Contents - Plantlife
Transcription
Contents - Plantlife
Cicerbita alpina (L.) Alpine Blue-sow-thistle, Bliochdan Gorm Ailpeach ASTERACEAE SYN.: Alpine sow-thistle, Blue sow-thistle.C. alpina (L.) Wallr., Sched. Crit. 434 (1822). Sonchus alpinus (L.), Sp Pl. 794 (1753). Mulgedium alpinum (L.) Less., Syn. Gen. Comp. 142 (1832). Sonchus caeruleus Sm., Fl. Brit., 2: 815 (1800) nom. Illegit. Superfl. Pro Soncho alpino (L.). Lactuca alpina (L.) A. Gray, Syn. Fl. N. Amer., 1(2): 444 (1884). Status Restricted to 4 sites in Scotland UK BAP Long List key species Schedule 8 Wildlife & Countryside Act (1981) Plantlife Back from the Brink species 4 10km squares Status in Europe: not threatened Contents 1 2 3 4 5 6 Morphology, Identification, Taxonomy & Genetics.................................................... 2 1.1 Morphology & Identification ............................................................................ 2 1.2 Taxonomic Considerations .............................................................................. 3 1.3 Genetic Implications ...................................................................................... 4 Distribution & Current Status ............................................................................... 5 2.1 World .......................................................................................................... 5 2.2 Europe ........................................................................................................ 5 2.3 United Kingdom ............................................................................................ 6 2.3.1 Scotland................................................................................................. 6 Ecology & Life Cycle............................................................................................ 9 Habitat Requirements ....................................................................................... 10 4.1 The Landscape Perspective ........................................................................... 10 4.2 Communities & Vegetation ........................................................................... 10 Management Implications .................................................................................. 11 Threats / Factors Leading to Loss or Decline or Limiting Recovery............................ 11 Work on Cicerbita alpina is supported by: 1 7 Current Conservation Measures .......................................................................... 7.1 In Situ Measures........................................................................................ 7.2 Ex Situ Measures ....................................................................................... 7.3 Monitoring Cicerbita alpina & the Common Monitoring Standard......................... 7.4 Recommendations for Future Work ................................................................ 8 References ...................................................................................................... 9 Acknowledgements ........................................................................................... 10 Contacts ......................................................................................................... 11 Links .............................................................................................................. 1 1.1 12 12 13 13 14 15 17 17 17 Morphology, Identification, Taxonomy & Genetics MORPHOLOGY & IDENTIFICATION Cicerbita alpina is a tall, robust, rhizomatous perennial with attractive blue-flowers. In late summer its pale blue-violet, dandelion-like flowerheads open in tight clusters at the top of the stem. Cicerbita alpina has an erect stem 50-250cm, simple or branched, usually with dense, reddish glandular hairs on the upper part. Alternate leaves are 8–25 x 2-12 cm, glabrous, glaucous beneath; lowest lyrate or runcinate-pinnatifid with a large, broadly triangular, acuminate terminal lobe and a few pairs of much smaller triangular lateral ones, with the base narrowed into a winged petiole; upper smaller and less divided, with a winged periole widened into a cordate-amplexicaul base. Flowerhead is an elongated panicle; peduncles with dense reddish glandular hairs. Involucre is 10-15 x 7-10 mm; bracts linear, usually with numerous reddish glandular hairs. Petals are pale blue. Achenes 4.5-5 mm, linear (Sell, 1986). Specimens are held at a number of institutions including the Smith Herbarium (LINN), Royal Botanic Garden Edinburgh (RBGE) (E) and the Natural History Museum (NHM). George Don sent specimens from Lochnagar to Sir James Edward Smith who published a description of the plant as Sonchus caeruleus SM. (Smith, 1811). Some of Don’s specimens are housed in the Smith herbarium (LINN) while others were given to private collectors one of whose sheets is now in the herbarium of the Royal Botanic Garden Edinburgh (E). Chromosome number is 2n=18 (Clapham et al, 1987). No hybrids have been reported. 2 Figure 1 - Cicerbita alpina (L.) Alpine Sow-thistle. (Photograph by Bob Gibbons, © Bob Gibbons/Natural Image). 