PLANTLIFE SCOTLAND LOWER PLANTS AND FUNGI PROJECT

TRAINING DAYS: 13- 15 OCTOBER 2009:

OCEANIC BRYOPHYTES IN ATLANTIC OAKWOODS AND RAVINES

INCLUDING IDENTIFICATION, ECOLOGY AND MANAGEMENT ADVICE
NB These notes are available to download free of charge on the Plantlife Website: www.plantlife.org.uk

Plantlife Scotland management courses for agency staff

Plantlife is the UK's leading charity working to protect wild plants and their habitats. The charity
has 10,000 members and owns 23 nature reserves. Plantlife works hard to conserve threatened
plants and fungi in the places where they grow. Conservation of these species is delivered through
the charity's Back from the Brink species recovery programme, which is jointly funded by
Countryside Council for Wales, Natural England, Scottish Natural Heritage, charitable trusts,
companies and individuals. The series of training days running as part of the Lower Plants and
Fungi Project are tools to assist land managers and conservation staff to conserve Back from the
Brink and other species and their habitats. Plantlife's head office is in Salisbury and the charity
has national offices in Wales and Scotland. HRH The Prince of Wales is our Patron.

Who is the event for ?

This event is for professional conservation and land management staff such as foresters,
ecologists or conservationists, involved directly or indirectly in management of habitats
containing Atlantic oakwoods and ravines.
The course trains participants to identify a range of species and their habitats, as well as
identifying suitable management prescriptions. There are short introductory talks supported by a
field visit.

NB The courses also covered lichens but these are covered in separate course notes also available
on the Plantlife website.

What are Bryophytes?

Bryophytes are ancient plants dating back 450 million years and are the basal groups of all land
plants. It has been said, admittedly by a bryologist, that flowers are just mosses with a highly
adapted lifestyle. Mosses, liverworts and hornworts are not closely related, having diverged long
before flowering plants appeared, but they have obvious similarities of lifestyle and ecology so are
usually studied together. The term “lower plants” is certainly pejorative – these plants have
evolved to survive in deserts and under-water, on trees and asbestos roofs, buried by snow for 10
months or in crevices of city pavements. They are brilliant at what they do.

What we are concerned with here are the bryophytes of our ‘temperate rain forests’, a scarce
habitat on a global scale. Definitions of the temperate rain forest biome vary and the map below is
a little conservative and should probably include the upland forests in the Himalaya area and parts
of the Japanese archipelago.
Map 1. The world distribution of the temperate rain forest biome.

Diversity and distribution of oceanic bryophytes

The oceanic bryophyte element of the vegetation of British and Irish Atlantic oakwoods is not only
the richest in Europe but is also one of the richest bryophyte floras in the world. According to the
most recent European checklists there are 1769 bryophyte species of which Britain has
approximately 1150; this is about 65% of the European flora, a far higher proportion than for any
other plant group. For instance we have a meagre 15% of the European flowering plants. With
estimates of the global total of bryophyte taxa in the range of 20-25,000 species, this means that
Britain may have as much as 5% of the world’s bryophyte flora, an astonishing figure. Within this
large flora, the assemblages of bryophytes found in our Atlantic oakwoods and ravines are of
particular importance in international terms. These woodlands, particularly those with bouldery
ravines have as diverse a bryophyte flora as almost anywhere else on the planet, rivalling the
tropical-montane cloud forests (Porley and Hodgetts, 2005). Many of these woodlands and ravines
are SSSIs or SACs but a large number of important sites have no legislative protection. Similarly a
number of the species have a Species Action Plan and appear in the bryophyte Red Data List but
none have any legislative protection.

This flora is concentrated in the west of Britain and Ireland and the distribution is best illustrated
by a few maps of typical oceanic woodland species. The best of these sites in the west of Scotland
are all within the West of Scotland Important Plant Area. Species like Spotty Featherwort
(Plagiochila punctata) and Prickly Featherwort (Plagiochila spinulosa) can also occur on humid
sites outside of woodland, particularly in ravines but also in shaded block scree and montane
heath and Map 2 reflects that. The distribution of the good woodland sites is better illustrated by
the distribution of more strictly woodland species like Deceptive Featherwort (Adelanthus
decipiens) and Western Featherwort (Plagiochila heterophylla) shown in Map 3.

