The Symposium was chaired by the President and six presentations were made. During the lunch break a short tour of the RBGE bryophyte herbarium was led by David Long and Sally Rae, who had laid out a selection of specimens from the bryological collections: specimens from R.K. Greville’s herbarium, including some original paintings of mosses (e.g. Aplodon wormskioldii); some historical type specimens, such as mosses collected in New Zealand in the late 18th century by Archibald Menzies, including many types which are being curated and databased; and recent acquisitions, such as a collection of British mosses made by H.N. Dixon, and specimens from the herbaria of Jean Paton, Cliff Townsend and Alan Crundwell.
DR MONTSERRAT BRUGUÉS (Autonomous University of Barcelona, Spain): The Bryophytic Richness of Spain
The Spanish bryoflora consists of 807 moss species, 256 liverworts and five hornworts. The number of bryophytes, and particularly mosses, recorded from Spain has recently increased considerably in parallel with the increase in the number of researchers, and especially due to the study of arid zones, of great interest but little known until now. These zones have yielded the majority of the species new to science, although there are regions where much greater study is required than that undertaken to date.
Climatic variety, accompanied by altitudinal differences within the Spanish territory, determines the interesting and diverse bryophyte flora. The flora includes arctic-alpines, characteristic of the higher areas of the Pyrenees, such as Gymnomitrion corallioides, Marsupella brevissima and Pleurocladula albescens, or such as Anthelia juratzkana and Amphidium lapponicum which are also found in the Sierra Nevada, species with oceanic distributions, such as Sphagnum pylaesii, Jubula hutchinsiae and Breutelia chrysocoma, and steppe species characteristic of arid, continental zones, such as Crossidium aberrans, C. seriatum, Tortula revolvens var. obtusata and Phascum vlassovii.
There are about 20 Iberian endemics, including Triquetrella arapilensis, Anomobryum lusitanicum, Schizymenium pontevedrensis, Pterygoneurum sampaianum and Phascum cuynetii. Some of these, such as Acaulon casasianum, Didymodon bistratosus, Orthotrichum ibericum or Phascum longipes, may eventually be found in other Mediterranean countries, as is the case with Acaulon fontiquerianum and some Orthotrichum species, or else in the Canary Islands, as with Goniomitrium seroi. Mediterranean endemics and Iberian Macaronesian endemics are also abundant. The catalogue also contains species with interesting disjunctions in their distributions, such as Oedipodiella australis, Antitrichia californica, Claopodium whippleanum, Schistidium occidentale and Phascum piptocarpum.
DR HEINJO DURING (University of Utrecht, The Netherlands): The Diaspore Bank of a Zimbabwean Savannah
The ‘Matopos Sandveld Fire Plots’ in the savannas of south Zimbabwe are experimental plots with a consistent management (12 different regimes) since 1947. The above-ground vegetation appears to contain few bryophytes: in February 1997, in the middle of the rainy season, only isolated plants of Exormotheca holstii, two Riccia species, two Archidium species and a Bruchia were found. However, from superficial soil samples taken in plots with four different management regimes, including annual fire, at least 11 liverworts and hornworts and 21 mosses emerged, often in large numbers. Many of the species found are hypothesised to possess an ‘episodic’ strategy much like that described for Physcomitrium sphaericum. Perhaps for that reason, the assemblage contained several species new to Zimbabwe or new to Africa, including Micromitrium tenerum, and at least one taxon new to science: Neophoenix matoposensis R.H. Zander & During gen. et spec. nov. Annual burning of the above-ground vegetation did not appear to be harmful to the diaspores in the soil.
DR ANGELA E. NEWTON (Natural History Museum, London): Mosses of the Maya Mountains: Research and Exploration in Central America
The Neotropical area, including Mexico, Central America, the Caribbean and northern South America, is probably one of the better known tropical regions for bryophytes, since there are now several complete or partial floristic works available. There is also a significant number of national and international researchers working on the floristics and systematics of bryophytes in the region. However, there are still enormous gaps in our knowledge of the distribution of mosses and liverworts in most of the countries involved, and many of the taxa are poorly understood, so that a large amount of research is still necessary.
