magellanicum and Leucobryum. Encouraged by these, and a steady trickle of other, less unusual, records, the group continued southwards towards the pools at Eilean na Muice Duibhe. These were also of interest, not least for the presence here and there of Calypogeia sphagnicola* among the sphagna. After this, members wended their way back to the cars for lunch. Most went back by the track; Mark, pursuing a solitary route across the moss, was rewarded with several good colonies of Polytrichum longisetum on the sides of old peat diggings. The conifer plantation by the roadside attracted some attention, as it was seen to contain a few sycamores; these produced some common basicolous epiphytes for the list. Nowellia curvifolia was also noted here at one spot, on a rotting log. The total list for the morning was 77 (54 mosses, 23 liverworts).

North of Loch Fada (NR4063, NR4064, NR4164)

The area between Loch Fada and Mullach Dubh has considerable areas of limestone, which forms incipient pavement in at least one site. The party gathered after lunch by the track-end at NR400639; this spot was obviously on limestone and a good list of calcicoles soon resulted, perhaps the best being Didymodon ferrugineus and Brachythecium glareosum. Progressing eastwards along the track, the group became somewhat scattered. David Holyoak, Rod and Mark digressed onto moorland a little to the north, where the ground was much more acidic and had a number of boggy areas. Mylia anomala, M. taylorii, Odontoschisma denudatum and Pleurozia purpurea were found here, as were no less than eight Sphagnum species, including S. magellanicum. Andreaea rupestris, wanted as a vice-county record, was seen on a rock outcrop hereabouts, but was not collected as it was in small quantity (it was subsequently seen, and collected, on Jura the following week). The three ‘separatists’ rejoined the others at the limestone pavement (NR411642), finding Sphagnum squarrosum in a flush on the way. The main party had given the limestone area a thorough search, and had been rewarded with (among others) Porella arboris-vitae, Bryoerythrophyllum (Barbula) ferruginascens, Hypnum lacunosum var. tectorum* and Thuidium philibertii. In addition, Tom had found Bryum subapiculatum* (B. microerythrocarpum) and Pohlia camptotrachela on damp soil near the track during the walk in. Due largely to the variety of habitats examined, the afternoon’s list included 131 taxa (25 liverworts, 106 mosses).


Ballygrant woodlands etc. (NR36, NR46)

The Ballygrant woodlands promised to be an interesting habitat, providing more shelter than is usual over much of Islay and also possessing a varied geology including much limestone. Cars were parked by the roadside just south of Ballygrant village (in NR36) and a useful list of common species was compiled from the roadside walls. Of more interest, however, was an adjacent Brassica field. This quickly produced Dicranella staphylina* (almost unbelievably a new vice-county record); further meticulous research by Harold and others turned up Bryum klinggraeffii*, B. sauteri* and Ditrichum cylindricum*. After such a start the woodlands could easily have been an anticlimax; this was prevented, however, by their sheer bryophyte biomass, by Christine’s discovery of Frullania teneriffae almost at the entrance, and by the finding of Neckera pumila*, on beech, soon afterwards.

Members followed the woodland track along the north side of Loch Ballygrant. Much of the ground here was decidedly acidic; the species seen, while providing a useful boost to the day’s list, were in the main common ones. The best were Lejeunea patens, Plagiochila britannica (abundant), Riccardia palmata and Hypnum lindbergii. Spirits leapt when the very obvious limestone around Loch nan Cadhan was reached. Rock outcrops and walls here had Anastrophyllum minutum, Plagiochila britannica, Scapania aspera, Campyliadelphus chrysophyllus (Campylium chrysophyllum), Entosthodon obtusus, Syntrichia (Tortula) intermedia*and Zygodon viridissimus var. stirtonii An area of partly basic marsh produced Sphagnum flexuosum* (S. recurvum var. amblyphyllum), S. inundatum (S. auriculatum var. inundatum), Drepanocladus cossonii and splendid Philonotis calcarea, while among the heather on a presumably acid rock outcrop nearby Tom and David turned up another new vice-county record in the shape of Barbilophozia atlantica*.

Lunch was taken, in hazy sunshine, on a breezy knoll above the loch; the group then continued north-eastwards along the main track. Limestone was much in evidence beside the path at first, but apart from Marchesinia and Porella arboris-vitae it produced nothing unusual which had not been seen already. Some well-grown sycamores were searched in hopes of finding Zygodon rupestris (Z. baumgartneri) but unfortunately all the specimens collected were found when checked to be Z. viridissimus. One sub-group spent some time in an unsuccessful search for Cryptothallus, but were consoled by finding a good colony of Lesser Twayblade Listera cordata. Once away from Loch Ballygrant, the track passed through relatively uninteresting country until Loch Allan was reached. Even so, there was some interest; Mnium stellare was recorded on a shaded wall, Didymodon ferrugineus was locally abundant on the track, Riccardia chamedryfolia* was found on a soil bank, and Fossombronia wondraczekii* (accompanied by F. pusilla and Pohlia flexuosa (P. muyldermansii)) on soil in a gateway. Sallow by Loch Allan sported Cololejeunea minutissima, while waterside rocks produced a good (but unconfirmable) candidate for Didymodon spadiceus (Barbula spadicea). As a final parting shot, David Long found Phaeoceros laevis* on a bank by the track near Dunlossit House.

As expected, the day’s total was high (129 mosses and 52 liverworts).


Rubh’ a’ Mhàil and the coast to the west (NR47, NR37)

The headland of Rubh’ a’ Mhàil (pronounced ‘Ruvaal’) lies at the extreme north-eastern corner of the island. Following good reports of the sea-caves to the west, at least some of the party hoped to reach that area. Others, put off by the thought of the four-mile approach walk and (perhaps) by the wet morning, opted to sample the bryophytes of the area around Bunnahabhain, a small distillery hamlet at the end of a road to the south.

Both groups initially met at Bunnahabhain. The ‘headland party’ (Tom, Agneta, Blanka, Richard, David Holyoak, David Long, Seán, Mark and Graeme) duly set off northwards in blowing drizzle, only slightly disconcerted by having to wade a small sea-inlet before following the ‘path’ up a low, but vertical, quartzite crag. From then on the path was clearer, if sometimes boggy; as the drizzle turned to driving rain soon afterwards, most of the group were soon too wet to care anyway. Due to a combination of the weather and the rapid pace set by the fitter members of the party, little bryology was done during the walk in. By the time the Rubh’ a’ Mhàil lighthouse was reached, the rain had stopped and the sun was coming out, so heads emerged from cagoules and started to take an interest in the surroundings.

The coast west of the lighthouse is spectacular in parts, with steep-sided inlets and many sea-caves. David Long, Blanka and Mark descended into the first of these gullies, which sported a total of three sizeable caves. These were all searched throughly in hopes of finding Cyclodictyon laetevirens, which is reported from this area, but without success. There was, however, consolation in Dicranum scottianum, which grew in large cushions on quartzite rock faces; Isothecium myosuroides var. brachythecioides was also present locally. The caves themselves had, among other things, Jubula hutchinsiae and Lophocolea fragrans; one contained a baulk of rotting timber which was well colonised by Riccardia palmata.

The rest of the group had carried on westwards to investigate the caves and cliffs of Bàgh an Dà Dhoruis. They had still not found Cyclodictyon, but there were compensations. These included Isopterygiopsis pulchella* (Isopterygium pulchellum), found in fruit by Tom in a deep crevice on a cliff, more Dicranum scottianum (providing a habitat for Microlejeunea (Lejeunea) ulicina at times), Leiocolea bantriensis (on a flushed slope), Marchesinia mackaii (found by David in a cave-mouth) and Plagiochila punctata (on rock). The party reunited for lunch on the windy headland of Aonan an Dà Dhoruis, subsequently splitting again into a number of small sections. David Holyoak found the rare Lejeunea holtii in a sea-cave (which also produced Trichomanes gametophytes). Mark undertook a solitary exploration of several caves, but again failed to find any Cyclodictyon; there was, however, another baulk of timber, this one with a thriving population of Nowellia as well as Riccardia palmata. An unexpected feature of one of these caves was a seepage of obviously basic water from the joints of the normally acid quartzite; this produced Eucladium verticillatum and Didymodon tophaceus (Barbula tophacea). David Long, Tom and others had gone on further west as far as Port a’ Chotain (in NR37), before heading inland up the valley of the Allt na h-Uamha Móire. Port a’ Chotain had Lejeunea patens and more Isothecium myosuroides var. brachythecioides, together with a patch of Wood Vetch Vicia sylvatica. The valley produced a number of western hepatics, the most notable being Cephalozia catenulata, C. leucantha, Jungermannia subelliptica, Kurzia trichoclados, Mylia taylorii, Odontoschisma denudatum (on peat) and Pleurozia purpurea. Members made their way back to Bunnahabhain by a variety of routes; the weather was by now fine and sunny, and David Long’s find of Campylopus subulatus* on a rough track near the village rounded off a very satisfying day.