1.2 TAXONOMIC CONSIDERATIONS The genus Cicerbita Wallr. belongs to the family Asteraceae (Compositae) subfamily Lactucoideae tribe Lactuceae. It consists of about 18 species native to the mountains of Europe, Asia, North Africa and North America. In the character of the pappus, Cicerbita is closely allied to Wall Lettuce Mycelis Cass., Cephalorrhynchus Boiss and Steptorhamphus Bunge. Mycelis is distinguished by its fewer involucral bracts, which are in two series, the outer very inconspicuous, and by its yellow flowers. The long beak to the achene distinguishes Cephalorrhynchus and Steptorhamphus. In Cicerbita the beak is absent or not more than 0.5mm. The large genus Lettuce Lactuca L., in which all these small genera including Cicerbita were once included, is distinguished by its pappus hairs all being of one kind and more or less equal in length. C. alpina is the only native species to the British Isles but three others, Common Blue-sowthistle C. macrophylla, (Willd) Wallr. subsp uralensis (Rouy) P.D.Sell, Hairless Blue-sowthistle C. plumieri (L.) Kirschleger and Pontic Blue-sow-thistle C.bourgaei (Boiss.) Beauverd, have been recorded as naturalised (Sell, 1986). These three naturalised species are distinguished by the following characteristics. C. plumieri is glabrous with a very scattered distribution in England and Scotland as a result of garden escapes. C. bourgaei is not rhizomatous with sparse, simple non-glandular hairs. It has a very scattered distribution in England, Scotland and the Isle of Man as a result of garden escapes and appears to be on the increase. C. macrophylla has lower leaves hairy on veins beneath with a cordate terminal lobe and usually only a single pair of lateral lobes, panicles are wider, more or less corymbose and achenes narrowly elliptical. This is another garden escape with a generally lowland distribution but reaching 320m at Tomintoul. 3 1.3 GENETIC IMPLICATIONS It is possible that there is a genetic issue that needs to be considered with regard to the conservation of C. alpina in Scotland. Despite repeated efforts, collection of viable seed over the last 20 years has not been successful. Seed collected has consistently been described as ‘shrivelled’ and has not germinated. Historically there is some evidence of viable seed being produced in the latter part of the 19th century and the early part of the 20th century at Coire Fee and Caenlochan, with samples of seedlings preserved at the Natural History Museum, London. Seed production is described as sporadic in other parts of C. alpina’s continental and Scandinavian range (J. Birks, pers. comm.), and viable seed production may depend on seasonal variation. However, C. alpina of continental origin growing in a garden south of Edinburgh is reported to seed prolifically every year (I. Bainbridge, pers. comm.). Cicerbita plants of continental origin grow easily at the Royal Botanic Garden Edinburgh, from seed that is sown in January or February in pots, and grown on in a cold glasshouse or frame (R. McBeath, pers. comm.). As the species has a continental distribution, this would suggest optimum conditions of hot summers and cold winters. Seedlings at Lochnagar were observed in 1977 after the long, hot summer of 1976, which may have been significant. Self-incompatibility (SI) is common in the Compositae (Richards, 1997). Work has been undertaken by RBGE to assess whether populations are uniclonal, and to what extent they are differentiated from one another, in order to assess if inbreeding depression could result in reproductive failure (Oliver et al, 2000). The study used Randomly Amplified Polymorphic DNA (RAPDs), analysed from plants sampled at the four locations in Scotland at which Cicerbita alpina occurs. Although the number of individuals sampled from each extant population limited interpretation of the results (see Table 1), the work showed that variation exists between all populations and that all populations are genetically unique from each other. Table 1 - Number of genotypes identified at the four locations for Cicerbita alpina in Scotland (Oliver et al, 2000). SITE Coire fee Lochnagar Coire Kander Caenlochan NO. INDIVIDUALS SAMPLED 5 4 OF NO. OF GENOTYPES IDENTIFIED 1 2 9 3 1 n/a However, even though different clones between populations were identified, these clones may be of the same self-incompatible type and may not be compatible mating types. Furthermore, the low level of within-population genetic variation, compared to betweenpopulation variation, indicates that mating events between close relatives are more likely. Further work is required for clarification; although it is possible that population isolation is a factor in the lack of seed production in Scotland. 4 2 2.1 Distribution & Current Status WORLD Cicerbita alpina grows in all the main mountain ranges of Europe, including the Pyrenees, Alps, the Apennines and Balkans and throughout the mountains of Scandinavia, as well as the Scottish highlands. 2.2 EUROPE Cicerbita alpina was first described from Lapland in 1720 (Nordsted, 1920). It is a widespread and locally abundant species of submontane birch and pine forest in Scandinavia and in European mountain ranges southwards to the Pyrenees, northern Apennines and the mountains of the Balkan peninsula (see Figure 2). Figure 2 - European distribution of Cicerbita alpina. © Koeltz Scientific Books. Nordhagen details associated species in Norway in fern-rich sub-alpine birch woods, lowalpine tall-herb vegetation, in sub-alpine Willow Salix scrub and in herb-rich sub-alpine birch woods (Nordhagen, 1943). In western Norway, Cicerbita alpina seems to avoid the extreme 5 oceanic areas and tends to be found in the more continental inner fjord areas. It avoids the west coast; like many boreal species, it tolerates or even needs high summer temperatures, occurs in areas of cold winters and tends to be absent from areas with mild winters. Its European distribution coincides with temperatures of –4oC in the coldest month for the highest point in the 50 x 50 km grids of Atlas Flora Europaeae and annual maximum temperatures for the highest point of 28oC (Dahl, 1998). Eastern Scotland is therefore right at the western edge of a continental boreal climate with cold winters and warm summers (Birks, pers. comm., 1996). Its distribution in Europe differs from Scotland in being a mainly submontane species more typical of damp, open, lightly grazed woodland and scrub in contrast to the steep mountain crags of the Scottish sites. It is a characteristic species of Scandinavian tall herb meadows and one of its synonyms, Lactuca alpina (L) A. Gray, lends its name to the alliance Lactucion alpina (McVean & Ratcliffe, 1962) where moist conditions prevail. In Scandinavia, C. alpina is commonly a ditchside plant. It is not a priority for conservation throughout the rest of its range due to its common and widespread status. 