Key biodiversity issues include a) cataloguing what is present, b) providing ‘quality control’ for names so that databases contain meaningful information, c) investigating the natural distribution and ecological associations of bryophytes before too much more of this information is destroyed by development, and d) developing keys, guides and inventories to enable ecologists, conservationists and others to undertake research and implement management plans. Systematic taxonomic revisions are necessary to provide a better estimate of the number and nature of species and higher level taxa in the area, and to provide a basis for evolutionary and biogeographical research into the processes influencing the origin and maintenance of biological diversity.
One of the countries in this area that is very poorly known is Belize, on the southern side of the Yucatan Peninsula between Mexico, Guatemala and the Caribbean. It is mostly lowland, but in southern Belize there is a large limestone and sandstone plateau with some peaks reaching over 1000 m (Doyle’s Delight is the highest point at 1140 m). There have been few visits by bryologists to the country, possibly because the low altitudes promise little diversity in contrast to the high peaks in adjacent areas. Recent checklists give 250 species of moss (Townsend & Allen, 1998) and 65 species of liverwort (Whittemore & Allen, 1996). In connection with other bryological research projects, Fred Rumsey and I undertook to survey the bryophytes around the Las Cuevas Research Station, which is run jointly by the Natural History Museum, London and the Forestry Department of Belize. The area around the station is rather uniform, being situated in mature secondary forest on the limestone plateau at about 450 m, surrounded by numerous small hills rising to 800 m. There is little surface water in the immediate area, but frequent heavy rain and overnight mist are conducive to bryophyte growth.
Preliminary results from our three-week visit produced a list of 132 moss species and at least 43 hepatic species. This represents 32% of the Townsend & Allen (1998) moss list, but with at least 19 additional species and five additional families. The liverworts collected have only been partially identified, but represent 44% of the genera in Whittemore & Allen (1996), with at least 14 additional genera. Interesting moss taxa new to the country, although common elsewhere in the neotropics, include Cryphaeaceae (Schoenobryum concavifolium), Anomodontaceae (Anomodon rostratus, A. attenuatus) and Fabroniaceae (Fabronia ciliaris var. polycarpa). In the liverworts, a large number of epiphyllous taxa were collected, including representatives of the genera Aphanolejeunea, Colura and Leptolejeunea. Terrestrial taxa included Marchantiaceae (Marchantia, Dumortiera and Asterella), Aneura, and also the hornwort Notothylas.
This number of additional taxa found in the course of a short visit to a rather uniform area indicates that the bryoflora of Belize is still far from completely known, and that additional visits here, especially to a range of different habitats and altitudes, are likely to produce additional records. The liverworts in particular would repay further work.
Townsend, C., Allen, B. 1998. A checklist of the mosses of Belize. Tropical Bryology 15: 89-100.
Whittemore, A.T., Allen, B. 1996. The liverworts and hornworts of Belize. Bryologist 99: 64-67.
PROF. JEFFREY G. DUCKETT (Queen Mary College, University Of London) & PROF. ROBERTO LIGRONE (Seconda Università Di Napoli, Caserta, Italy): What we Couldn’t Have Done if we’d Stayed in Europe: Selection and Serendipity in the Southern Hemisphere!
We are indebted to David Long for dreaming up the exploration theme for the 2002 BBS Symposium and for inviting us to speak. This has provided a unique opportunity to pause and to reflect on the various fruits of our visits, over the last decade or so, to Lesotho, South Africa, Borneo, Uganda and New Zealand. This turns out to be a curious tale of setting out with one set of objectives and discovering very different things.