Bunnahabhain area and Ardnahoe Loch (NR47)

The Bunnahabhain group was made up of John, Frank, Jean, Christine, Phil, Rod, David Rycroft and Harold. They started by checking the valley of the Abhainn Araig, north of the village; this produced such species as Cephalozia lunulifolia, Frullania teneriffae (found by Christine), Metzgeria conjugata, Plagiochila punctata, P. spinulosa and Tritomaria quinquedentata. From here the party struck southwards, initially to the old quarries at NR422721; these had (among other things) Pohlia annotina, Leiocolea turbinata and Scapania irrigua. Loch Ardnahoe, checked next, produced a useful record of fruiting Orthotrichum rupestre.


Beinn Bheigeir and the Ardtalla area (NR45)

Beinn Bheigeir is in many ways an unremarkable quartzite hill. Most of the party were, however, hoping to visit it, as it is the only non-Irish site in the British Isles for the rare liverwort Adelanthus lindenbergianus. The party gathered at Claggain Bay and the majority promptly set off for the hill; the others (of whom more later) stayed low, to look at some of the old woodlands of the area.

The approach to Beinn Bheigeir, as with so many similar hills, starts with boggy moorland and then progresses to steep heathery slopes. Little serious bryologising was done on the ascent; David Long did, however, find Sphagnum contortum in a Juncus flush. Higher up, a gale-force wind was blowing but this did not seem to deter the group, most of whom were soon investigating the numerous patches of ‘mixed hepatic mat’. Liverworts noted here, in addition to the Adelanthus, included Anastrepta orcadensis, Bazzania tricrenata, Herbertus aduncus ssp. hutchinsiae, Nowellia curvifolia (on peat) and Pleurozia purpurea, while the best of the mosses were Andreaea rupestris and Dicranodontium denudatum.

Lunch was taken in a sheltered spot just below the crest of the ridge; afterwards the party moved back towards the coast. Some time was spent in the ravine of the Allt nam Bodach (NR460562); this looked promising initially but turned out not to be rich in species. The reason was not obvious, but might be a combination of the ravine’s relative openness and its north-easterly exposure. The most interesting species seen were Lejeunea lamacerina, Metzgeria conjugata, Plagiochila spinulosa and Saccogyna viticulosa. Progressing roughly southwards, the group reconvened in some damp coastal scrub near Ardtalla House. A potential candidate for Ulota drummondii was collected from here, but unfortunately keyed out as U. bruchii (U. crispa var. norvegica) when checked later. Large rock outcrops nearby produced Frullania fragilifolia (its distinctive turpentine-like smell very noticeable here) and Plagiochila killarniensis; coastal rocks and sand at Traigh Bhàn had Schistidium maritimum, Tortella flavovirens and (as so often) a large amount of a promising-looking, but sterile, Bryum sp.

Various woodlands of south-east Islay (NR45, NR44)

The ‘lowland party’ had looked first at the wooded ravine of the Claggain River and the surrounding wet heathland. This produced 45 mosses and 21 liverworts, but none was rare or particularly unusual. Drepanocladus revolvens, Heterocladium heteropterum and Plagiochila punctata were perhaps the most interesting. From here they progressed to the rocky oak/birch woodland (an SSSI) near Trudernish Point (NR4652); this had rather more interest, with Thuidium delicatulum, Cephalozia lunulifolia, Cephaloziella hampeana, Fossombronia wondraczekii, Scapania umbrosa and Tritomaria exsectiformis heading a list of 28 mosses and 20 liverworts. This group’s final visit of the day was to a wood near Calumkill, north-west of Ardbeg (NR4046). This again was not of spectacular interest, but 29 mosses and 21 liverworts were noted. The wood appeared to be slightly more basic than the other two; Palustriella commutata var. falcata (Cratoneuron commutatum var. falcatum) and Neckera complanata were both seen here, while highlights included Hylocomium brevirostre and Trichocolea tomentella.


Killinallan area (NR3071 to NR3374)

The day began fine and dry, if a little cloudy. Cars were parked by the track south-west of Killinallan, and it is some indication of the interest and variety of the area that the party immediately fragmented. David Holyoak and others went to look at some incipient salt-marsh on the edge of Loch Gruinart; this produced plentiful Hennediella (Pottia) heimii, some of it fertile, together with Drepanocladus aduncus and a sterile Bryum which was believed to be the rare B. marratii but which was still under investigation at the time this report was written. This party then went into the dunes to the north, finding Climacium dendroides, Entodon concinnus and a certain amount of Thuidium philibertii, in addition to common basicole species such as Homalothecium lutescens. The main group was rejoined in a decidedly marshy area which had, among other things, Drepanocladus polygamus* (Campylium polygamum) (found by David Long) and Calliergon giganteum. Interest soon shifted, however, to a nearby rock outcrop; this provoked some discussion, as it appeared to consist of acid quartzite but sported a number of calcicole species (examples being Didymodon rigidulus, Encalypta streptocarpa and Zygodon viridissimus var. stirtonii), as well as others typical of coastal rocks hereabouts (e.g. Frullania fragilifolia and Porella obtusata). The conclusion was that the calcareous influence was provided by blown sand, which can build up on the tops of outcrops to provide basic flushing after rain; this effect was to be noted on several occasions later in the day.

Moving on in a basically north-easterly direction, the party fragmented again. Mark found a good colony of Reboulia hemisphaerica growing on flushed, sheltered rocks; after this he teamed up with Rod to see sheets of Entodon, with frequent Thuidium philibertii, on the slopes of calcareous dunes. Dichodontium pellucidum was also common here, a surprise for those of us accustomed to finding it by hill streams. The party reassembled in a fascinating area of the aforesaid ‘basic quartzite’, outcropping from ground which had been heavily poached by cattle and which, to judge from the presence of Cinclidotus fontinaloides on the base of a rock, held standing water in winter. This poached ground produced one of the week’s best discoveries in the form of Riccia cavernosa*. The rocks nearby had a considerable variety of calciphiles, of which the most noteworthy were Orthotrichum rupestre and fruiting Marchesinia mackaii.

Lunch was taken, in bright sunshine, near these rock outcrops; afterwards most of the party investigated a large and promising dune slack nearby. David Holyoak said it looked an ideal Petalophyllum habitat, but of course nothing could be seen of the plant (even if it was present) at this time of year. Some banter was exchanged over the possibility of a March meeting in the area, but nothing definite materialised; in the interim the Society had to be content with an abundance of Didymodon (Barbula) fallax, Bryum algovicum var. rutheanum and other interesting, but sterile and therefore unidentifiable, Bryum spp.

The dune belt narrowed eastwards from here, and appeared less bryologically interesting. One small group of diehards (Seán, Mark and Graeme), continued as far as Gortantaoid Point, where there was a local abundance of stunted Porella obtusata together with Tortella flavovirens and a small quantity of Isothecium myosuroides var. brachythecioides. A splinter group, including Tom, Jean and others, visited a patch of birch-hazel woodland east of Bun-an-uillt (NR3069), but found it infested with Rhododendron and bryologically unpromising. They nonetheless managed to produce a list of 65 species (35 mosses, 30 liverworts) from the wood and the surrounding moorland. The highlight was Cephalozia macrostachya var. spiniflora*, found by Jean; others included Hylocomium brevirostre, Sphagnum magellanicum, Cephalozia catenulata (on peat), Chiloscyphus pallescens, Mylia anomala, Odontoschisma denudatum and Trichocolea tomentella. The rest of the group returned to the vehicles by way of the dunes, but do not appear to have added much new to the list. This eventually totalled 92 species (78 mosses, 14 liverworts), excluding those found by the Bun-an-uillt group.