2.3 UNITED KINGDOM Cicerbita alpina is restricted within the UK to Scotland. Figure 3 – Scottish distribution of Cicerbita alpina. 2.3.1 SCOTLAND Cicerbita alpina was first discovered in Scotland in 1801 by George Don on one of the precipitous ledges in the northeastern corrie of Lochnagar. The species is extremely local and confined to 4 x 10 km squares at 4 locations in the Eastern Grampian mountains in 6 Angus (v.c. 90) and South Aberdeen. (v.c. 92) (see Figure 3). The species grows on north or northeast facing or well-shaded ledges or gullies moistened by late snowmelt, between approximately 600m at Coire Fee and 1000m at Lochnagar. Few other montane vascular plants are confined to the eastern highlands of Scotland with the exception of Purple Colt’sfoot Homogyne alpina, which is a possible introduction known from one location in the Clova Mountains, Angus. There is evidence (Gardiner, 1848 and Ewing, 1887 in Marren et al, 1986) that C. alpina occurred at more locations in the past within Caenlochan Glen, within Glen Doll and at the lost site at Canness Glen. C. alpina in Scotland will have occurred as part of the tall herb communities of the sub-alpine tree line communities. It is therefore likely that its current limited distribution is a result of the increase in grazing pressure from the late 19th century onwards, which has forced its retreat to inaccessible ledges. SITE DESCRIPTIONS Site details for the locations of Cicerbita alpina in Scotland are given in Table 2. C. alpina sites are difficult to access and ropes are required for safety at Lochnagar and Coire Kander. Once on site it can be difficult to locate the colonies and reference should be made to a number of site reports (Alexander, 2000; Clarke & Clarke, 1995; Marren et al, 1986). Table 2 - Site details for locations of Cicerbita alpina in vice-counties in Scotland. LOCALITY VICE COUNTY (VC) VC NO. NO. OF COLONIES 6 Coire Fee: Glen Doll Angus 90 Coire Kander South Aberdeen 92 Caenlochan Glen Angus 90 2 “several” FLOWERHEAD COUNT DATE RECORDER W. Gardiner A.G. Payne L. Alexander 200 67 1848 1979 1999 1866 1958 1977 c.155 c.165 c.58 1978 1979 1995 A.G. Payne A.G. Payne P & J Clarke 1999 1848 1975 1976 1977 1978 1979 1980 1981 1983 1984 L Alexander W. Gardiner F.H. Perring C. Geddes A.G. Payne C. Geddes NCC NCC C. Geddes “ anon. 44 239 a) 176 b) 66 12 3 30 20 8 41 14 45 c. 35 plants 7 Ratcliffe R.E.Randall SOURCE Marren et al 1986 “ Alexander 2000 Ratcliffe 1958 “ Marren et al 1986 “ “ Neiland & Wilcock 1996 Alexander 2000 Gardiner 1848 Marren et al 1986 “ “ “ “ “ “ “ SNH files LOCALITY Lochnagar VICE COUNTY (VC) South Aberdeen VC NO. NO. OF COLONIES 92 2 sites 3 sites 2 Canness glen Angus 90 1 FLOWERHEAD COUNT “Abundant” “Part of dense vegetation” A: few; B: many B: 30+ B: 42 A: 6; B: many B: many; C: 2+ A: 3; B: 120+; C: 22+ A: 6; B: 160; C: 51 B: 222; C: 75+ A: 8; B: 311; C: 73+ B: 53; C: 50+ A: 7; B: 46; C: 15+ A: 3; B: 163; C: 85+ A: 50+; B: leafy shoots only; C: 110 a) 110 b) 239 “large number” 2-4 annually DATE RECORDER 1801 1927 1932 G. Don 1972 1973 1974 1975 1976 1977 A.G. Payne; C. Geddes SOURCE LINN. E. Cairngorm Club Journal Xi; p.314 & XIII p.91 Marren et al 1986 “ “ “ “ “ 1978 “ 1979 1980 “ “ 1981 1982 L Farrell; F Leney 1983 “ SNH files Marren et al 1986 1985 AG Payne; P Rothwell SNH files 1999 1885 1937 1947-1976 L. Alexander Alexander 2000 BM, Oxford Holden 1952 Marren et al 1986 R.M. Adams COIRE FEE In the 19th century, Glen Doll was the best known and the most accessible of the C. alpina sites with a least 6 colonies recorded. It was thought to have been lost from the glen until it was re-discovered in 1979 by A. G. Payne in a narrow north-facing gully now referred to as the Coire Fee site. The gully is cold, and the wet substrate comprises loose rock. Associated species include Purple Saxifrage Saxifraga oppositifolia, Yellow Saxifrage Saxifraga aizoides and Alpine Lady-fern Athyrium distentifolium. Plants grow profusely within the main gully and on a ledge above the main gully to the right while climbing. A total of 239 flowering stems were counted in 1999 (Alexander, 2000). A few young plants have also been recorded below the chockstone. COIRE KANDER Cicerbita alpina grows on 2 ledges at Coire Kander. The site is steep, wet and very slippery in places and ropes are required for safety. The plant grows vigorously on the lower northfacing ledge at about 840m and is co-dominant with Alpine Lady-fern Athyrium distentifolium. In addition, a large colony of Athyrium distentifolium also grows just beneath the ledge. Other associated species include Alpine Lady’s-mantle Alchemilla alpina, Wood Crane’s-bill Geranium sylvaticum and Mountain Sorrel Oxyria digyna. A total of 176 flowering stems were counted in 1999 (Alexander, 2000). The upper ledge is above and to the west and is smaller with a total of 66 flowering spikes counted in 1999 (Alexander, 2000). CAENLOCHAN GLEN Several colonies were known in the 19th century and according to Gardiner (1848) it grew “in several places