The first question that immediately comes to mind is: why should two botanists, whose research can best be described as the cell, developmental and evolutionary biology of bryophytes (with a bit on the side on protonemata and bryological exploration of uncharted territories (Hodgetts et al., 1999)), want to travel to the southern hemisphere in the first place?
Together, Britain and Italy have remarkably good bryophyte floras for our kind of work; most groups are well represented and can be fixed immediately after collection. The most notable lacunae are Takakia, giant members of the Polytrichales (Dawsonia, Dendroligotrichum), Schuster’s primitive metzgerialean taxa (Treubia, Phyllothallia, Allisonia, Verdoornia) and those with highly differentiated gametophytes (Symphyogyne, Hymenophyton), Monoclea and Neohodgsonia in the Marchantiales, only one species of Haplomitrium, and the hornwort genera Dendroceros and Megaceros.
Thus in the mid 1990s we set off to Lesotho to study the ontogeny of the water-conducting cells in Symphyogyne and to make comparative observations on fungal endophytes in the Marchantiales. The discovery that the formation of the large pits in the former is associated with callose (Ligrone & Duckett, 1996), a capricious carbohydrate better known in association with phloem sieve plates in angiosperms, was the catalyst for wide-ranging comparative studies on water-conducting elements in bryophytes. Ultimately, this has led to the conclusion that these conduits are polyphyletic in both liverworts (Calobryales, Metzgeriales) and mosses (Takakia, Bryales and Polytrichales) (Ligrone et al., 2000, 2002). Developmental and, most recently, immunocytochemical evidence that moss hydroids are not homologous with tracheary elements has major implications for interpreting vascular plant lineages.
From the same expedition the chance observation that the central thallus cells in Asterella contain longitudinal arrays of endoplasmic microtubules associated with a pleomorphic vacuole system (Ligrone & Duckett, 1994a) caused us to re-examine the cytology of food-conducting leptoids in mosses (Ligrone & Duckett, 1994b). This highly characteristic ‘food-conducting cytology’ has now been found, not only in polytrichalean leptoids, but also in other mosses such as Neckeraceae, Orthotrichaceae, Hookeriaceae and most notably Sphagnum (Ligrone & Duckett, 1998), previously thought to lack specialised conducting elements. It also occurs in moss caulonemata and rhizoids (Duckett et al., 1998), in the axes of Takakia and Haplomitrium, and is widespread in the thalli of Marchantiales.
Our studies on conducting elements have extended to a re-examination of the possible functional significance of dimorphic rhizoids in the Marchantiales, apparently last studied by Kamerling (1897). Whereas smooth rhizoids are living and frequently contain hyphae of the fungal endophytes, the pegged variety are dead and function as an internalised, external water-conducting system in carpocephala grooves. The pegs prevent total collapse when the thalli dry out and permit recovery after rehydration. The absence of pegged rhizoids in Monoclea, Neohodgsonia and some aquatic Riccia species suggests that this is a derived condition – a conclusion in line with recent molecular evidence (Wheeler, 2000) but at variance with earlier Schusterian notions.
One of our principal original aims was the exploration of fungal endophytes in hepatics (Read et al., 2000). While some of our observations have been closely in line with predictions (e.g. the widespread occurrence of Endogonaceae in the Marchantiales, including Monoclea), others were unexpected, for example fungi in the rhizome systems of the Pallavicinaceae. Most remarkably, the thalli of Treubia contain both extracellular and intracellular endogaceous fungi, and the same are confined to the outer cell layers in the mucilage-invested ‘roots’ of Haplomitrium.
This distillation of some of the cytological gems originating from the southern hemisphere is just a preliminary progress report of the results from our travels. Just as floristic collecting expeditions result in hundreds of specimens requiring many years to identify, ours produce dozens of resin-embedded specimens – the bryological equivalent of sculptures waiting to be carved. So, when asked where we might go next, the answer should really be ‘absolutely nowhere – not until we’ve looked at most of what we’ve got already’. Nearing the top of the pile are investigations into the ontogeny of stomata and intercellular spaces in hornworts and mosses. Just as hydroid/tracheid homology was a key question in moss-tracheophyte evolution, monophyly versus polyphyly of stomata has now become the vexed issue in setting the basal lineages of land plants (Raven, 2002).