Map 2. The distribution of Plagiochila punctata in Britain and Ireland.

Map 3. The distribution of the oceanic liverworts Adelanthus decipiens and Plagiochila
heterophylla in Britain and Ireland.

Bryologists have long recognised the importance of the Atlantic flora and its limited extent in
Europe. In a celebrated paper, Ratcliffe (1968) listed the oceanic species of Britain and Ireland,
based on their phytogeography and his list of some 200 species, has only recently been
superseded by a more rigorous treatment which has produced a similar list (Hill & Preston, 1998).
Of these oceanic bryophytes about 50% are essentially woodland species or species with many
sites within woodland. The British and Irish distribution pattern of Atlantic woodland bryophyte
communities is well-illustrated by Map 3. In particular, Deceptive Featherwort (Adelanthus
decipiens) seems to be a good indicator of Atlantic oakwood with a continuity of canopy cover over
many, perhaps hundreds, of years and where internationally important bryophyte communities
are likely to be found. This oceanic community is now fragmented but does extend up into West
Sutherland but Deceptive Featherwort (Adelanthus decipiens) reaches its northerly world limit on
Raasay.
The global distribution of bryophytes is often very different to that of flowering plants with many
bryophyte species having an extraordinarily wide distribution. This also applies in a particular
way to the oceanic bryophytes, many of which have an affinity with the tropics and the global
distribution patterns of these species are remarkable to those used to vascular plants. To use
Deceptive Featherwort (Adelanthus decipiens) again, this liverwort is widespread in the Tropics and
also has a liking for isolated islands (Map 4).

Map 4. The global distribution of Adelanthus decipiens. (Map produced on DMap.)

Map 5. The world distribution of Radula aquilegia

60N

30N

30S

60S

150W 120W 90W 60W 30W 0 30E 60E 90E 120E 150E

Some species have a curiously disjunct distribution; Brown Scalewort (Radula aquilegia) for
instance is frequent in ravines in western Britain and Ireland and also occurs sparingly in western
Norway, the Faroes, NW Spain, Portugal and Macaronesia ( the Atlantic islands eg.the Azores,
Canaries etc) and then occurs again in the Himalaya (Map 5). This kind of disjunct distribution is
even more strongly marked in the liverworts of our oceanic montane heaths, some of which have a
few woodland and, more frequently, ravine sites.
In contrast, some of the species in our Atlantic oakwoods have a very restricted global distribution.
One of the most typical species of Atlantic oakwood is the European endemic liverwort Prickly
Featherwort (Plagiochila spinulosa) which can often form large cushions and patches on rocks and
tree trunks. It is widespread and locally abundant in western Britain and Ireland but is very rare
elsewhere, occurring sparsely in the Faroes, western Norway, and Belgium with a very similar
taxon in Macaronesia. This means that Britain and Ireland have the bulk of the world’s population
of this species and so, though it is locally common in the west, it is of high conservation priority.
It is the same general story with other oceanic liverworts that are common in the west of Scotland
like Western Earwort (Scapania gracilis) and Straggling Pouchwort (Saccogyna viticulosa) which
have very similar distribution patterns

At least as important as the individual species is the fact that only in the west of Britain and
Ireland do these oceanic bryophytes form distinct and widespread communities. An impoverished
version of the same flora can be found in western Norway, Brittany and the extreme western parts
of Spain and Portugal, although not always associated with oak trees. Many of the Atlantic
species also occur in Macaronesia (Canary Islands, Madeira and the Azores) and indeed some of
our rarities have their headquarters there, but are associated with very different vegetation, often
montane laurel forest.

The flora of our Atlantic oakwoods is reasonably well-known. The most comprehensive survey of
over 400 woodlands was carried out by Ben Averis (Averis, 1991), and this has been substantially
added to since then. More detailed surveys of individual woods have also given exhaustive species
lists so that we have a clear idea how rich some of the best woodlands are. To use some local
examples, Taynish NNR is an extensive Atlantic oak wood with a lot of rock outcrops, some of
which are calcareous, and some boggy areas and a bryophyte list that runs to 250 taxa. So this
one woodland has just under 25% of the British bryophyte flora. Another fine, large woodland
extends along the coast at Ellary in Knapdale and has over 200 species and perhaps more rarities
than Taynish. Perhaps even more remarkable are the narrow strips of woodland associated with
deep, bouldery ravines where total numbers of species can rival those of large areas of woodland.