by the sides of streams at the head of Caenlochan Glen”. Only one colony is known today and this is probably the best-known site for the species in Britain (Marren et al, 1986). This population occupies a north west-facing ledge at about 840m with steep overhanging rock walls on the southern and western sides. The site is very open and 8 exposed and limited by the extent of the ledge. The soil is stony but deep and rich, flushed by water from ledges above. Associated species include Alpine Lady-fern Athyrium distentifolium, Great Wood-rush Luzula sylvatica, Sheep’s-fescue Festuca ovina and Hogweed Heracleum sphondylium. A total of 31 flowering stems were counted in 1999 (Alexander, 2000). LOCHNAGAR All of the known sites lay within the north east corrie in at least three main areas, all of which are damp, shady gullies and rock chimneys or broad ledges kept moist by percolation, and which often carry snow until later in the summer. Two colonies were counted in 1999. The first lies in a cool, dark, west-facing gully with little direct sun. The main group in this colony had 99 flowering stems with two separate smaller groups of five and six flowering stems making a total of 110 flowering stems. The Cicerbita alpina is co-dominant with Alpine Lady-fern A. distentifolium. The second colony is a broad north east facing ramp sloping at about 45o and is one of the best known localities for C. alpina as the flowers can easily be viewed from a distance. Associated species at this site include Tufted Hair-grass Deschampsia cespitosa, Red Campion Silene dioica and Broad Buckler-fern Dryopteris dilatata. A total of 239 flowering stems were recorded for this colony in 1999 (Alexander, 2000). THE LOST SITE – CANNESS GLEN This site for Cicerbita alpina was photographed in 1937 (Holden, 1952) when a large number of flowering stems are evident. The site occurred beside the Canness burn and was described as “well shaded with ‘bushy’ vegetation”. Access was described as difficult for sheep and deer. The colony occurred on a rich, stony soil with Great Wood-rush Luzula sylvatica. A small population of two to four flowering shoots persisted each year to the 1970s but since 1977 C. Geddes has recorded no plants during three monitoring visits in August 1978, July 1995 and September 1999. 3 Ecology & Life Cycle Flowering shoots rise annually from a rhizome and the flower buds usually open in late July. The plant typically grows in large clumps and the broad, sagittate leaves cast a heavy shade. When the flowers first appear they are an attractive shade of blue but they soon wither to a dull bluish violet. Flowers open for at least one day but last longer if pollination does not occur (Mejias, 1994), and wind and rainstorms often batter the plants by late summer. The seeds are surmounted by a pappus of long, stiff hairs, but ripe seed is not usually produced in Scotland. The species is pollinated by long-tongued bees and butterflies, which seek the nectar in deep-seated nectaries (Willis & Burkhill, 1902). Hoverflies have been observed on flowers (Alexander, 2000), and small tortoiseshell butterfly observed by A.G.Payne (Marren et al, 1986). Knuth (1909) states that self-pollination within the flower does not occur, although fertilisation between adjacent flowers may be possible. Experiments at Aberdeen University (Neiland & Wilcock 1996a) have shown that seed formation in Scottish populations is a rare event and examination of herbarium material, dating back to 1837 from Glen Doll, suggests that this has been the case for a long time. In contrast, seed set in central European and Scandinavian populations appears to be much more frequent, although Cicerbita alpina is more common there and cross fertilisation is more likely. Neiland and Wilcock (1996a) concluded that the reproductive biology in Scotland is probably different, although the mechanisms remain unclear. For example, it is not clear whether pollen from C. alpina in Scotland is intrinsically infertile or whether, as Sell (1986) has suggested for C. macrophylla, subsp. uralensis, there is a selfincompatibility system operating to prevent fertilisation by self-pollen (see section 1.3). 9 While Stace (1997) states that C. alpina is not rhizomatous, Neiland and Wilcock (1996a) demonstrated that the plant has a rhizome, which in the examined specimen was at least eight years old. Marren et al (1986) also reflect that the clumped habit of a Scottish colony is likely to signify clonal propagation. 4 4.1 Habitat Requirements THE LANDSCAPE PERSPECTIVE GEOLOGY & SOILS The geology of the C. alpina sites is complex. Although the soils associated with C. alpina have not been exhaustively analysed, these soils are probably analogous to those of other tall herb associations on mountain ledges which are typically periodically irrigated, skeletal brown loams and mull humus and slight podsolization. The ledges on which C. alpina grows usually carry luxuriant vegetation. The parent rocks are all acidic but they commonly contain base-rich intrusions. The parent material at Coire Kander is syenite, acid granite with prophyritic crystals of quartz and felspar. The Lochnagar sites occur within or close to gullies containing crushed rock associated with fault zones where the typical grey or pink Lochnagar granite is replaced by cream or greenish, heavily weathered rock in which the base-rich minerals epidote and calcite may be frequent (Marren et al, 1986). There is a strong likelihood that nutrients are transported to the C. alpina sites from base-rich rocks both by percolation and by rock falls. Ellenberg Indicator values (Hill et al, 1999) for C. alpina are as follows: Light – LO=6; LO=7+ Moisture – FO=6; F=6 Reaction – RO=6; R=6 Nitrogen – NO=8; N=6 Salt – SO=0; S=0 ASPECT Cicerbita alpina grows in moist mountain rock-ledges and gullies. It is found in Scotland in areas with richer soils, winter snow accumulations and summer moisture at high altitudes with severe winters. Its sites in Scotland are all on north and east facing slopes where snow cover is likely to be more reliable and longer lasting. As a consistent feature of its European habitats, winter snow cover protects plants from winter frosts and additional moisture in early summer. However, C. alpina does not seem suited to the exposures it receives in these montane locations and by the time flowering occurs in July and August the plants are often battered and have browning leaves. 