We thank the British Council, The Royal Society and the Natural Environment Research Council without whose financial support this work would not have been possible. Equally crucial have been laboratory facilities at the Universities of Lesotho and Canterbury, Christchurch, New Zealand, and the camaraderie of the worldwide bryological community who shared experiences including men with guns, bribery of officials, life-threatening weather, kea-damaged cars, and a universal hatred of possums in New Zealand.
Duckett JG, Schmid AM, Ligrone R. 1998. Protonemal morphogenesis. In: Bates JW, Ashton NW, Duckett JG, eds. Bryology for the twenty-first century. Leeds: Maney, 223-246.
Hodgetts NG, Matcham HW, Duckett JG. 1999. Bryophytes of Lesotho and South Africa: results of the 1995 British Council Expedition. Journal of Bryology 21: 133-155.
Kamerling Z. 1897. Zur Biologie und Physiologie die Marchantiaceen. Flora 84: 1-68.
Ligrone R, Duckett JG. 1994a. Thallus differentiation in the marchantialean liverwort Asterella wilmsii (Steph.) with particular reference to longitudinal arrays of endoplasmic microtubules in the inner cells. Annals of Botany 737: 577-586.
Ligrone R, Duckett JG. 1994b. Cytoplasmic polarity and endoplasmic microtubules associated with the nucleus and organelles are ubiquitous features of food conducting cells in bryalean mosses (Bryophyta). New Phytologist 127: 601-614.
Ligrone R, Duckett JG. 1996. Development of water-conducting cells in the antipodal liverwort Symphyogyne africana (Metzgeriales). New Phytologist 132: 603-615.
Ligrone R, Duckett JG. 1998. The leafy stems of Sphagnum (Bryophyta) contain highly differentiated polarized cells with axial arrays of endoplasmic microtubules. New Phytologist 140: 567-579.
Ligrone R, Duckett JG, Renzaglia KS. 2000. Conducting tissues and phyletic relationships of bryophytes. Philosophical Transactions of the Royal Society B 355: 795-814.
Ligrone R, Vaughn KC, Renzaglia KS, Knox JP, Duckett JG. 2002. Diversity in the distribution of polysaccharide and glycoprotein epitopes in the cell walls of bryophytes: new evidence for the multiple evolution of water-conducting cells. New Phytologist 156: 491-508.
Raven JA. 2002. Tansley Review 131. Selection pressures on stomatal evolution. New Phytologist 153: 371-386.
Read DJ, Duckett JG, Francis R, Ligrone R, Russell A. 2000. Symbiotic fungal associations in ‘lower’ land plants. Philosophical Transactions of the Royal Society B 355: 815-832.
Wheeler JA. 2002. Molecular phylogenetic reconstructions of the marchantioid liverwort radiations. Bryologist 103: 314-333.
DR MARK HILL (Centre for Ecology and Hydrology, Monks Wood, Huntingdon) & BRYAN EDWARDS (Dorset Environmental Records Centre, Dorchester): A Very Slow Exploration: 40 Years of Bryology in Dorset, England
Dorset (Watsonian vice-county 9) is on the south coast of England, just west of the line separating English Nature’s South-West region from the South-East region. It measures about 80 km from east to west and 60 km from north to south. Topographically, it is dominated by a ridge running south-west to north-east. The highest hill, Pilsdon Pen, is 277 m high. Geologically, the county is underlain by relatively young rocks, mostly of Jurassic and Cretaceous age. Tertiary sands, clays and gravels characterise the Poole Basin, an area of heathland on acid soils in the south-east near Poole. Annual rainfall is lowest on the coast, being 700 mm on Portland, rising to about 1300 mm on the chalk ridge.