Why so rich?
Besides the buffering provided against changes in humidity by the tree canopy of the woodland,
the western side of Britain and Ireland has other features which make it suitable for bryophytes.
The oceanic climate typical of Atlantic oakwoods with high rainfall and relatively small difference
in summer and winter mean temperatures is ideal for bryophytes, particularly liverworts. It is not
the absolute amount of rain that is critical but rather the absence of long periods of dry weather,
best expressed as the number of wet days (24 hours with more than 1mm of rain). The greatest
variety of the oceanic bryophytes occurs in those areas which have over 200 wet days per year.

There are other bryophyte-friendly characteristics on the west side of Britain and Ireland. Of
prime importance is the recently glaciated landscape characterised by lots of crags and scree,
steep-sided burns and raised-beach cliff lines giving suitable substrates and a buffer against
changes in humidity. The rocky terrain and the ravines also hamper intensive management,
particularly heavy grazing, allowing semi-natural woodland remnants to survive. The geology is
also favourable, with old, hard and often gritty rocks prevailing but with some variety in rock
chemistry. Finally, the whole Atlantic area is well away from the main centres of population and
has been relatively free of pollution from industry in the recent past.

Different micro-habitats
A rocky woodland or wooded ravine presents an attractive image of a landscape swathed in a
carpet of green ‘moss’ but in reality there are a large number of different species exploiting subtly
different habitats within the woodland. Virtually all of the interesting species need a firm
substrate so there are few oceanic species growing on the woodland floor. Most are too small to
compete with robust common species like the poorly named Little Shaggy-moss (Rhytidiadelphus
loreus) or Greater Fork-moss (Dicranum majus) but the handsome oceanic Bottle-brush moss
(Breutelia chrysocoma) can form substantial wefts in more heathy woodland The trees and rocks
provide the most obvious niche, many being completely covered in bryophytes, the most
ubiquitous being Slender Mousetail-moss (Isothecium myosuroides).
In the best woodlands, this Slender Mousetail-moss (Isothecium myosuroides)community on
rocks and tree bases also includes Prickly Featherwort (Plagiochila spinulosa) and Western
Scalewort (Scapania gracilis), often with the oceanic Wilson’s Filmy Fern, and with Spotty
Featherowrt (Plagiochila punctata) particularly common on trees. The presence of this
assemblage on tree trunks and rocks is probably the best indicator of old Atlantic oakwood. In
Sunart, on the Ariundle NNR and on a few other sites in that area, Prickly Featherwort (Plagiochila
spinulosa) is frequently replaced by Western Featherwort (Plagiochila heterophylla) which occurs
on nearly every rock and tree. This handsome species was once thought to be a British endemic,
Plagiochila atlantica, and thus attracted some attention from conservation bodies but it was then
found at a few other Atlantic woodland sites in the British Isles and Brittany and is now thought to
be conspecific with a widespread tropical species. The best woodlands often have both Western
Featherwort (Plagiochila heterophylla, as we must now call it), and Deceptive Featherwort
Adelanthus decipiens on the rocks and this indicates a woodland of high conservation value.

A less obviously mossy habitat, but one which contains a number of interesting species, is the
lower rocks, often with sloping faces down which ground water flows during rain. One moss of
interest here is Prostrate Signal-moss (Hageniella micans), uncommon but widespread in NW
Britain and SW Ireland. This habitat is constantly encroached upon by more robust species of the
woodland floor and depends on erosion or the passage of grazing animals (and sometimes
humans) to keep it open. Despite not producing any sporophytes, Prostrate Signal-moss
(Hageniella micans) seems able to persist and shift about as new sites open up. We know so little
about the autecology of virtually all of these oceanic species that such apparent anomalies remain
unexplained.