4.2 COMMUNITIES & VEGETATION Cicerbita alpina typically occurs with a mix of tall herbs and Alpine Lady-fern Athyrium distentifolium, a species of late snow–beds that is a frequent associate of C. alpina in its higher sites. Other species which are usually present in abundance are Hogweed Heracleum sphondylium, Lady’s-mantle Alchemilla glabra, Tufted Hair-grass Deschampsia cespitosa, Broad Buckler-fern Dryopteris dilitata, Great Wood-rush Luzula sylvatica, Wood-sorrel Oxalis acetosella, Common Sorrel Rumex acetosa and Goldenrod Solidago virgaurea. A fuller list of close associates is given in Marren et al (1996). The ledge vegetation with which C. alpina grows is related to the NVC CG13a Dryas octopetala – Carex flacca heath community (Rodwell, 1992). On more accessible areas next to current C. alpina populations, modified versions of this community continue to exist, suggesting that C. alpina may also have 10 occurred here. C. alpina is listed as a component of the U16 Luzula sylvatica – Vaccinium myrtillus tall herb community (Rodwell, 1992; Averis et al, 2004). In Europe, C. alpina is more widespread above the tree line, but also occurs below the tree line in open upland woodland, alpine meadows and avalanche trails. GRAZING In Scotland, Cicerbita alpina only grows in places where grazing is minimal. The plant is extremely palatable to large herbivores, notably sheep and deer, and mountain hares have been observed eating it in Scotland. The tall herb communities of which C. alpina is a component can be visually dramatic in terms of their lush growth, in stark contrast to surrounding plant communities that are accessible to grazing animals. 5 Management Implications Cicerbita alpina has managed to sustain itself at four of its five historic sites in Scotland over the last few hundred years, reflecting some kind of stability. However, the habitat in which it survives is climatically fragile, and under current climate change predictions, will decline in range. Its rock ledges are also susceptible to rockfall and unstable slopes. In addition, current populations may be too isolated to allow sexual regeneration and the production of viable seed. However, conservation of C. alpina in Scotland depends largely on grazing management that would: Maintain populations by ensuring that current populations are not grazed. No other management seems necessary to protect current populations, given their longevity to date. Create suitable, adjacent habitat for future spread and increase chances for population survival in the future. Management for this species offers very real opportunities to improve the conservation status of this species: Habitat conservation should focus on restoring a lightly grazed mountain habitat that could ultimately include areas of open upland woodland with tall herbs. This may enable C. alpina to expand out of its relic sites into areas that are more akin to its stronghold sites in Scandinavia, and allow it to increase population size and genetic integrity in the face of environmental change. This would mean controlling grazing at current sites, and preferably at adjacent sites that are suitable for colonisation and spread either naturally or by introduction of plant material. At its current locations C. alpina grows vigorously in association with other tall herbs and it is likely that it would compete successfully against other herbaceous species if grazing pressure were reduced. Species management needs to ensure sufficient stock is held ex situ to allow continued experimentation and research, and to support translocation trials where appropriate and possible. 6 Threats / Factors Leading to Loss or Decline or Limiting Recovery Although of an extremely restricted distribution today, Cicerbita alpina may have been less restricted for most of the Holocene period, before the increase in grazing pressures from the 11 19th century. However, evidence of early to mid Holocene distribution of this plant is difficult to find as the pollen of C. alpina is indistinguishable from certain other members of the Asteraceae family (Bennett, 1994). It is clear that colonies have been lost since records began and there may be a number of explanations for this, including: the isolation of populations; increased sheep and deer grazing pressure; over collecting; one-off events, and the vagaries of climate. The key factors limiting the current and future distribution of C. alpina are as follows: C. alpina appears to be at the western limit of its climatic tolerance in Scotland. Grazing pressure by sheep and deer restricts the distribution of the plant to inaccessible ledges and gullies: control of grazing has been initiated at three of the sites and this may reap benefits in the longer term, as long as robust monitoring protocols are in place. Population isolation, genetic self-incompatibility and inbreeding depression may be resulting in no viable seed