Progress of recording
Mark Hill became interested in bryophytes in 1962 when he began recording them in the west of the county near Wootton Fitzpaine. Initially, his aim was to learn to identify the species and to record them in west Dorset for the newly-launched BBS Mapping Scheme. During the following 40 years, the project transformed itself gradually from a local flora of the west of the county (1966), via a flora of the whole county, based on 10-km squares (1970), to a tetrad flora (1995). BBS meetings were held at Lyme Regis (1969), Wareham (1977) and Weymouth (1995). During the 1970s, Monica Milnes-Smith recorded bryophytes extensively near Child Okeford. In the 1980s, Rod Stern sent numerous records from the east of the county. In the 1990s, Bryan Edwards became active and kept systematic records from sites all over the county. In 1995, he and Mark Hill decided to pool their records and (finally) to write up a bryophyte flora of the county. This is due to be published by Dorset Environmental Records Centre as a book.
In the 1960s there was already a long history of recording in the county, starting with Leptodon smithii, found at Weymouth in 1800. There was no further activity until 1867, but thereafter mosses were recorded rapidly, with half the moss flora discovered by 1890. Liverworts trailed and did not reach the halfway mark until 1935. Notable finds during the 19th century were Myrinia pulvinata (1879), Sphagnum pulchrum (1880) and Eurhynchium meridionale (1881). The discovery of E. meridionale, one of only two bryophyte species confined in England to Dorset, was delayed because E.M. Holmes misidentified it as E. striatulum. Its other finder, Sir William Medleycott, got a correct identification from H.H. Wood, but Wood died shortly afterwards and the record did not come to light until 1917. From that time onwards, bryologists have regularly visited Portland, where Southbya nigrella was found new to Britain in 1921.
During the 1930s, Eustace Jones recorded bryophytes on Studland Heath and in several woods. Many of his best discoveries were liverworts, including Jungermannia hyalina, Lophozia incisa, Marsupella funckii and Riccia huebeneriana, which have never been refound. Since 1945, many other interesting plants have been added to the county list, of which Plagiochila norvegica must surely be the most remarkable. It is known nowhere else in Britain, and in Dorset is confined to a single stone.
Composition of the flora
With 95 liverworts, three hornworts and 329 mosses, the Dorset bryophyte list comprises about 42% of the British and Irish total. Arctic-montane species are completely lacking, but there are 18 Boreal and Boreo-arctic montane species, including both calcicoles, such as Leiocolea badensis and Thuidium abietinum, and calcifuges, such as Calypogeia sphagnicola and Pogonatum urnigerum. In comparison with the average British county, Dorset has more Mediterranean-Atlantic and Submediterranean-Subatlantic species than the average, with 65% of the total.
The most similar vice-county is South Hampshire (VC 11), which has a nearly equal number of species. Among the species that occur in Dorset but not in any adjacent county, perhaps the most surprising is Sphagnum pulchrum. It is common in Dorset but does not quite reach the Hampshire border. Myrinia pulvinata and Weissia rostellata are also not known from any adjacent counties but are more widespread in England. In the other direction, Campylophyllum calcareum and Orthotrichum stramineum are known from all six adjacent vice-counties but not from Dorset.
Changes in the flora
There are just five recorded aliens in the Dorset bryoflora. Campylopus introflexus, now very common on heathland and occasional elsewhere, was first found in 1950. Orthodontium lineare was found in 1961 but has remained uncommon. Didymodon umbrosus, Lophocolea bispinosa and Telaranea murphyae were discovered in the last 15 years. Only L. bispinosa appears likely to become common. It is confined to sites disturbed by quarrying or military activity, and has also been found in adjacent parts of Hampshire.