On acid-barked trees where Featherwort (Plagiochila) species are frequent there may also be
patches of the more cryptic wedge flapwort (Leptoscyphus cuneifolius) a nationally scarce species
locally frequent in the west of Scotland and Ireland but elsewhere in Europe only known from one
site in Norway. Trees and shrubs with more neutral bark may have a number of tiny oceanic
liverworts as epiphytes. The most common is probably pearl pouncewort, Lejeunea patens, but
two other, even smaller species, pointed pouncewort, Harpalejeunea molleri, and toothed
pouncewort, Drepanolejeunea hamatifolia, can be frequent, particularly on hazel in very humid
sites.

The steep faces of rocks in and by burns in wooded ravines provide another extremely important
habitat, particularly for the Pounceworts (Lejeunaceae), a mainly tropical family of tiny liverworts.
The large common species here tend to be Yellow Fringe-moss (Racomitrium aciculare) and Rusty
Feather-moss (Sciuro-hypnum plumulosum) on most of the rocks, often with the oceanic moss
Flagellate Feather-moss (Hyocomium armoricum) at the margins. Where the rocks are more base-
rich there can also be large patches of Long-beaked Water Feather-moss (Platyhypnidium
riparioides) and Fox-tail Feather-moss (Thamnobryum alopecurum).

Some large rocks in ravines, regularly inundated but not scoured, can be covered in a weft of Pearl
Pouncewort (Lejeunea patens) and, further from the water, the even smaller stems of Fingered
Cowlwort (Colura calyptrifolia), Toothed Pouncewort (Drepanolejeunea hamatifolia) and Pointed
Pouncewort (Harpalejeunea molleri) make a delicate green tracery against the dark stone. All of
these tiny liverworts have an extremely restricted oceanic distribution in Europe but are
widespread in the tropics. This community often also has the dark green patches of Hutchins’
Hollywort (Jubula hutchinsiae), with toothed leaves resembling a tiny holly leaf, and the bronzed
stems of Brown Scalewort (Radula aquilegia). There are some rarities in this habitat as well,
including Wilson’s Pouchwort (Acrobolbus wilsonii) which has some 30 sites in Scotland (including
a number in the Sunart area), a few more in SW Ireland and elsewhere in the world only occurs in
the Faroes and the Azores. The tiny liverwort Atlantic Pouncewort (Lejeunea mandonii) has always
been very rare in Scotland and now seems to be restricted to one site in each of three ravines. All
of these species are particularly sensitive to water-quality, especially to silt loading, and increases
in acidity and changes to these qualities can severely damage this community.

Oceanic species of Atlantic oak woodland and associated ravines in Scotland

Typical species Zygodon conoideus
Aphanolejeunea microscopica
Breutelia chrysocoma Scarce species
Colura calyptrifolia Adelanthus decipiens
Dicranum scottianum Fissidens polyphyllus
Drepanolejeunea hamatifolia Hageniella micans
Harpalejeunea molleri Leptoscyphus cuneifolius
Hyocomium armoricum Plagiochila heterophylla
Jubula hutchinsiae Radula voluta
Lejeunea lamacerina Ulota calvescens
Lejeunea patens
Lepidozia cupressina Rare species
Metzgeria leptoneura Daltonia splachnoides
Plagiochila bifaria Acrobolbus wilsonii
Plagiochila exigua Dumortiera hirsuta
Plagiochila punctata Lejeunea holtii
Plagiochila spinulosa Lejeunea mandonii
Radula aquilegia Radula carringtonii
Saccogyna viticulosa
Scapania gracilis
Ulota phyllantha

THREATS TO BRYOPHYTES IN ATLANTIC OAKWOODS AND RAVINES

Invasive non-native species

The most depressing management problem for some of the best of our woodlands for oceanic
bryophytes is the seemingly inexorable spread of Rhododendron. Initially the growth of scattered
bushes of Rhododendron increases humidity and may actually improve conditions for some
oceanic species but eventually, as the canopy closes, the shading and litter fall exclude all but a
few tolerant common species. At Ellary, the fine woodland, part of the Knapdale Woods SAC, is
being engulfed by Rhododendron, with the problem having become significantly worse over the
past ten years despite some clearance in a limited area. There is now some hope that a
management plan to deal with the Rhododendron can be agreed with the owner In Sunart, a more
comprehensive effort to clear Rhododendron is being made and the situation looks more hopeful
though there is still a long way to go. Away from listed sites, the problem is becoming
exponentially worse and unless more effort is made and very large sums of money are spent, we
will lose this battle.