being produced: research undertaken by the Royal Botanic Garden, Edinburgh has given some insight into the genetics of the four populations but further research is required with larger numbers of samples. Populations are susceptible to rockfalls potentially wiping out entire colonies. Over-collection in the past, although this is no longer considered an issue. As C. alpina is not a UK Biodiversity Action Plan Species, the resources available for research and management are limited. The unknown effects of climate change: Kerr et al (1999) used the UKCIP98 climate scenarios to investigate climate impacts in Scotland for the Climate Change: Scottish Implications Scoping Study. These scenarios (Hulme and Jenkins, 1998) suggest that over the next century, Scotland will become warmer. Average temperatures are likely to rise by between 1.2 to 2.6°C, with relatively more warming in winter than summer. It is not known what impact this kind of scenario would have on C. alpina but the effect may be to minimise the continental type conditions which prevail within the area of distribution of C. alpina and on which the species appears to depend. 7 7.1 Current Conservation Measures IN SITU MEASURES COIRE FEE Coire Fee is included in a Section 7 area agreement for deer control, which includes Coire Kander and Caenlochan Glen. This has been in operation since 2002 and the effects of this will become apparent in the longer term. Another current proposal for the species at Coire Fee being discussed between Scottish Natural Heritage (Airlie) and the Angus Glens Ranger Service is to uproot some plant material under licence for introduction into a 64ha exclosure in Corrie Sharroch (K. Slater & I. Langford, pers. comm.). COIRE KANDER Coire Kander is included in the deer control area, as above, and the C. alpina may benefit in the longer term being able to establish on other more accessible ledges. 12 CAENLOCHAN GLEN There is much evidence that grazing intensity has increased markedly in Caenlochan Glen over the past 200 years. The main evidence for this comes from pollen analysis, which shows that during the last two hundred years, grass pollen has increased significantly while pollen from other species has decreased (Loizou, 1999). Reduction of grazing is a priority for the SSSI and this is being addressed through the Section 7 agreement. Additional grazing pressure from sheep coming in from the neighbouring estate of Invercauld is being addressed through the Scottish Executive. There does appear to be a very welcome concerted effort to reduce grazing pressure over a huge area covering these three Cicerbita sites. LOCHNAGAR There is currently no monitoring of C. alpina populations at Lochnagar. The deer management strategy on the Balmoral Estate aims to sustain a level of 12 deer per km2, which has almost been achieved, but it is unlikely that this level of grazing would allow C. alpina to expand to any great extent. 7.2 EX SITU MEASURES SEED BANKING Cicerbita alpina seed is not currently stored in the Millennium Seed Bank at Kew, owing to the lack of production of viable seed in the field. MATERIAL HELD IN CULTIVATION Rootstock was collected in 1996 from Lochnagar and Coire Fee and grown on at the Aberdeen University Cruikshank Gardens. The plants doubled in sized between 1996 and 1998 but had not flowered during this time. Unfortunately these 2 plants have since been lost. Plant material collected from Lochnagar and Coire Kander in 1998 is still held at the Royal Botanic Garden, Edinburgh and this is being bulked up for re-introduction where possible. It would, in addition, be useful to collect material under licence from the remaining 2 sites – Caenlochan and Coire Fee. The best method of propagation is by division of the rhizomes in spring, i.e. March or April, or taking root cuttings, which should allow approximately 12 plants to be generated from each source plant over a period of 2 years (J. Mitchell, pers. comm.). 7.3 MONITORING CICERBITA ALPINA & THE COMMON MONITORING STANDARD Methods for monitoring have changed over time and previously a combination of counting flowering stems, non-flowering stems and the number of clumps was undertaken. It is recognised (Geddes pers. comm.), that monitoring by counting flowering and/or nonflowering stems can result in damage to the plants and disruption to the substrate. In order to clarify whether the plants are still present and flowering it is possible to observe the colonies from a distance during the second half of July and August when the blue of the flowers can be seen through binoculars. However, field counts have been shown to vary by between 10-15% between individuals monitoring colonies using this method, so standardised vantage points plus the use of a margin of error should be included (Scott, 1996). The distant count method is the technique used by Scottish Natural Heritage for monitoring the Coire Kander site. C. alpina is a notified feature for the Glen Callater SSSI of which Coire Kander is a part and which will be included in Site Condition Monitoring as part of European requirements for monitoring on a 6-yearly cycle. Last monitored in 2002 this will be repeated in approximately 2008. C. alpina should also form part of the national vascular 13 plant tall herb monitoring contract in 2004 for the notified features on the Glen Callater SSSI. The population at Coire Fee is not routinely monitored but the Angus Glen Rangers may be able to undertake this work. There is currently no monitoring at Lochnagar. The Balmoral Estate Ranger Service may undertake monitoring but this is yet to be confirmed. Monitoring by Scottish Natural Heritage at Caenlochan Glen is