In addition to the official excursions, various unscheduled visits were made to other sites during the course of the week. David Holyoak and Rod carried out a full survey of Richard and Mavis’s garden at Imeravale; they found a total of 54 taxa (46 mosses, 8 liverworts), including Bryum bornholmense*, Isothecium myosuroides var. brachythecioides and Schistidium maritimum. Harold discovered Bryum rubens* in two separate arable fields, one about 3 km north-east of Bridgend (NR359641) and the other a similar distance south-west of Bowmore (NR284579).

The week had been more than satisfactory, despite the rather indifferent weather. Although it appears to lack the best of the western Scottish oceanic-montane bryoflora, Islay was found to be a varied and interesting island. No doubt work still remains to be done, but with a total of 25 new vice-county records (16 mosses, 9 liverworts) found during the week there should be less than formerly! Our sincere thanks go to Richard and Mavis Gulliver, both for organisation of the week and for the superb catering on the Sunday night. Thanks are also due to the management and staff of the White Hart Hotel for providing the headquarters, and to the various landowners for granting permission to visit (and, in most cases, collect) on their land. My own thanks as writer of this report go to the following people, all of whom helped by sending in lists: John Blackburn, Tom Blockeel, David Holyoak, David Long, Jean Paton, Christine Rieser, Rod Stern and Harold Whitehouse.


Account of meeting - Week 2 - Isle of Jura, 1-7 August


A reduced party from that on the Isle of Islay caught ferries for the short crossing to Jura (VC 102) during the morning: John Blackburn, Agneta Burton, Blanka Buryová, David and Geraldine Holyoak, Jean Paton, Mark Pool, Phil Stanley, Rod Stern and Harold Whitehouse.

From the ferry crossing the summits of the Paps of Jura, three conical quartzite mountains that dominate the southern part of the island, seemed quite close and binoculars revealed Red Deer scattered on the long moorland slopes leading up to the higher rocks, as well as a Golden Eagle soaring over a distant ridge.

Abhainn Mhór to Daimh-sgeir, NR4468 to NR4467

Agneta, Jean and John caught an early ferry and began work in earnest before the rest of the party reached Jura. A substantial list of bryophytes was recorded by them from stream banks, including Frullania teneriffae and Diphyscium foliosum.

We then moved on to the Jura Hotel at Craighouse, which was to be our (very comfortable) headquarters for the week. In warm, sunny weather, we enjoyed a picnic lunch on the hotel lawns.

Craighouse ‘ravine’, NR5266

The afternoon was devoted mainly to exploration of the wooded, rocky valley of the small river that reaches the sea near the Jura Hotel. Luckily the water-level was low and, except for a climb to avoid the waterfall, there was little difficulty in following the course of the river upstream, giving access to a luxuriant bryophyte flora that included Aphanolejeunea microscopica, Bartramia ithyphylla, Dumortiera hirsuta (in several places), Fissidens celticus, F. curnovii, Heterocladium heteropterum var. heteropterum, lots of Jubula hutchinsiae, Lejeunea lamacerina, L. patens, scattered Lophocolea fragrans, Metzgeria conjugata and also Ulota drummondii. On the walk back, an old track into a pasture near the roadside produced Fossombronia wondraczekii, Philonotis caespitosa*, Pohlia bulbifera, P. camptotrachela, Riccia sorocarpa and R. subbifurca.


Corran River valley, NR5472 to NR4973

During a dry morning the whole party followed the Corran River upstream across open moorland to Loch an t-Siob. Peaty banks along the way produced Cladopodiella francisci* and Lophozia bicrenata* found growing very close together by Jean, and other finds included Anastrophyllum minutum, Calypogeia neesiana, Campylopus atrovirens, Cephaloziella divaricata, C. hampeana, Kurzia trichoclados, Pohlia bulbifera and Sphagnum quinquefarium. The edge of Loch an t-Siob had Scapania umbrosa in addition to patches of Water Lobelia Lobelia dortmanna.

During the afternoon the slopes of two of the Paps of Jura around Gleann an t-Siob and an adjacent hill were visited by small groups. The rather unprepossessing quartzite screes of the southern slopes of Beinn Shiantaidh were covered by Agneta, Jean, Harold, Phil and Rod, who found Glyphomitrium daviesii and Hedwigia stellata* there on small boulders of basaltic rocks, whereas the quartzite boulders were bare. John and Mark tackled the wetter northern slopes of Beinn Mhearsamail, and found extensive ‘mixed liverwort mats’ that included Bazzania tricrenata, Herbertus aduncus ssp. hutchinsiae and Plagiochila carringtonii, as well as Racomitrium ellipticum*. Blanka and I searched the north-eastern slopes of Beinn Chaolais, finding similar liverwort mats but with the addition of Anastrepta orcadensis, Barbilophozia floerkei and Dicranodontium asperulum; Andreaea alpina* occurred there on rock.


Inver Cottage area, NR4469 to NR4473

A large group braved steady light rain with occasional heavier outbursts to explore the coastal slopes northwards from the Feolin Ferry, with its stretches of raised beach and sea-caves. Cyclodictyon laetevirens was admired at its well-known site inside a damp cave entrance, and Calypogeia azurea was noticed growing nearby. Despite the poor weather, energetic recording resulted in a total list of over 100 bryophyte species, including Blasia pusilla and Dicranella cerviculata, and new records for Pohlia flexuosa* (P. muyldermansii) and Splachnum sphaericum* found by Rod.

Jura House Gardens, NR4863

Smaller groups visited the attractive gardens and woodland around Jura House, Ardfin, and amassed large lists of bryophytes, but with few real surprises among them. The ‘Tea Tent’ in the gardens provided welcome respite from steady light rain. The garden was created a hundred years ago, and much of it is sheltered by a wall up to five metres high. It contains many Australian and New Zealand plants especially collected for the garden.

Coast near Daimh-sgeir, NR4468

Finally, Jean and Phil searched coastal rocks and caves above the road in continuing rain, locating Calypogeia muelleriana, Gymnocolea inflata, Plagiochila killarniensis, P. punctata, Dicranoweisia cirrata and Dicranum fuscescens


Lealt, NR6690

A morning with good weather was used to explore the ravine along a small river. The steep-sided ravine above the road had several low waterfalls and required some scrambling, but it proved to be rich in bryophytes with, among others, Bazzania trilobata, Blepharostoma trichophyllum, Fissidens taxifolius var. pallidicaulis, Harpalejeunea molleri (H. ovata) and Lepidozia cupressina, as well as masses of Jubula hutchinsiae, and plenty of Frullania teneriffae epiphytic on hazels. The lower part of the ravine produced Barbilophozia attenuata on the trunk of a living oak tree, Cephalozia catenulata on a log, C. lunulifolia, Harpanthus scutatus and Microlejeunea (Lejeunea) ulicina, but this part of the ravine also had steep and inaccessible sides.

Tramaig Bay, NR6588

Woodland on a low rocky slope near the shore had a varied bryophyte flora, including Adelanthus decipiens, Barbilophozia attenuata, Cephalozia lunulifolia, Frullania fragilifolia and Thuidium delicatulum. The best find was made just as we were about to leave, a patch of Plagiochila atlantica* found by Blanka.

Tarbert Bay, NR6082

Brief searching around the shores of the bay resulted in finds of Bryum subapiculatum (B. microerythrocarpum), B. rubens* (found by Rod), Drepanocladus aduncus, Pellia neesiana, Pohlia camptotrachela and Tortella flavovirens.

Loch na Mile, NR5471

The edges of several sandy bays were searched in the hope of finding Bryum salinum, but only unidentifiable, non-fertile material was found. Only slight consolation was obtained from finds of Pohlia bulbifera and Bryum rubens


Jura House Gardens, NR4863

A second visit to the gardens by a larger group resulted in two new finds there: Anthoceros punctatus* in garden plant pots and Marchantia polymorpha ssp. ruderalis* in garden beds and in pots. Other bryophytes were studied in an adjacent marshy field. Blanka, Harold and Phil left for the mainland later in the day. The remainder of the party explored a short distance along the coast to the east, then sat on the beach for lunch, watched by three wild goats high on the cliff.

Inver Cottage area, NR4469 to NR4473

A return visit to this rich area with Mark, in fine weather, resulted in a few additions to the list, including Hedwigia stellata, Orthodontium lineare and Thuidium delicatulum. The occurrence of numerous calciphiles, including Eucladium verticillatum, in a few places along the base of the quartzite sea-cliff was attributed to flushing by base-rich ground water.