Figure 1. Rhododendron jungle under mature oaks at Ellary, Kintyre.

Figure 2. Shade and deep and persistent litter cast by beech trees.

Beech trees are often magnificent and were a popular addition to many policy woodlands and
from there they have spread into oceanic broadleaf woodland where the dense shade cast by the
canopy and the copious and persistent leaf litter present problems for bryophytes and lichens.
This is particularly true of ravines where beech seedlings find it easy to establish and soon make
an already shaded habitat very dark indeed. A similar problem can also occur when non-native
conifers seed into and become established in ravines.
Overgrazing
Where regeneration in a woodland is prevented by grazing by domestic livestock, deer or
occasionally feral goats, the woodland will become moribund and eventually disappear. This is
not so apparent with long-lived oak trees but many areas of birch woodland, a tree with a shorter
lifespan, can become fragmented alarmingly quickly. The loss of the trees will eventually lead to a
loss of at least some of the oceanic species, usually the least common and most sensitive.

Under-grazing
In order to give tree regeneration a chance, some areas of woodland have been fenced to exclude
all grazing animals. While this may well increase regeneration rates, it can be very bad news for
bryophyte communities. In the south of Scotland, in the absence of grazing animals, the growth
of bracken and bramble under an open canopy can be prodigious and these plants and their litter
smother the rocks, particularly the low rocks on the woodland floor. On some sites bramble has
smothered boulders up to 2m in height, wiping out interesting bryophyte communities. In the
north-west of Scotland, within the exclosures, there can be rapid growth of coarse plants like
Tufted Hair-grass (Deschampsia cespitosa), Woodrush (Luzula silvatica) or heather (Calluna
vulgaris) again smothering the lower rocks to the detriment of bryophyte communities.

Figure 3. Growth of coarse field and shrub layer after exclusion of grazing in Knapdale (left)
and West Sutherland (right).

A case can be made that, in time, the regeneration will lead to a closed canopy and this will
repress the bracken, bramble and coarse grasses and that the rocks will become available again for
colonisation by bryophytes. However, many of the rarer species now have such fragmented
populations and such limited means of propagation that, once lost from a site, the plant may
never manage to become re-established. The lack of information on how species spread (or in
many cases, do not spread), also hampers any attempt at modelling the performance of the
community under differing management regimes.

There is no easy answer to this problem of regeneration within existing woodland and not radically
changing the conditions on the woodland floor, but woodland managers need to ask themselves,
when framing a woodland management policy, what kind of woodland they are trying to create
and over what timescale.

Fragmentation of habitat
Many of the oceanic species rarely produce sporophytes and few have specialised means of
vegetative propagation. This means that most spread is by fragments of the plant being blown or
washed, or occasionally carried, to a new patch of suitable habitat. This means that spread is
sporadic and slow and the likelihood of it occurring is inversely proportional to the size of the
extant population. So fragmentation of suitable habitat is a real barrier to the spread of these
species. Each area of oceanic woodland acts as an island in a sea of unsuitable habitat which
makes it more likely that small populations will die out and severely curtails the likelihood of re-
colonisation from other sites.

Hydro-electric schemes
A particular threat to oceanic ravines comes from the development of large numbers of small-
scale run-of-the-river hydro-electricity schemes. These schemes take water out of the river, pipe it
down to the powerhouse and then return it to the river. This leaves a stretch of the watercourse
between the intake and the powerhouse with the hydrology radically altered. There are no long-
term studies of the effects on bryophytes and lichens of a lower level of discharge down ravines
but knowledge of the ecology of the plants involved and observations on the extant communities
in ravines altered by hydro-schemes tens of years ago gives some pointers. The most likely effects
are

• a downward shift to the new median water level of riparian species

• a growth of more robust, woodland floor species on the rocks above this
• a loss of habitat for the community of small liverworts on rock faces as a result of
increased competition
• a decrease in frequency of spate flows and resultant erosion which frees-up habitat for the
smaller liverworts

The interplay between the various factors is complex and it is not always clear what the limiting
factors for any one plant are, but the precautionary principle suggests that hydro-schemes should
not be sited on ravines with nationally and internationally important bryophyte and lichen
communities

Forestry
In the past insensitive planting of commercial conifer crops too close to the edge of ravines has
meant that a large number of ravine sites, particularly smaller ones, have lost a significant
proportion of their bryophyte and lichen interest. Guidelines are better now but we are left with a
legacy of damaged sites which may never recover; observing these ravines after clear-felling is a
depressing prospect. The run-off from commercial conifer plantations may also have an effect on
water quality in the ravines both in terms of silt loading, thus increasing scouring, and acidity
which seems to encourage algal growth at the expense of bryophytes.