problematic due to restrictions on access imposed by the estate and no monitoring has been undertaken since 1999 at this site. 7.4 RECOMMENDATIONS FOR FUTURE WORK GENETIC RESEARCH Further work is required to sample more plants, to sample any material which could originate from seed, and to undertake experimentation in controlled crosses in cultivation to either produce viable seed or to confirm inbreeding depression. If there are no differences in levels of seed set between crosses involving genetically clearly differentiated parents, and those involving genetically similar parents, then other explanations should be sought for lack of successful seed set in the wild. Genetic research would further clarify genotypes present and the extent of intra-population variation. RESEARCH INTO POLLINATION AND SEED SET The reasons behind the lack of seed set from fertile seed in Scotland need to be clarified. For example, seed set may be restricted by self-incompatibility and/or resource limitations. Pollen from C. alpina in Scotland may either be intrinsically infertile, or there could be a self-incompatibility system operating to prevent fertilisation of self-pollen, as proposed by Sell (1986). Alternatively, low seed set could be a result of problems at a post-pollination or postfertilisation stage. It would also be useful to set up hand pollination experiments in the field to test these hypotheses, as well as conducting experiments on material already in propagation at the Royal Botanic Garden, Edinburgh. RESEARCH INTO VEGETATIVE SPREAD It would be useful to assess whether vegetative spread does occur in Scotland as this has important management implications. POPULATION MONITORING There is a need for a standardised procedure that would be based on counting flowering heads from a distance to minimise disturbance, but from a permanent vantage point (recognised through a Global Positioning System [GPS] reading for example) and including a 10% margin of error. It might be possible to find a calibration for this technique by comparing distance counts with actual counts conducted by an appropriately trained climber. EX SITU MEASURES Efforts should be made to collect material from all sites, to preserve the genetic variation of Scottish populations, and to provide material for bulking up for further research and translocation. This material should in the first instance be vegetative. MONITORING OF GRAZING Monitoring of the effects of exclusion of large herbivore grazers, and mountain hares, should be standardised and maintained. 14 TRANSLOCATION EXPERIMENTS Suitable sites for translocation of bulked up material from the Royal Botanic Garden Edinburgh, subject to IUCN guidelines, should be identified. Established plants should be used in these translocations, as meticulous after-care is likely to be impossible. If cross-pollination is demonstrated to be important in the production of viable seed, then translocations of mixed populations into current sites should be considered. RE-INTRODUCTION INTO THE LOST SITE AT CANNESS GLEN C. alpina was last recorded at Canness Glen in 1976, when four flowering stems were recorded (C. Geddes, pers. comm.). There is potential to establish plant material from the Caenlochan ledge in the tall herb community where it was originally recorded. WOODLAND INTRODUCTION The feasibility of an introduction of C. alpina into a woodland site more typical of its continental and Scandinavian habitat was undertaken as part of the Action Plan research for Scottish Natural Heritage (Alexander, 2000). Two woodland sites were proposed in this report – Morrone National Nature Reserve and a wooded gully on the Balmoral Estate. Neither of these proposals has been further investigated and there is resistance in principle to introductions where there is no recorded evidence of past presence. Further debate on this issue would be useful to resolve how we conserve and manage species whose habitat has been completely altered by human activity over time. UPLAND WOODLAND & TREE-LINE REGENERATION C. alpina should be included at appropriate regeneration sites where the relevant lost habitat is being re-instated. This would be useful both as a conservation practice to extend and consolidate its range but also at some sites to increase public understanding. ECOLOGICAL RESEARCH Research into the ecological requirements of C. alpina on the continent and any management work being undertaken there would be useful in informing conservation activity in Scotland. 8 References Alexander, L. (2000). Action Plan Research on Alpine blue-sowthistle, C. alpina. Report to Scottish Natural Heritage. Alexander, L. (1997). Recovery investigation on Cicerbita alpina. Report to Scottish Natural Heritage. Averis A., Averis, B., Birks, J., Horsfield, D., Thompson, D. & Yeo, M. (2004). An illustrated Guide to British Upland Vegetation. JNCC, Peterborough. Beauverd, G. (1910). Contribution a L'etude des Composees. Bulletin Societe Botanique de Geneve, 2me Serie, Volume II, No 6. Bennett, K. D. (1994). Annotated catalogue of pollen and pteridiophyte spore types of the British Isles. Unpublished. Benson, E.E. (2000). Assessment of the applicability of in-vitro conservation methods to Scottish populations of Cicerbita alpina (Linn.) Wallr. Pers. comm. Clapham, A. R., Tutin, T.G. & Moore, D.M. (1987). Flora of the British Isles. 3rd edition. Cambridge University Press, Cambridge. Charlesworth, D. & Charlesworth, B. (1987). Inbreeding depression and its evolutionary consequences. Annual Review of Ecology and Systematics, 18: 237-268. Clarke, J. & Clarke, P. (1995). Action for Alpine sowthistle. Report to Scottish Natural Heritage and Plantlife. 