Ardfernal, NR5570 to NR5671

Jean, John and Rod explored steep natural woodland, scrub, rocky grassland and salt-marsh around this coastal headland and bay. The most significant finds from a long list of bryophytes were Bryum dunense* and Barbilophozia floerkei found by Rod, B. subapiculatum by John, and Ulota drummondii by Jean. The eastern part of the same area was revisited briefly the next morning.


Beinn an Oir, NR4774 to NR4975

The remoteness of this area from the road led to arrangements being made for transport by Land Rover along rough tracks to the base of the mountain. The only day when this transport was possible unfortunately turned out to be very wet. Hence the small group (Agneta, Mark and me) who decided to walk into the mountains enjoyed steady rain and views of less than fifty metres throughout. The northern slopes of Beinn an Oir had areas with well-developed ‘mixed liverwort mat’, including Herbertus aduncus ssp. hutchinsiae and Plagiochila carringtonii, but there were no surprising finds. The long walk back to the road through the Inver Cottage area saw the weather improve, and allowed a few more additions to the bryophyte list for that by now well-worked area, including Sphagnum girgensohnii found by Mark.

Near Knockrome, NR5571

During the morning the ‘lowland group’ (Jean, John and Rod) walked from the airstrip near Loch na Mile along the shore and up over Rubha Bhride to search salt-marsh and dunes around the bay before returning by way of Ardfernal. Searches for Bryum salinum were again unsuccessful, but the bryophytes found included Blasia pusilla, Cephaloziella hampeana and Campylium stellatum var. protensum.

Jura Forest, NR5370

The ‘lowland’ group later explored the grounds of Jura Forest House, surrounding woodland and open moorland above. Despite rain and midges, the more significant finds included Anthoceros punctatus (outside the walled garden) and Phaeoceros laevis in the garden.


West of Tarbert, NR6082, NR5982, NR5984, NR6083

John and Rod left early for the mainland. The four bryologists remaining drove north to the narrowest part of Jura and worked an area of moorland, old tracks and the sea shores of Loch Tarbert, in mostly light rain. Substantial lists of species were recorded for two 10 km squares, including the first Sphagnum molle to be noticed on the trip, but the best finds were of Cephalozia macrostachya var. macrostachya* (discovered by Jean on a Leucobryum tussock in open moorland) and Campylopus subulatus. Other bryophytes recorded included Bryum subapiculatum, Calypogeia sphagnicola, C. neesiana, Cephalozia catenulata on a peat bank, Cephaloziella hampeana, Fissidens osmundoides, Kurzia trichoclados and Splachnum ampullaceum. Mark made a long walk to patches of woodland on the western side of the head of Loch Tarbert, where numerous species were added to the list, including Riccardia palmata and Sphagnum quinquefarium.

Partly bare patches in the top edges of the salt-marsh at the head of the loch had many patches of non-fertile Bryum, again raising hopes of B. salinum, but the absence of sporophytes prevented any of it from being identified.

Glen Shiel (VC 105), NG9217, NG9317, NG9318

After leaving Jura, Blanka, Harold and Phil travelled northwards on the Scottish mainland to search for Philonotis cernua (Bartramidula wilsonii) at a locality where it had been found by Dr Warburg 50 years ago. Although the P. cernua was not relocated, a rich bryophyte flora was recorded on the eastern slopes of Sgurr Mhic Bharraich, including Douinia ovata, Herzogiella striatella, Marsupella sphacelata, Plagiochila carringtonii, Scapania nimbosa and Sphagnum strictum.

The Isle of Jura is geologically less varied than Islay, being composed mainly of quartzite, so it supports a less varied bryophyte flora. Larger parts of Jura are also difficult to reach, since there is access by road only along the southern and eastern coasts, and long hikes or the use of a boat are necessary to reach many northern and western parts of the island. Hence, although the Isle of Jura does not have an especially rich bryophyte flora, there are doubtless still some species awaiting discovery there. The presence of such strongly Atlantic species as Cyclodictyon laetevirens on Jura and Islay, and Adelanthus lindenbergianus and Lejeunea holtii on Islay, may well hint that other Atlantic species still await discovery in VC 102, such as perhaps Bazzania pearsonii or Mastigophora woodsii. Nevertheless, the ‘northern mixed liverwort mats’ of the mountains of Jura are species-poor compared to examples elsewhere in western Scotland, and the impression of a rather restricted bryophyte flora on the island as a whole is probably correct.

Unfortunately, the famous island distillery adjacent to our hotel was closed for the week. The frequent rain was only heavy enough to restrict bryological fieldwork on some of the days and perhaps partly as a result of the wet weather, we were less troubled by midges, horseflies or deer ticks than might be expected in some parts of Scotland. Clear spells revealed some splendid mountain and coastal scenery, and allowed sightings of Merlins, Hen Harriers, Black Guillemots and Arctic Skuas.

Thanks are due to various estates for allowing access to their land, and to Richard Gulliver for making many of the arrangements for this access. John Blackburn, Jean Paton, Mark Pool, Rod Stern and Harold Whitehouse helped with providing lists for this report.


Annual General Meeting and Symposium 1998

University of Loughborough

The University of Loughborough was the venue for this year’s AGM and symposium meeting. Thanks are due to the local secretary, Dennis Ballard, who organised the weekend very ably and made sure that everything ran smoothly (or sorted it out when it didn’t!). As seems to be the norm in recent years (global warming?), the day of the field excursion was warm and sunny, and a good contingent of members were shown that even the Midlands can be bryologically interesting.



The Charnwood Forest area of Leicestershire, where John Ray reported the finding of rare cryptogams in the 17th century, was described. Dr Richard Pulteney (1730-1801), an apothecary, provided the first real account of the vascular and cryptogamic flora of the Charnwood Forest, Loughborough and Leicester. The Rev. George Crabbe (1754-1832) collected and recorded in north-east Leicestershire. Their works were published in Nicol’s 1795 History of Leicestershire. Several clergymen continued the collecting and recording in the 19th century. With the formation of the Leicester Literary and Philosophical Society in 1835, botany was put on a firm base, with a few other botanists continuing to advance bryology. The publication of their work was quoted. The activities of the Society and the Leicester Museum, which opened in 1849, resulted in the production of the 1863 Flora. Separate herbaria were maintained by the two organisations. In the 20th century a new generation of botanists carried bryology forward. Floras were produced in 1909 and 1941, and several papers published. It was not until 1984 that recording was carried out in a systematic way on a tetrad basis. More work is required before a comprehensive bryophyte Flora can be produced. Records are available at present from about one third of the 617 tetrads in Leicestershire.


Bryum argenteum is probably the world’s most widely distributed plant species, extending from continental Antarctica through the lowland tropics to the high Arctic. Something of the wide variation in microclimatic conditions experienced by different populations of this species was demonstrated by data showing summer moss-level temperature regimes at three polar and boreal forest sites; modal values for temperature during the day ranged from 0ºC on Ross Island, Antarctica, to 20ºC at Pinawa, Manitoba. Clones isolated from these populations nevertheless showed strikingly similar relationships between temperature and growth. All grew slowly at a day/night temperature regime of 9/1ºC, and fastest at 22/15ºC, with evidence of heat stress at 30/30ºC. This is consistent with the temperature relationships of CO2 exchange in an Antarctic population, in which the optimum temperature for net assimilation at light saturation was 25ºC (Rastorfer, 1970). Antarctic and tropical (Hawaii) clones both survived but grew very slowly at 5/-5ºC. Similarly, North American B. argenteum from normal and metal-contaminated soils showed similar levels of tolerance to copper and cadmium, whereas in Funaria hygrometrica there was evidence of genetic adaptation in this respect (Shaw, 1991).

These physiological responses thus give the impression that B. argenteum may exist as a small range of ‘multi-purpose genotypes’. However, collateral cultivation experiments have demonstrated extensive morphological variation in features such as nerve length and shape of the leaf apex. Some of this variation is apparently random, but plants with strikingly obtuse leaf apices and abundant bulbils have been isolated from both Arctic and Antarctic sites. There is also evidence that the excurrent nerve giving many colonies in dry habitats a characteristically hoary appearance (var. lanatum) is maintained in cultivation, in isolates derived both from gametophytes and from spores, although with no discontinuity in the range of variation between these forms and var. argenteum. Moreover, a study of the nuclear-encoded rRNA internal transcribed spacer (ITS) region in B. argenteum and related species showed that much of the total variation among these taxa resides within B. argenteum. Indeed, genetic distances between some populations of this species were greater than between those of some species traditionally assigned to different genera, a finding with important implications in terms of conservation strategies (Longton & Hedderson, in press). High levels of within- and between-population variation in RAPDs has also been reported in B. argenteum from the Antarctic and Australasia (Stotnicki, Ninham & Selkirk, 1998), emphasising the complexity of genetic variation in this most familiar of moss species. Beware lest familiarity breeds contempt!