Pollution
All of the bryophytes that grow in and near ravine burns are susceptible to changes in water
chemistry, particularly to eutrophication by run-off from fields or from ineffective sewage
systems. Some are directly affected but most are just out-competed by the growth of robust
common species, both higher plants and bryophytes, and algae in response to the nutrient
enrichment.

MANAGEMENT RECOMMENDATIONS FOR BRYOPHYTES IN ATLANTIC OAKWOODS AND

RAVINES

Invasive non-native species

Eradication of Rhododendron should be a priority action in any management plan for Atlantic oak
woodland. Removal of Rhododendron is the essential action but if there is a possibility of using
methods which impact less on the bryophyte flora, then these should be used. Where a good
bryophyte flora persists and access to Rhododendron bushes is not too difficult the method of
choice would be stem injection. This leaves the bush standing and causes little disturbance,
mediates a little the loss of humidity and the dead bush, though a little unsightly, may actually
provide a habitat for some species. This method has been used to great effect in North Wales.
Unfortunately ravines are often difficult of access and have Rhododendron growing high on crags,
so removing Rhododendron will be both expensive and involve rope-access techniques. Failure to
remove Rhododendron from ravines because of access difficulties means that not only is the
ravine itself under threat but the site provides a seed source that can infect the wider area.

Beech and non-native conifers are a less obvious problem and there is often some understandable
resistance to removing mature trees which are often very fine. However, the threat that they pose
should not be underestimated and where possible, periodic removal of seedlings, particularly from
ravine sites would go some way towards reducing this threat. Unfortunately, some models of
climate change suggest that conditions in Scotland are likely to favour the beech and it is
probable that this problem will get worse. Non-native conifers should be removed.

Forestry and ravines

Fortunately, the current guidelines with regard to commercial plantations and watercourses are
far better now and no longer should good ravine sites for bryophytes and lichens be shaded out.
However, even with current guidelines, mature conifers will still have some shading effect and
there is a strong argument for a much wider buffer zone for those ravines which have an existing,
full or partial, broadleaf canopy and which are thus likely to have good bryophyte and lichen
diversity. Conifers should be planted far enough back from the break in slope that forms the
ravine so that no shading of the ravine occurs when the trees are mature. The effect on ravine
sites of direct run-off from commercial conifer plantations can be mitigated by not having ditches
running directly into water-courses as is current good practice. This does need occasional
maintenance and does not deal with older plantations.

One dilemma that is apparent in a number of forest areas in the west of Scotland is the presence of
oceanic species, including some that are nationally scarce, on broadleaf trees by burns that flow
through a narrow corridor of non-native conifers. The mature conifers have effectively created a
ravine in terms of the buffering to changes in humidity and it is reasonable to assume that the
oceanic species would not be there if the conifers were absent. Clear-felling will almost certainly
mean that these populations will be lost but leaving a protecting barrier of conifers may not be
realistic because of wind-throw. The hope must be that after restocking with conifers, leaving a
much wider strip for colonisation by broadleaves, these more mobile species will return.

Woodland regeneration and exclosures

There is no easy answer to the problems posed for bryophytes and lichens by exclosures, although
where possible, a reduction in the number of grazing animals in the general area and so removing
the need for an exclosure should be considered as an option. Some grazing is helpful in sustaining
good habitat for both lichens and bryophytes but there is no doubt that in the long term,
expansion of broadleaf woodland will also be of considerable benefit. Exclosure fences should be
seen as a temporary measure, to be removed once tree regeneration has reaches a stage where it
can sustain some grazing. Where dense regeneration threatens old trees with good lichens and
bryophytes, some sort of ‘halo-thinning’ might be required. In the long term, if all that can be
achieved within existing resources is dense regeneration of birch then that is better than no
regeneration as long as it does not damage bryophyte and lichen species of conservation concern.