A few species appear to be increasing, but older records are not in the main good enough for us to be certain. Didymodon nicholsonii was first found in 1980 but was not seen again until 2000. It now appears to be common in the county, especially on tarmac in villages. Although it is rather inconspicuous, it would almost certainly have attracted our attention if it had been common in the past. Likewise Cololejeunea minutissima, now found frequently inland, was formerly almost confined to the coast.
A rather larger number of species appear to be declining. Some, such as Bartramia pomiformis, Dicranum spurium and Splachnum ampullaceum, are generally decreasing in southern England. More surprising are Plagiothecium denticulatum and Pohlia nutans, which were frequently found in the 1960s, but since 1980 have been seen in only seven and three tetrads respectively. At least seven species are thought to have been short-persisting casuals in the county. Notable among these are Riccardia palmata and Scapania gracilis, found in 1970 on rotting wood in a small wetland called Aunt Mary’s Bottom. R. palmata was refound in 1972 but neither species has been seen thereafter in spite of searching.
Far more species appear to have been stable than to have increased or decreased. This is partly due to the efforts of conservationists. Much of the coast has been acquired by the National Trust, and many heaths now have statutory protection. Epiphytes were little affected by acid rain, even when it was at its height in the 1960s. Thus, we see at the present a flora which in its general character is remarkably similar to that which existed when H.H. Wood began systematically to record mosses in 1876.
GORDON ROTHERO (DUNOON, ARGYLL): Exploration at the Gallop: 10 Years of Bryology in Assynt, Scotland
Assynt is a parish in Sutherland, in the far north-west of Scotland, botanically best known for the interesting vascular plant flora on the outcrops of Cambrian limestone at Inchnadamph. You need to think big for parishes in the Highlands: Assynt is slightly larger than the Isle of Wight and even so is not the largest parish in Sutherland. Other comparisons with the south also shed light on the nature of the area. The Isle of Wight has a population of some 130,000 and hundreds of roads criss-crossing the landscape; Assynt has about 1000 people and one main road through the middle and a minor road around the coast. The area is very rugged with spectacular hills like Suilven and Quinag and ground over 950 m on Conival, but much of the ground is an undulating plateau with a myriad of crags, lochans and wooded ravines. The coast is wild out on the Point of Stoer but there are shell-sand bays and quiet coastal woodland as well. The climate is extremely oceanic: wet and mild in the winter and wet and slightly warmer (if you are lucky) in the summer.
The Assynt area, and particularly the limestone, has attracted botanists over the years – the first bryophyte records date from 1767 – but recording has always come from fleeting visits, so the total number of records before my survey was surprisingly small. The survey which culminated in the bryophyte section in the Flora of Assynt (Evans et al., 2002) grew out of records made during the BBS visit in 1992 and the friendship I struck up with Pat and Ian Evans who live in the Assynt township of Nedd. Pat and Ian had already embarked on survey work for their vascular plant flora and, with fieldwork scheduled to end in 2000, a tight timetable was set in an attempt to cover at least a representative sample of the 164 tetrads in the parish. In the end, 116 tetrads had some sort of survey but the vast majority had just the one visit.
The area has a rich bryophyte flora with 156 liverwort and 345 moss taxa. In addition to this there are some 14 liverworts and 20 mosses reliably recorded for Assynt which were not refound during this survey.
The limestone, centred on Inchnadamph, has a number of interesting species, notably an abundance of Orthothecium rufescens and a number of other uncommon montane species at low altitude, including Pseudoleskeella rupestris, Encalypta alpina, Schistidium trichodon and Hypnum bambergeri. Dripping limestone also has stands of Seligeria trifaria and Hymenostylium insigne. There are surely more good discoveries yet to be made on the limestone.