Longton RE, Hedderson TA. In press. What are rare species and why conserve them? Lindbergia.

Rastorfer JR. 1970. Effects of light intensity and temperature on photosynthesis and respiration in two East Antarctic mosses, Bryum argenteum and Bryum antarcticum. Bryologist 73: 544-556.

Shaw AJ. 1991. Ecological genetics, evolutionary constraints, and the systematics of bryophytes. Advances in Bryology 4: 29-74.

Stotnicki ML, Ninham JA, Selkirk PM. 1998. High levels of RAPD diversity in the moss Bryum argenteum in Australia, New Zealand and Antarctica. Bryologist 101: 412-421.


Recording of the distribution of bryophytes in West Cornwall and the Isles of Scilly (vice-county 1) by tetrads (2 x 2 km squares) was begun in 1993. Well over half of the tetrads have now been visited, with a total of 340 full days spent in 242 tetrads.

By means of numerous repeated visits to each of several tetrads, it was found that two hours recording effort per tetrad was sufficient to find at least 50% of the bryophyte species revealed in the same tetrads by ten hours of searching, but that a few additional species continued to be added even after 15 hours of searching. These results emphasise that field recording of bryophytes is a process of sampling, and that complete coverage of any large area is unattainable. Hence, in order to offer the prospect of increased reproducibility in any future surveys, the time (hours) spent searching in each tetrad and the range of habitats searched are being recorded. Practical considerations and the aim of achieving reasonably even coverage have led to a ‘rule’ that each tetrad is sufficiently searched when all the main bryophyte habitats have been investigated, a total of at least two hours has been spent searching, and more than 50 moss species have been found. Allowing for travel, microscopic checking of material, and input of data onto computer, this ‘rule’ allows one tetrad to be completed in a single, rather long, working day.

The Atlas of the Bryophytes of Britain and Ireland (Volume 1, 1991) described Cornwall as ‘exceptionally well covered’, but recent fieldwork has yielded 22 new and five updated vice-county records (approximately one for every 34 hours of fieldwork). These new finds emphasise that knowledge of our bryophyte flora is still much less complete than that of vascular plants. In volume 2 of the Atlas (1992), Alan Crundwell’s ‘guess’ was that the distribution maps gave two-thirds of potentially available records in 10 km squares, and this now seems approximately correct for VC 1 if we make the reasonable assumption that there must still be much more left to find. However, given that VC 1 was ‘exceptionally’ well covered it would appear that the estimate of national coverage of 10 km square records should perhaps be reduced from two-thirds to a half or less.

Mapping distributions on a tetrad scale allows patterns of occurrence in relation to environmental factors to be studied in more detail than is possible using data mapped at 10 km or 5 km grid scales. Nevertheless, many of the more detailed patterns found from tetrad studies in VC 1 were to be expected, such as Grimmia trichophylla being found mainly on hard rock outcrops (and walls and gravestones) or Syntrichia ruraliformis (Tortula ruralis ssp. ruraliformis) occurring mainly on dunes. Likewise, the occurrence of certain species (Schistidium maritimum, Tortella flavovirens) only on the coast was easily predicted, although their presence beside sheltered tidal creeks well ‘inland’ was not. More surprisingly, several species appear to avoid a zone within several kilometres of exposed coasts, among them Ditrichum cylindricum, Pseudephemerum nitidum, Orthotrichum affine, Pleurozium schreberi and Ulota bruchii (U. crispa var. norvegica), possibly because they are intolerant of excessive salt-spray. When ‘standardised’ data have been recorded for all tetrads it is intended to investigate some of these distribution patterns statistically.

Mrs Jean Paton collected detailed data on bryophytes over much of Cornwall during the 1960s that she has generously made available for comparisons with the data from the past five years. Several species show marked increases or decreases over this period of approximately 30 years. Among the most obvious increases are those of Campylopus introflexus, which has ‘filled in’ a pattern of distribution the outlines of which were already nearly complete by 1960. A more surprising increase is that of Zygodon conoideus, which has become much commoner relative to Z. viridissimus. Increase of Z. conoideus elsewhere in southern England has been attributed to a reduction in sulphur dioxide pollution, but this explanation cannot be true over much of West Cornwall where there were never high levels of SO2. A recent increase in nitrogen deposition on bark seems a more likely explanation for the increased abundance of Z. conoideus, at least in Cornwall, since this species commonly occurs on nutrient-rich bark types such as that of Elder Sambucus nigra. Among the most conspicuous decreases since the 1960s are those of several species that grow on acidic rocks, especially Andreaea rothii, Racomitrium aquaticum and R. fasciculare. Increased nitrogen deposition on rocks may offer the most likely explanation for the decreases of these species.


After a long absence from England and the British Bryological Society, twelve years of study, research and field work in the Americas were summarised, with slides of bryological habitats in different areas visited.

Duke University, in the Piedmont of North Carolina on the east coast of the United States, is situated in an area of mixed deciduous and coniferous forests. The very hot, humid summers (often over 35ºC and 95% humidity) are not conducive to bryophyte growth in the heavily-shaded forests, but in more open areas such as rocky outcrops, and in the mountains, there is more diversity. Research projects included a phylogenetic study of relationships in the tropical moss family Pterobryaceae, especially of the largely neotropical genus Pireella. Characters useful for defining the species show conflicting patterns of relationships between the species. These and additional characters were studied in depth for use in cladistic analysis, allowing the relationships between the species to be resolved (Newton, 1993 & papers in prep.) This work led to further detailed study of characters such as branching architecture in pleurocarpous mosses, and of relationships between the families of the Leucodontales.

A different group of studies involved the influence of mature plants on the germination of spores and vegetative propagules, and the evolutionary role of asexual reproduction (Mishler & Newton, 1988; Newton & Mishler, 1994). Fieldwork included long trips to the south-western states and to Costa Rica, Panama and Colombia, where many fine bryophytes were seen in a range of desert, forest and montane habitats.

A move to the National Museum of Natural History (NMNH) in Washington DC, and involvement in the Biological Diversity of the Guianas Program, led to opportunities for field work in Guyana, in northern South America. Guyana currently has very large areas of untouched lowland rainforest, but like many such areas, is threatened by logging companies. Travel to the interior involves flights on small planes to very short airstrips, where cows on the runway can be a serious hazard. Further travel is then normally by dug-out canoe or on foot, and sleeping accommodation consists of hammocks, with mosquito nets an absolute necessity. One can get very stiff after eight hours sitting on a wooden plank in a canoe, broken only by portage of equipment round a waterfall. But the continually changing riverbank, with towering forest trees, riverine birds, and numerous pale yellow butterflies following each other in long undulating lines, more than compensates. Although the recorded moss diversity is not high (ca 315 species - Boggan et al., 1997) new distributional records include species very common elsewhere in the neotropics (Hypopterygium tamariscinum and Schoenobryum gardneri), indicating that more collecting needs to be done before we can be confident that the flora is well known. Floristic studies based on political rather than biogeographic regions do not necessarily reflect biological processes, so this project also includes collating records from eastern Venezuela and northern Brazil, which together with the Guianas constitute the Guayana Highlands. An additional small project at NMNH involved studies of mosses and liverworts in amber from the Dominican Republic. The age of these collections is uncertain, but they are probably between 20 and 30 million years old. However, many of the plants found are very similar or virtually identical to extant species.