Within existing woodland, particularly heavily managed oak woodland with even aged stands and
no understory, it may be sensible to undertake some thinning to create glades for regeneration to
diversify the stand structure. While this may be detrimental to the local bryophyte community
initially, by reducing canopy cover and reducing humidity, particularly on south-facing slopes, in
the long term it will be beneficial. This benefit will be much greater if under-story species like
hazel are encouraged along with trees with more base-rich bark like ash or elm, rather than the
more usual dense regeneration of birch. It would seem sensible that any operation of this nature
was preceded by a lower plant survey to make sure that rare species were not damaged.

Dead wood
Dead wood is an excellent habitat for bryophytes, lichens and fungi and has its own typical
communities which may include nationally important species. The best development of these
communities is almost always on the larger logs. Where possible all dead wood should be left on
site and there should be a policy of increasing the amount of dead wood aiming for the Forestry
commission suggested target of 5m3 per hectare of dead wood over 20cm in diameter.

Pollution
Most of the oceanic ravines are in parts of the country where levels of pollution, both current and
historic, are low. Pollution from run-off from agriculture or sewage are likely to be of very local
occurrence but may have a significant effect and all such run-off should be prevented from
entering ravine sites.

Hydro-electric schemes
Any bryophyte or lichen interest on potential sites for hydro-schemes is best identified as early as
possible in the planning process and the initial environmental assessment should involve
surveyors sufficiently competent to assess or at least indicate that this interest is present.
Inevitably most if not all oceanic ravines will fall into this category. Much can be done with
careful planning and micro-siting to avoid damaging bryophyte or lichen interest during the
construction process, by avoiding certain trees and boulders and by carefully re-siting rocks.

The change in hydrology will change both the riparian habitat and the humidity regime in the
ravine and this will have an effect on the plants. There is no obvious mitigation for this.
Unfortunately what little monitoring of these effects that has taken place has been ‘after the
event’ and without baseline data for comparison, and this should be rectified. If the scheme
precludes all but the highest spate flows then it is likely to have a greater effect than one that
allows through more than just peak flows. The uncertainty is with regard to the extent of the
effect of the reduced flow on humidity-demanding oceanic species and this suggests that the best
ravines for oceanic bryophytes should not be used for hydro-schemes.

There should be a bryophyte survey of all sites with ravine or incised sections that fall within the
West Coast IPA and on the larger islands, particularly those with some trees. This essentially
covers the whole of the watershed to the west coast of Scotland north of the Clyde. The
precautionary principle suggests that those sites with nationally important oceanic bryophyte
communities should not be used for hydro-schemes because of the uncertainty over the long term
effects of the changes in humidity.

References and further reading

Averis ABG, 2000. Bryophytes and run-of-river hydro-electric power schemes in the British Isles.
Unpublished paper for Scottish Natural Heritage.

Averis, A B G, 1991. A survey of the bryophytes of 448 woods in the Scottish Highlands. Sottish
Field Unit Report, NCC, Edinburgh.

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British Bryological Society.

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

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Ratcliffe, DA, 1968. An ecological account of Atlantic bryophytes in the British Isles. New
Phytologist 67, pp365-439.

Rothero, G.P. 2005 Oceanic Bryphytes in Atlantic Oakwoods. Bot. Journal Scot. 57, pp.135-140.

Rothero GP, 2005. Mosses and Liverworts. Part of the ‘Naturally Scottish’ series published by
Scottish Natural Heritage, Battleby.
Worrel R, 2009. Management of woodland plants in Atlantic broadleaved woodland: a conservation
framework. Plantlife 2009.

Photographs on the following page:

Page 1: mosses and liverworts of Atlantic oak woodland

Page 2: mosses and liverworts of Atlantic ravines.

Plantlife Scotland’s Lower Plant and Fungi project is a PLINKS project supported by the
British Bryological Society, British Lichen Society, Forestry
Commission Scotland, Scottish Natural Heritage (SNH) and the Scottish Environmental
Protection Agency (SEPA)