15 Conti, F., Manzi, A. and Pedrotti, F. (1997). Liste Rosse Regionali delle Piante d’Italia. Associazione Italiana per il World Wildlife Fund & Società Botanica Italiana, Camerino, Italy. Dahl, E. (1998). The phytogeography of northern Europe: British Isles, Fennoscandia and adjacent areas. Cambridge University Press, Cambridge. Gardiner, W. (1848). Flora of Forfarshire. Edinburgh & London. Geddes, C. (1996). Monitoring of rare montane vascular plants on Ben Lawers National Nature Reserve and Caenlochan National Nature Reserve. Report to Scottish Natural Heritage. Hill, M.O., Mountford, J.O., Roy, D.B. & Bunce, R.G.H. (1999). Ellenberg's indicator values for British plants. Centre for Ecology and Hydrology Publications. Available online at http://www.ceh.ac.uk/products/publications/untitled.html Hobbs, R. (1982). The Flora of the NE Coire. Brathay Field Studies Report, 37. Hogarth, B.G. & Marren, P.R. (1996). Cicerbita alpina (L.) Wallroth. In: Wigginton, M.J. (ed.) British Red Data Books 1: Vascular Plants. JNCC, Peterborough. Holden, A.E. (1952). Plantlife in the Scottish Highlands. Oliver and Boyd, Edinburgh. Hulme, M. & Jenkins, G. (1998). Climate change scenarios for the United Kingdom. UK Climate Impacts Programme Technical Report No. 1. Climatic Research Unit, University of East Anglia, Norwich. Kerr, A., Shackley, S., Milne, R. & Allen, S (1999). Climate change: Scottish implications scoping study. Scottish Executive, Edinburgh. Kotiranta, H., Uotila, P., Sulkava, S. & Peltonen, S.-L. (1988). Red Data Book of East Fennoscandia. Ministry of Environment, Finnish Environment Institute and Botanical Museum, Finnish Museum of Natural History. Helsinki. Knuth, P. (1909). Handbook of Flower Pollination. Vol. I – III. Claredon Press, Oxford. Loizou, T. (1999). Ecological aspects and management of Caenlochan Glen. Report to Scottish Natural Heritage. Lusby, P. & Wright, J. (1996). Scottish Wild Plants. Royal Botanic Garden Edinburgh. Meijas, J. A. (1994). Self-fertility and associated flower head traits in the Iberian taxa of Lactuca and related genera (Asteraceae: Lactuceae). Plant Systematics and Evolution, 191: 147-160. McVean, D. N. & Ratcliffe, D. A. (1962). Plant Communities of the Scottish Highlands. Monograph no. 1 of the Nature Conservancy. HMSO, London. Marren, P.R., Payne, A.G. & Randall, R.E. (1986). The past and present status of Cicerbita alpina (L). Wallr. in Britain. Watsonia, 16: 131–142. Neiland, M. R. M. & Wilcock, C. C. (1996). Cicerbita alpina. SNH report. Unpublished. Neiland, M. R. M. & Wilcock, C. C. (1996a). The reproductive biology and conservation of 3 rare Scottish plant species: Linnaea borealis, Cicerbita alpina and Moneses uniflora. Plantlife report. Unpublished. Nordhagen, R. (1943). Sikilsdalen og Norges Fjellbeiter. Bergens Museum Skrifter 22. Norstedt. (1920). Prima loca plantarum suecicarum. Första litteraturuppgift om de i Sverige funna vilda eller förvildade kärlväxterna. Bilage till Botaniska Notiser 1920: 1-95. in http://linnaeus.nrm.se/welcome.html.en Oliver, C., Hollingsworth, P., Griffiths, R., & Sydes, C. (2000). Genetic Diversity in Scottish Populations of Cicerbita alpina (L.) Wallr. (Compositae). Unpublished. Ratcliffe, D. A. (1958). Cicerbita alpina (L.) Wallr. in Glen Callater. Transactions of the Proceedings of the Botanical Society of Edinburgh, 37: 287-289. Richards, A.J. (1997). Plant Breeding Systems. 2nd edition. Chapman and Hall, London. Rodwell, J. S. (1992). British Plant Communities, Vol. 3. Grasslands and Montane Communities. Cambridge University Press, Cambridge. Schierup, M.H. & Christiansen, F.B. (1996). Inbreeding depression and outbreeding depression in plants. Heredity, 77: 461-468. Scott, M. (1996). Action proposals for Alpine (Blue) Sowthistle Cicerbita alpina in Scotland. Consultative report to Scottish Natural Heritage. 16 Sell, P.D. (1986). The genus Cicerbita Wallr. in the British Isles. Watsonia, 16: 121-129. Smith, J.E. (1811). An account of several plants, recently discovered in Scotland by Mr George Don A.L.S. Trans. Linn. Soc. Lond., 10: 333-346. Stace, C. (1997). New Flora of the British Isles. 2nd edition. Cambridge University Press, Cambridge. Williams, J.G.K., Kubelik, A.R., Livak, K.J., Rafalski, J.A., & Tingey, S.V. (1990). DNA Polymorphisms amplified by arbitrary primers are useful as genetic markers. Nucleic Acids Research, 18: 6531-6535. Willis, J. C. & Burkill, I. H. (1902). The alpine flora and rarer plants of the Glenshee District. Transactions of the Proceedings of the Botanical Society of Edinburgh, 23: 83-91. 9 Acknowledgements Thanks to Claire Geddes, Michael Scott, Chris Sydes, Lynne Farrell and Peter Marren for their valuable comments on this dossier, to Peter Hollingsworth for generously providing additional information, and to Plantlife Scotland volunteer Jill Williams for her editorial work. 10 Contacts Back from the Brink contact: Dr Deborah Long, Conservation officer, Plantlife Scotland SNH Species Recovery Programme: Dr Chris Sydes, Species Group Manager, Scottish Natural Heritage. Plantlife Scotland, Balallan house, Allan Park, Stirling, FK8 2QG Tel: +44 (0) 1786 478509 Fax: +44 (0) 1786 479382 [email protected] Scottish Natural Heritage, 2 Anderson Place, Edinburgh, EH6 5NP Tel: +44 (0) 131 447 4784 Fax: +44 (0) 131 446 2405 [email protected] 11 Links Plantlife International wishes to acknowledge the financial support of Scottish Natural Heritage, the Countryside Council for Wales and English Nature, for the Back from the Brink (species recovery) programme. ISBN: 1 904749-15-1 Original draft by Leonie Alexander Edited by Plantlife Scotland First draft dated March 2004 Last revised 20 March 2006 17