The last year was spent in Mexico, working at the Institute of Ecology in Xalapa, Veracruz. The Institute is 19º north of the equator, at 1400 m altitude (slightly higher than Ben Nevis), so all year round there is normally hot sunshine, cool nights, and frequent rain and mist, making very good conditions for mosses. The natural environment includes forests of deciduous trees, such as oaks, elms and beeches, with tropical evergreen trees and epiphytic bromeliads, and at higher altitudes forests of alders and conifers. A lot of the forests have now been replaced by coffee and citrus plantations, but nevertheless the area is rich in tropical moss families such as the Meteoriaceae, Pterobryaceae, Hookeriaceae and Calymperaceae. Research carried out here, in collaboration with Dr Efrain DeLuna, was a continuation of work on morphological characters and family relationships in the Leucodontales, started at Duke, and forms part of an international collaborative project studying the relationships of all green plants. Both morphological data and molecular sequence data from rbcL were used in cladistic analyses of the pleurocarpous mosses. Part of this work involves further studies of complex character systems, including rhizoid distribution and structure, axillary hairs, the elements of branching architecture, and the ontogeny of the foot-vaginula-calyptra complex (Newton & DeLuna, papers in revision). Field trips to collect plants for research produced several new distributional records for Mexico, Veracruz or Chiapas. The most recent field trip, to the rainforests of Chiapas in south-west Mexico, involved flights in a six-seater, single-engine Cessna, to land on airstrips that looked like rough pasture complete with tall weeds and grazing horses! But solar cells on the roofs of the field stations provide the amazing luxury of 24-hour, silent electricity, with no diesel fumes or roaring generators. The representation of moss families in the areas visited seemed quite different from the Guianas, with more species of Hypnales but relatively few species of Hookeriales and Calymperaceae.

Back in England, at the Natural History Museum, work will continue on several of these projects, with collaborators both here and abroad.


Boggan J, Funk V, Kelloff C, Hoff M, Cremers G, Feuillet C. 1997. Checklist of the plants of the Guianas, 2nd edition. Washington: Biological Diversity of the Guianas Program, Smithsonian Institution.

Mishler BM, Newton AE. 1988. Influences of mature plants and desiccation on germination of spores and gametophytic fragments of Tortula. Journal of Bryology 15: 327-342.

Newton AE. 1993. Phylogenetic systematics of the tropical moss genus Pireella (Pterobryaceae: Musci). PhD thesis. Department of Botany, Duke University.

Newton AE, Mishler BM. 1994. The evolutionary significance of asexual reproduction in mosses. Journal of the Hattori Botanical Laboratory 76: 127-145.


This was a miscellany of the more unusual or attractive bryophytes to have been enjoyed by bryologists attending field courses I have taken. To qualify for consideration, a bryophyte had to have captured the attention of someone on the course and to have added to the pleasure of everyone. It was hoped that, in presenting these slides, some of that pleasure could be conveyed to BBS members present at Loughborough. From a wealth of qualifying species representing all nine of the major biome categories in Britain (Hill & Preston, 1998), just 57 were chosen from seven of those geographical assemblages.

It was argued that even the most experienced bryologist could not fail to admire the attractive colours and forms of many extremely common species. Ceratodon purpureus, Bryum argenteum, Lophocolea heterophylla, Dicranum majus and Thuidium tamariscinum were therefore included alongside such species as Orthothecium rufescens, Bryum weigelii, Cinclidium stygium, Dicranum bergeri (D. undulatum), Sphagnum lindbergii and Petalophyllum ralfsii.

A larger number of species, however, demanded attention for reasons of rarity, geographical distribution and factors involved in reproductive behaviour and/or genetic isolation. The rare Bryum cyclophyllum, for instance, had been discovered new to Wales, whereas the nationally scarce Anastrophyllum hellerianum had been found further east than any of its previous records in Wales. Not only was the gross morphology of Pellia borealis illustrated but also the cytological features which reveal its genetic isolation from the similar P. epiphylla. Their chromosomes differ numerically as well as structurally, there being 18 unique ones in gametophytes of the former species and nine in the latter (Newton, 1986). Both these species are monoecious and generally fertile but, in contrast, a number of dioecious bryophytes excite attention because of the rarity of sporophyte production, among them being fertile Plagiomnium undulatum, Climacium dendroides and Barbilophozia floerkei. Indeed, this was, I believe, the first illustration of sporophyte production by British B. floerkei.


Hill MO, Preston CD. 1998. The geographical relationships of British and Irish bryophytes. Journal of Bryology 20: 127-226.

Newton ME. 1986. Pellia borealis Lorbeer: its cytological status and discovery in Britain. Journal of Bryology 14: 215-230.


This paper, subtitled ‘NMR (nuclear magnetic resonance) fingerprinting and chemotype classification of British Plagiochilae’, contains some of the results obtained since the first applications of NMR fingerprinting of liverworts were presented in 1996 as a poster at the Centenary Symposium in Glasgow. Further information, including details of the chemical structures, can be found in the references cited.

This work has evolved out of the chemical studies of the secondary metabolites of liverworts that have been going on at Glasgow for over 25 years. In considering both our and other people’s work in this area, a recurring question was: are the secondary metabolites characteristic of the species or just of the particular specimen? Our efforts to answer this question for particular liverworts have involved using NMR spectroscopy to develop NMR fingerprinting (Rycroft, 1996). The novel aspect of this technique is to extract the plant material with deuterochloroform, the deuterated solvent that is normally used to measure NMR spectra; many benefits arise from the simplicity and directness of the procedure compared to more conventional methods (Rycroft, 1998a). In favourable circumstances, the amount of plant material required is only a fraction of many herbarium specimens, and the method opens up the possibility of undertaking comparative chemical studies using a wide range of fresh and/or herbarium material.

To illustrate what a NMR spectrum can tell us, we may consider the case of an extract from Calypogeia azurea. The blue compound responsible for the colour of the oil bodies was shown to be 1,4-dimethylazulene over 30 years ago (Meuche & Huneck, 1966); the signals in the NMR spectrum arise from different parts of this molecule and could be used to elucidate the structure. For the purposes of liverwort characterisation, identification of 1,4-dimethylazulene in the NMR spectrum is equivalent to seeing the blue oil bodies in the liverwort, and whatever weight is attached to the character of blue oil bodies could in principle also be attached to other compounds identified, with the advantage of course that they do not have to be coloured. An additional point to emphasise is that the case of C. azurea is one where a secondary metabolite can be demonstrated unequivocally to be localised in the oil bodies. It is commonly assumed that this is the norm, despite the fact that what are regarded as normal liverwort compounds have also been isolated from liverworts that have no oil bodies.

Inter alia, we have been using NMR fingerprinting to study all (except the most recently discovered) British species of Plagiochila. Our work on P. spinulosa started originally in 1982, and the poster at the Centenary Symposium reported that there was little chemical variation between different samples of P. spinulosa. Subsequent work (Connolly et al., 1999) has resulted in identification of more of the components: eight are 9,10-dihydrophenanthrenes (DHPs), two are methyl orsellinates, and one is a bibenzyl. In comparison with P. spinulosa, the P. punctata spectra have fewer signals; the major components have been identified, and in addition to three of the DHPs found in P. spinulosa there is a flavonoid.

One aspect of chemical interest in Plagiochila killarniensis is the characteristic smell. The proton NMR spectrum quickly revealed that the extract was dominated by methyl everninate, which is known in the perfumery world as one of the odour-impact compounds of oakmoss (the lichen Evernia prunastri). Another compound present is b-phellandrene, a monoterpenoid with a strong smell, present in essential oils from various Eucalyptus, Abies and Pinus species. b-Phellandrene is also present in Plagiochila spinulosa and P. punctata, and is responsible for the smell obtained when fresh material is crushed in the field, whereas methyl everninate is responsible for what makes P. killarniensis distinctive after it has been collected. Our isolated methyl everninate contained two minor constituents that were also methyl orsellinate derivatives, one of which we already knew from P. spinulosa. In six Scottish specimens and four from the Azores, another minor component that was always present was a more unusual type of compound, a 3-benzylphthalide that we have named ‘killarniensolide’ (Rycroft et al., 1999). The structure of killarniensolide is related to lunularic acid, the dormancy-inducing factor that seems to be ubiquitous at low levels in liverworts. There is also a structural relationship with the bibenzyl that we isolated from P. spinulosa. P. killarniensis did not contain any of the DHPs from P. spinulosa, but two of the minor components corresponded to two of the DHPs reported from a Costa Rican Plagiochila (Anton et al., 1997).