Away from the limestone, much of the ground is rather unproductive wet heath with large populations of Campylopus atrovirens and Pleurozia purpurea, but there are also more undisturbed mires. There are a number of populations of Sphagnum austinii and S. fuscum, and species such as S. molle and S. strictum are probably more frequent than this survey suggests. My survey found only one stand of Campylopus shawii; this seems unlikely to reflect reality, though it is undoubtedly rare. The most important heath community is found under ericaceous shrubs at moderate to high altitudes on shaded slopes and amongst scree. Here there is an oceanic-montane liverwort community, usually associated with Sphagnum capillifolium and Racomitrium lanuginosum, composed of robust species with extraordinarily disjunct global distributions and including Anastrophyllum donnianum, A. joergensenii, Scapania ornithopodioides, S. nimbosa, Plagiochila carringtonii, Mastigophora woodsii, Bazzania pearsonii and Herbertus aduncus subsp. hutchinsiae.
The woodland and ravines also have an excellent oceanic flora with a good development of the Scapania gracilis–Hymenophyllum wilsonii community on rocks, large stands of Plagiochila spinulosa and P. punctata, and less frequent, but often large, stands of Lepidozia cupressina. The smaller Lejeuneaceae are also well represented with an abundance of Lejeunea patens, and with Drepanolejeunea hamatifolia, Colura calyptrifolia, Aphanolejeunea microscopica and Harpalejeunea molleri all reasonably frequent. The smaller scale of the woodland and the northerly latitude seem to result in the absence of some of the rarer oceanic species that occur on Skye although there is one population of Geocalyx graveolens.
Because much of the survey work took place in the winter months, the highest hills have not had as good a coverage as they justify. Some arctic-alpine species have already been mentioned as occurring on the limestone. There are also records for Arctoa fulvella, Kiaeria blyttii, Philonotis seriata and Ditrichum zonatum var. zonatum, and there are older records for K. falcata and Aulacomnium turgidum. After the flora was published, one trip to the hills north of Conival this year produced Barbilophozia lycopodioides and Paraleucobryum longifolium, indicating that there is much still to be found. Crags lower down can also have an interesting flora, with Glyphomitrium daviesii proving to be frequent on the gneiss; there are a few oddities such as Cynodontium jenneri and Dicranoweisia crispula here as well. Rock exposures by the many lochs have an interesting bryophyte community with several localities for Grimmia longirostris and a few sites for G. ovalis, as well as more common plants such as Pterigynandrum filiforme, Orthotrichum rupestre and Antitrichia curtipendula. Many lochs also have good populations of Odontoschisma elongatum along the flushed margins, and two lochs on the limestone have large stands of Cinclidium stygium in the same habitat.
The riparian flora, particularly where the run-off is from the limestone, is also interesting, and includes Hygrohypnum duriusculum and Rhynchostegiella teneriffae, usually with abundant Rhynchostegium riparioides, Thamnobryum alopecurum and Cinclidotus fontinaloides. On the gneiss, one fall has good populations of both Schistidium agassizii and Bryum dixonii, and on the sandstone there are at least two populations of Rhynchostegium alopecuroides.
The coastline is extensive and varied; where it is sheltered and wooded there are often similar oceanic species to the ravines with the addition of large stands of Frullania teneriffae and occasional F. microphylla. On the exposed Stoer peninsula, pride of place goes to the few stands of Myurium hochstetteri, a very beautiful hyper-oceanic species here in one of its few mainland localities. There are shell-sand beaches at Achmelvich, Clachtoll and Clashnessie, and these have some species typical of western dune systems, including Ditrichum gracile, Entodon concinnus, Didymodon ferrugineus, and occasionally Amblyodon dealbatus, Encalypta rhaptocarpa and Distichium inclinatum.
There are lots of holes still to be filled in Assynt and I suspect that more rare species will be found on the limestone. A number of species recorded by H.N. Dixon, W.E. Nicholson and E.S. Salmon on their visit in 1899 have not been refound, and the bigger hills need more attention. And this is just one of Sutherland’s parishes.
Evans PA, Evans IM, Rothero GP. 2002. Flora of Assynt. Nedd: PA & IM Evans.