In discussions with the Saarbrücken and Göttingen bryophyte groups we learnt of the synonymy of P. killarniensis with the Neotropical species P. bifaria (Heinrichs, Grolle & Drehwald, 1998). We have now examined several Neotropical specimens, but the extract of the only sample of P. bifaria was dominated by a compound that is suspected to have arisen from fungal contamination. Until more material becomes available, the secondary metabolites of Neotropical P. bifaria remain obscure. The only work in the literature is a GC-MS study of a single Peruvian specimen that appears to differ chemically from the ten Scottish and Azores specimens studied by us. Two Madeiran samples sent from Göttingen were very similar chemically to Scottish and Azores material; however a third sample (Drehwald 960277) turned out to be very different. The NMR spectrum showed mainly a ca 1:1 mixture of two compounds, one of which was one of the DHPs in P. killarniensis and the other the bibenzyl derivative in P. spinulosa. Morphological examination of the plant then revealed that it was very different from P. killarniensis as we know it in Britain: the leaves were relatively longer, were not truncate, and the line of teeth on the postical leaf margin continued around the apex and sporadically along the antical leaf margin. An interesting development since the AGM is the discovery that the extract of a 45 year-old sample of P. sharpii (R.M. Schuster 40723 ex GL) contains a ca 1:1 mixture of the same two compounds. These observations may be placed in perspective by noting that chemically the difference between Drehwald 960277 and P. killarniensis is at least as great as that between P. spinulosa and P. punctata.

The NMR spectra of extracts of three samples of Plagiochila exigua from different sites in Glen Creran were fortuitously simple in the methoxyl region in giving one large peak. In the aromatic region, things were not so simple, but a simulated spectrum could be calculated to match the line positions of the experimental spectrum and show that the new compound 3,4-dihydroxy-3´-methoxybibenzyl was present (Rycroft, Cole & Aslam, 1998). This result differs from the only other work reported for P. exigua, from a GC-MS study of a sample from Peru: none of the compounds was identified, although two of them were subsequently recognised by Asakawa & Inoue in P. spinulosa (from Belgium), and we have since shown that they are two of the methyl orsellinate derivatives mentioned earlier. A study of a wider range of material would be necessary to discover if there is any significance in these differences.

Historically at least, it appears that Plagiochila atlantica was liable to be confused with P. spinulosa. In 1996 the chemistry of only one sample of P. atlantica, from its now classic locality of Ariundle, had been investigated. It was shown that it was very different from P. spinulosa in that P. atlantica contained plagiochiline C, a 2,3-secoaromadendrane sesquiterpenoid, rather than DHPs (Rycroft, 1996). Many 2,3-secoaromadendranes are easy to recognise in a proton NMR spectrum because they give a doublet for H-2 and a singlet for H-3 in a characteristic region of the spectrum. Subsequently we have extended the range of the study to include samples from as far as the English Lake District to the south and the Gruinard River in Wester Ross to the north, and discovered a uniform chemical pattern consistent with the idea that the population consists of one clone only, with reproduction occurring asexually.

A set of signals from a second compound was consistently present at ca 10% of plagiochiline C; the compound responsible is a new sesquiterpenoid alcohol that we have called ‘atlanticol’ (Rycroft & Cole, 1998b). It is a derivative of bicyclogermacrene, the sesquiterpenoid found most frequently in liverworts (and also present in P. atlantica); a closely-related compound has been reported by Hashimoto et al. (1993) from Japanese P. fruticosa, but without details or characterisation data.

P. carringtonii also contains 2,3-secoaromadendranes. The NMR spectra of five extracts are all very similar, and in this case there are two H-3 singlets, two H-2 doublets and one aldehyde and one methyl ester signal forming two sets of signals in the ratio 2:1. These signals do not correspond to any of the known plagiochilines, and the new compounds responsible were isolated using thin layer chromatography (Rycroft, Cole & Lamont, to be submitted). They are a pair of closely-related plagiochilines, T and U, that have one of the methyl groups on the cyclopropane ring of the 2,3-secoaromadendrane skeleton oxidised to either an aldehyde or a methyl ester respectively. Previously reported plagiochilines do not have these methyl groups oxidised beyond the alcohol stage, but otherwise T and U are typical plagiochilines. Therefore the Plagiochila that for a long time was placed in Jamesoniella has secondary metabolites that, although new, are of a type characteristic of other Plagiochila species. Lewis (1970) demonstrated a similar point by studying sugar alcohols.

Plagiochila porelloides has been recorded as new for China from Changbai Mountain, at a latitude of ca 42ºN and less than 350 km west-south-west of Vladivostok (Söderström et al., 1999). In comparison with the foregoing, the NMR spectra of this Chinese material are more complicated, but it was possible to determine that there was a major compound that did not correspond to the known plagiochilines. Again, the new compound, plagiochiline V, was isolated, and we have been able to propose a structure where one half is the same as in the known plagiochiline M, but the other half is novel. British P. porelloides also contains 2,3-secoaromadendranes, but we have still to complete our studies and confirm whether it contains plagiochiline V. Our preliminary studies of British P. asplenioides and P. britannica have also found evidence of 2,3-secoaromadendranes.

Our present knowledge, though incomplete, is sufficient to produce a provisional classification of British Plagiochilae based on the chemotypes of Asakawa (1995). In the largest group, characterised by 2,3-secoaromadendranes, we can include P. asplenioides, P. atlantica, P. britannica, P. carringtonii and P. porelloides. P. exigua can be placed in the bibenzyl group, but we need to propose a new chemotype group (Rycroft, 1998b) characterised by the presence of DHPs (with possible sub-divisions) to accommodate P. killarniensis, P. punctata and P. spinulosa. This DHP group has Neotropical connections but DHPs have not (yet) been reported from any Asiatic Plagiochila. Of particular note is the observation that, on chemical grounds, P. atlantica and P. spinulosa would be classified in different sections.

Our work has by no means been confined to Plagiochila and one example concerns Cryptothallus mirabilis (Rycroft & Cole, 1998a). A specimen from near Glasgow gave a remarkably clear NMR spectrum (Rycroft, 1998a) of a new pinguisane sesquiterpenoid derivative, a result of chemosystematic interest in view of the fact that the first pinguisane was isolated from Aneura pinguis. Investigation of a second specimen involved a return visit to the site in Glen Creran where Jeff Duckett had unearthed plants in 1996. The new pinguisane was detectable in this material but the dominant component in this case was a sesquiterpenoid with a different carbon skeleton, a cyclocuparanol that has also been found in Marchantia polymorpha (the cyclocuparanol was also detectable in the first specimen of C. mirabilis). Investigation of further specimens is a prerequisite to attributing any significance to this difference.

Application of NMR fingerprinting has enabled us to obtain detailed chemical results from a range of specimens that were unimaginable until very recently. It would be interesting to apply these methods in parallel with DNA techniques to phytogeographical problems.


Others involved in this work are cited in the references and I should like to thank them, and in particular my GC–MS colleague Dr John Cole, for their collaboration. In addition I am grateful to the many BBS members who have given much advice and encouragement to a comparatively recent recruit to the Society.


Anton H, Kraut L, Mues R, Morales Z MI. 1997. Phenanthrenes and bibenzyls from a Plagiochila species. Phytochemistry 46: 1069–1075.

Asakawa Y. 1995. Chemical constituents of the bryophytes. In: Herz W, Kirby GW, Moore RE, Steglich W, Tamm Ch, eds. Progress in the chemistry of organic natural products. Vol. 65. Wien: Springer, 490.

Connolly JD, Rycroft DS, Srivastava DL, Cole WJ, Ifeadike P, Kimbu SF, Singh J, Hughes MP, Thom C, Gerhard U, Organ AJ, Smith RJ, Harrison LJ. 1999. Aromatic compounds from the liverwort Plagiochila spinulosa. Phytochemistry: in press.

Hashimoto T, Asakawa Y, Nakashima K, Tori M. 1993. Chemical constituents of 25 liverworts. Journal of the Hattori Botanical Laboratory 74: 121–138.

Heinrichs J, Grolle R, Drehwald U. 1998. The conspecificity of Plagiochila killarniensis Pears. and P. bifaria (Sw.) Lindenb. Journal of Bryology 20: 495–497.

Lewis DH. 1970. Chemotaxonomic aspects of the distribution of acyclic sugar alcohols in leafy liverworts. I. Chemical evidence for the taxonomic position of Plagiochila carringtonii (Balfour) Grolle. Transactions of the British Bryological Society 6: 108–113.

Meuche D, Huneck S. 1966. Azulene aus Calypogeia trichomanis. Chemische Berichte 99: 2669–2674.

Rycroft DS. 1996. Fingerprinting of plant extracts using NMR spectroscopy: application to small samples of liverworts. Chemical Communications, 2187–2188.

Rycroft DS. 1998a. Chemical comparison of liverworts using NMR spectroscopy. Journal of the Hattori Botanical Laboratory 84: 105–111.

Rycroft DS. 1998b. Plagiochila atlantica F. Rose newly identified in England: chemotype classification. Journal of Bryology 20: 240–242.

Rycroft DS, Cole WJ. 1998a. 15-Acetoxypinguisone and a cyclocuparanol from the liverwort Cryptothallus mirabilis Malmb. Journal of Chemical Research (Synopses), 600–601.

Rycroft DS, Cole WJ. 1998b. Atlanticol, an epoxybicyclogermacrenol from the liverwort Plagiochila atlantica F. Rose. Phytochemistry 49: 1641–1644.

Rycroft DS, Cole WJ, Aslam N. 1998. 3,4-Dihydroxy-3´-methoxybibenzyl from the liverwort Plagiochila exigua from Scotland. Phytochemistry 49: 145–148.

Rycroft DS, Cole WJ, Aslam N, Lamont YM, Gabriel R. 1999. Killarniensolide, methyl orsellinates and 9,10-dihydrophenanthrenes from the liverwort Plagiochila killarniensis from Scotland and the Azores. Phytochemistry: in press.

Söderström L, Rycroft DS, Cole WJ, Wei S. 1999. Plagiochila porelloides (Torrey ex Nees) Lindenb. from Changbai Mountain, new to China, with chemical characterization and chromosome measurements. Bryobrothera: in press.


The Sefton Coast, north-west England, is historically a well-known stronghold for several important bryophytes. In 1995 two surveys by Dr M. Newton (JNCC Report No 239) of Red Data List bryophytes and stoneworts found only four of the seven previously recorded Red Data List bryophytes. Bryum warneum and Petalophyllum ralfsii were found to have reasonably strong populations. However, B. neodamense was sparse, and B. mamillatum (tentatively identified) was confined to a single small site - its only recent British record. B. knowltonii, B. uliginosum and B. marratii were not refound. It is probable that B. marratii is now extinct on the Sefton Coast.

Further survey work on Petalophyllum was undertaken by myself in 1997, with some incidental records also made of B. neodamense. Populations of Petalophyllum were confirmed to be strong, albeit rather localised. B. neodamense was found to be somewhat more frequent than previously thought. Detailed maps were made of all known populations using a computerised Geographic Information System (GIS). This system allows data such as species location and abundance to be entered into a database format and then displayed and analysed by their geographic attributes. Aerial photography, either displayed on screen or printed out, is a rapid and accurate means of location in an otherwise featureless landscape. Species records can be represented by a chosen symbol on enlarged aerial photos. Relocation of sparse or scattered records benefits from the high resolution of the system, providing accuracy of a metre or less.

Primarily as a guide for the land managers, a report was produced giving precise locations and photographs of Petalophyllum populations with an estimate of plant numbers (based on counts of individual thalli) plus other field notes. This report also listed possible threats and management recommendations for each population. The report also assisted in providing the information required by the Sefton Coast LIFE Project to produce a conservation strategy for Petalophyllum at Sefton.

Current management of dune slack habitat on the Sefton Coast rarely shows any consideration toward the conservation of bryophytes. This may be a consequence of the lack of recording effort and limited awareness of the conservation requirements of bryophytes, but may also reflect the level of importance given to bryophytes by conservation managers compared with other groups such as herptiles and birds. Natterjack Toad conservation is the main reason for management of many dune slacks in the frontal dunes of the Sefton Coast. This often involves heavy machinery re-profiling or excavating new dune slacks, and the use of herbicides to treat scrub re-growth. While this form of management is likely to continue in the short term, it is hoped that future management will give greater emphasis to dune geomorphological processes. Such management may include the implementation of appropriate grazing regimes and the spatial and temporal zoning of activities associated with recreation, such as beach car parking. The environmental impact of beach cleansing activities on dune processes also requires careful consideration. Most dunes in Britain are mature and in a phase of sediment recycling. Natural dune slack creation and development should therefore be encouraged as a desirable quality.

Footpath management is also an important consideration, particularly in areas of little or no grazing. Light levels of trampling can keep some dune slack areas open in structure that may otherwise become dominated by scrub and course grass species. While it is necessary to ensure heavy trampling does not destroy valuable habitat, the construction of boardwalks should only be undertaken after an adequate impact assessment and for overriding recreational management reasons. This is because boardwalks cover valuable habitat, remove trampling pressure, and encourage scrub to spread right up to the edge of the boards.

The most critical factor on the Sefton Coast for several of the Red Data Book bryophytes is water quality. The stronghold for Petalophyllum and several other bryophytes at this site is flooded regularly by eutrophic water from a nearby lake. Aside from the direct effects this may be having on these species, indirect effects include an increased vigour of vascular plants and growth of an algal mat which coats the ground when the water subsides. This algal mat then either hardens on drying or is puddled into a muddy mess.

An innovative recording approach is required for important dune bryophytes in the face of the environmental stress and rapidly changing conditions that occur in sand dune systems. This must aim to provide an adaptable and updatable system with a level of detail that can translate the information gathered in a manner that is accessible to land managers.


In warm autumn sunshine, 24 of us explored the grassland and woods near Charnwood Lodge (VC 55). The way from the cars to the woods led past a dark Precambrian rock outcrop with Barbilophozia atlantica*, Ptilidium ciliare and Racomitrium heterostichum, showing at once that this was not an ordinary part of the English Midlands. Our leader, Dennis Ballard, walked us briskly for about half a mile to Burrow Wood, but we tarried by the way and found Ulota bruchii (U. crispa var. norvegica) In the wood, we saw several plants that are often thought of as western, including Cryphaea heteromalla* on elder, Hypnum andoi (H. mammillatum) on a stone wall, and Lejeunea lamacerina*, Scapania undulata and Heterocladium heteropterum var. heteropterum* in a small stream gully. Dicranum tauricum on dead wood, however, indicated that we were still some distance from the west coast, as did the presence of plentiful Plagiothecium curvifolium and Orthodontium lineare. In nearby Cat Hill Wood, there was a steep shaded rock outcrop with more Barbilophozia atlantica and Racomitrium heterostichum and also Barbilophozia attenuata, B. floerkei and plenty of a Cephaloziella without perianths.

Another party broke away at an early stage and visited some woodland near a reservoir south-west of Charnwood Lodge. There was some damp ground with Polytrichum commune and three species of sphagna. They also found Brachythecium plumosum and Scapania undulata. Richard Fisk and Rod Stern found Calypogeia arguta in a ditch in a nearby plantation, but did not collect it because they had no reason to suppose that the species would be bracketed in the new Census Catalogue.

Two parties went to look for Syntrichia (Tortula) amplexa. S. amplexa was duly refound in three separate places, none of which was its locus classicus, which has now become a deep hole in the ground with water at the bottom. On the advice of Dennis Ballard, one group went to look for it in an opencast mine area east of Moira (SK3215). After much searching, they eventually found it in the lower part of the site, and in the process recorded Cephaloziella divaricata, C. hampeana*, Fossombronia incurva*, Riccia sorocarpa, Aloina ambigua (A. aloides var. ambigua), Bryum dunense and a scrap of Sphaerocarpos, which was not identifiable to species. The other party found S. amplexa in small quantity near Lount (SK3819) and more plentifully in a ditch by a stone and earth dump near Boothorpe (SK3117). In the last locality, Harold Whitehouse found Ephemerum serratum var. serratum* with immature capsules on bare soil near a stream and successfully grew it on to achieve an identification.

While the cognoscenti were especially gratified to see that great speciality of the area, Syntrichia amplexa, there was pleasure for all in the beauties of Charnwood Forest, and for some in making new records for the vice-county. Some of these update old records that had been bracketed in the new Census Catalogue, but others are completely new. We are grateful to Dennis Ballard for organising an interesting meeting and for showing us that there are many good bryophytes to be found in the centre of England.


Bryological Workshop 1998

Beaulieu, Hampshire, 7-8 November

Local Secretary: Dr. June Chatfield, Anglefield, 44 Ashdell Road, Alton, Hampshire, GU34 2TA. Tel: 01420 82214.

This will be an workshop meeting with the emphasis on introducing beginners to bryophytes and bryological techniques. There will be excursions into the New Forest, so there is sure to be something to interest everyone.

Copyright © British Bryological Society .